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-VoLuMe 19° Numer 1 ‘SEPTEMBER, 2000 oo 


Barney L. Lipscomb, Editor 
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OSIDA, CONTRIBUTIONS TO BOTANY 
VOLUME 19, NUMBER 1, PAGES 1-234 
Copyright 2000 

Botanical Research Institute of Texas 


Printed in the United States of America 


ISSN 0036-1488 


TABLE OF CONTENTS 


BOTANICAL ACCOUNT OF THE JOSEPHSTAAL FOREST MANAGEMENT AGREEMENT AREA, PAPUA New GUINEA 


A FLORISTIC AND ET 
W. TAKEUCHI—1 


ERAGROSTIS ANCASHENSIS (POACEAE: CHLORIDOIDEAE), A NEW SPECIES FROM ANCASH, PERU 
PAUL M. PETERSON, NANCY REFULIO RODRIGUEZ, AND Oscar TOVAR—65 


WOOD AND BARK ANATOMY OF ACHATOCARPACEAE 
SHERWIN CARLQUIST—71 


BATOPILASIA (ASTERACE 
Guy 


: ASTEREAE), A NEW GENUS FROM CHIHUAHUA, MEXICO 
L. NESOM AND ee D. Noyes—79 


A NEW COMBINATION IN ARCHIBACCHARIS (ASTERACEAE: ASTEREAE) 
Guy L. Nesom—85 


REVISION OF LOBELIA SECT. TUPA (CAMPANULACEAE: LOBELIOIDEAE) 
THOMAS G. LAMMERS—87 


A NEW SPECIES OF TREPADONIA (ASTERACEAE: VERNONIEAE) FROM PERU 
HAROLD ROBINSON AND HAMILTON BELTRAN—111 


RHYNCHOSPORA ZACUALTIPANENSIS AND ELEOCHARIS MOOREI, [WO NEW CYPERACEAE FROM MEXICO 
Mark T. STRONG AND M.S. GONZALEZ-ELIZONDO— 115 


DISTINCTION BETWEEN VITIS BLANCO! AND V. CINEREA VAR. TOMENTOSA (VITACEAE) 
Barry L. COMEAUX AND JIANG LU—123 


BOLBITIS THOMMANKUTHIANA (LOMARIOPSIDACEAE), A NEW SPECIES OF FROM INDIA 
ANTHOSH NAMPY—133 

A NEW COMBINATION FOR SARCOSTEMMA CYNANCHOIDES VAR. HARTWEGII (ASCLEPIADACEAE) 

ALEXANDER KRINGS—137 
RHYNCHOSPORA LEPTOCARPA (CYPERACEAE), AN OVERLOOKED SPECIES OF THE SOUTHEASTERN UNITED STATES 

Bruce A. SoRRIE—139 


OCCURRENCE, DISTRIBUTION, AND ECOLOGY OF ALOCASIA, CALADIUM, COLOCASIA, AND XANTHOSOMA (ARACEAE) 


IN THE SOUTHEASTERN UNITED STATES 
Brett E. Serviss, SIDNEY T. MCDANIEL, AND CHARLES T. BRYSON—149 


GEOGRAPHIC DISTRIBUTION PATTERNS OF THE GENUS ARIOCARPUS (CACTACEAE) IN TAMAULIPAS, MEXICO 
Jost GUADALUPE MARTINEZ-AVALOS AND HUMBERTO SUZAN-AZPIRI—175 


PLATEILEMA (ASTERACEAE: HELENIEAE) A NEW GENERIC REPORT FOR THE UNITED STATES 
BILLig L. TURNER—185 


WHICH NON-NATIVE PLANTS ARE INCLUDED IN FLORISTIC ACCOUNTS? 
Guy L. NEsom—189 


CINNA AND LIMNODEA (POACEAE): NOT CONGENERIC 
David M. BRANDENBURG AND JOHN W. THIERET—195 


THEMEDA QUADRIVALVIS (POACEAE: ANDROPOGONEAE) IN KANSAS: AN EXOTIC PLANT INTRODUCED FROM BIRDSEED 
E. GENE TOWNE AND IRALEE BARNARD—201 


NODULATING LEGUMES FROM THE TAHOE BASIN, CALIFORNIA 
MOHAMMAD ATHAR AND JAMES HARDING——205 


NOTEWORTHY PLANTS FROM NORTH FLORIDA. VII 
LORAN C. ANDERSON—211 


WHO NAMED EUTHAMIA (COMPOSITAE: ASTEREAE) AND WHEN? 
JOHN L. STROTHER—-217 


JOSEPH ANDORFER EWAN, OcToOBER 24, 1909—DecemeBer 5, 1999. A MEMOIR 
ANNE S. BRADBURN—219 


BOOK REVIEWS AND NOTICES 64, 114, 132, 136, 148, 184, 188, 194, 204, 223 


ANNOUNCEMENT 234 


¥ 


A FLORISTIC AND ETHNOBOTANICAL ACCOUNT OF THE 
JOSEPHSTAAL FOREST MANAGEMENT AGREEMENT 
AREA, PAPUA NEW GUINEA 


W. Takeuchi 


Botanica Research Institute of Texas ay ue 
| and Panijia New Grinen Enrect Recenn h Inctitiste 
i eee Guinea 


PO. Box 314, Lae, Morobe Province 41 17, PAPUA NEW GUI NEA 


ABSTRACT 


Botanical survey results are presented from the Josephstaal Forest Management Agreement Area, 
a venue currently under planning evaluation for several development contingencies. Four new 
cS are e formally described: Ag! aia saxonii (Meliaceae), Barringtonia josephstaalensis 
Cal (Rubiaceae), and Psychotria mayana (Rubiaceae). A substan- 
tial number of distributional records and discoveries of rare taxa are reported. Compilations of 
Maian plant names and uses are also in 
The findings suggest that es habitats are refugia for restricted endemics which have 
been eliminated from other parts of their historical range. Judicious planning is necessary when 
evaluating the land-use options for this area. Populations of several susceptible taxa may consti- 
tute the only existing colonies. 


Key Woros: Aglaia, Barringtonia, botanical d ies, Calycosia, Josephstaal, Papuasia, 


Psychotria 


ABSTRACT 
RE. COLMA ARUMROSRA PCHSY av IAY IRAE Bis EME Ic 
SIF SHMBAOBRERATS, MAICSI\ CHR SNE 4 DOMMBICDINT 
(4. EORMSDBARAPICADHNTS. ENSORMES Aglaia saxonii(vvyv 
#4), Barringtonia josephstaalensis (tH U /\F#), Calycosia mamosei (7 AAR 
#4). Psychotria mayana(PHAREH)CHS, BBERICLMMBPHARAKEZEO 
RMS H RID SOBRMERENTING., 
YatkIay—Ibihitit, DOTODAMMRM5S HAKRDCLEDKHOPORA 
HOVILTY TS CHSACEMRMABRD SOHMMANS, COMMOLMARTHO 
fERRICISBRRAABHSVEMHS. Va IAD —I Ibe HRODORES 
SIF PT UMHO. HEFT SREOIOXM—THSAREHMHSD5 THS. 


INTRODUCTION 


The Josephstaal Forest Management Agreement Area (JFMAA) is an intended venue for 
logging operations based on the reduced-impact formats known as ‘ecoforestry.’ Prior 


SIDA 19(1): 1-63. 2000 


2 SIDA 19(1) 


to the project's implementation, botanical surveys were considered necessary to estab- 
lish biological baselines for the concessional areas. An ecological reconnaissance of the 
JFMAA (Fig. 1) was thus conducted by the Nature Conservancy (TNC) between Septem- 
ber 9 and 17,1998, followed by a general floristic evaluation during the period from July 
26 to August 25, 1999. The following paper is a synopsis of the initial findings from these 
investigations. 


SITE SUMMARY 


The survey tract is located for the most part, within territory covered by the Annanberg 
topographic sheet, but also overlaps the adjacent Adelbert, Manam,and Nubia map units 
(cf. Australian Survey Corps 1973, 1974a, 1974b, 1974c: 7888, 7889, 7988, 7989). This gen- 
eral area is part of the Mugumat-Yakiba Census Division of Bogia Subdistrict; and in- 
cludes the principal villages of Dumadum, Moresada, Mugumat, Roumirap, Wadaginum, 
and Wagadab (Dept. of District Administration 1968: 88-89). 

The 1999 survey was based at three camps established sequentially at map coordi- 
nates (GPS) 9504560 N x 281407 E;9497596 N x 280100 E;and 9498679 N x 284829 E; at 
elevations from ca.50 to 160 m. There was mature growth foothill forest at Camp 1,and 
alluvial terrace communities at Camps 2 and 3. The latter bases provided convenient 
access to both riverine and foothill vegetation. 

All investigated sites are within northern Papua New Guinea's (PNG) lowland forest 
life zone, where mean annual rainfall is 2,000 to 3,500 mm. The wettest months genera lly 
occur during January to April, when prevailing winds are northwesterly, and the driest in 
May to August when southeasterly trades become effective (McAlpine et al. 1983: 65). 
Even during the relative dry season, average monthly rainfalls are still generally around 
200 mm, so the vegetation is only infrequently subjected to soil moisture deficiencies 
under normal conditions (ibid: 140). Most climatic classifications would categorize the 
project sites as tropical everwet, perhumid, or some other equivalent descriptor empha- 
sizing the overall absence of water deficits. 

The survey sites are typical for a PNG wilderness area, in that information-gathering 
services are sporadic or altogether lacking. Meteorological summaries are necessarily 
developed by extrapolation from stations which may not be representative. Although 
the nearest station with published tables (Madang A/S) shows high annual rainfalls with 
moderate seasonality, severe droughts have been known to occur in this general region. 
Episodes of widespread fires and forest destruction have been documented (Johns 1986: 
349-351, 359). 

From a geological perspective, the Josephstaal Physiographic Province represents 
the crustal remnants of an island arc which collided with the Australian plate about 10 
m.y. BP (Pigram & Davies 1987). Parent substrates are generally derived from basalt 
volcanics of this now-disappeared arc Jaques & Robinson 1975: 12). Severe earthquakes 
occur frequently within the area covered by the TNC project, so plant speciation is prob- 
ably being encouraged by landslide-induced isolation of populations, particularly at the 
higher elevations (cf. Balgooy et al. 1996: 201-02). 


LORENGAU. 
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DO KAVIENG 
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NEW BRITAIN 


TWWLSHdISOf WOH SLINSIY TWDINVLO® ‘IHNINVL 


4 SIDA 19(1) 


Soils of the Bogia Subdistrict (including Josephstaal) are dyst ts and eutropepts, 
a group characteristic of lowland habitats and comprising the ioe common soil type 
in Papua New Guinea (Bleeker 1983: 98-9). There are no ultramafic or other specialized 
edaphic environments within the surveyed tract. 


HISTORY OF BOTANICAL ACTIVITY AT JOSEPHSTAAL 


Botanical work on the Josephstaal flora has been sporadic and opportunistic in nature, 
with involvement by a limited number of specialist collectors. Prior to the TNC surveys, 
the largest collection was the set made by KJ.White during the period September 1 to 15, 
1958.White obtained a total of 103 numbers under the NGF (New Guinea Force) series, 
from which three collections were later designated as type specimens (Endiandra magni- 
limba Kosterm., Horsfieldia basifissa de Wilde, and Syzygium madangense Hartley & Perry). 
Contemporary efforts at identification of historical localities have been generally 
complicated by the oftentimes limited data provided by earlier botanists in Papuasia. 
The K.J.White numbers are thus somewhat uncharacteristic for the period, since for all 
sheets the point of reference is clearly specified as Josephstaal,’ at'L.4 45 S and Lat. 145 
00 E.'The elevation is consistently given as ‘250 feet’and repeatedly indicates riverine or 
alluvial habitats. Pondoma, Naikum, and Tumbundi villages are mentioned on some la- 
bels (e.g., Thespesia fissicalyx in NGF 10297; Maniltoa rosea in NGF 10226; and Cryptocarya 
weinlandii in NGF 10306, respectively). From the information provided on White's gather- 
ings, there are consistent indications he was working in the lower basin connected to 
the Guam River. For at least part of the time, White was probably accompanied by R.G. 
Robbins, as suggested by an independent number (Endiandra squarrosa in Robbins 1625), 
which is cross-referenced against White’s NGF 10252,and which refers to the latter as a 
duplicate. Robbins's locality is specified as Josephstaal, lower Ramu-Atitau area, Madang 
District.’ Robbins 1667 (Barringtonia apiculata) adds further:‘near Josephstaal 400 ft.’ 
White's specimens have been intensively studied over the years, and the determi- 
nations assigned to them by specialists indicate the sort of taxa that would be expected 
from the JEMAA on general distributional and ecological grounds. The K.J.White collec- 
tions have been incorporated into the survey documentation (Appendix 1) because the 
species involved are present within and immediately around TNC’s project area. 
During the herbarium work phase, specimens collected by R. Pullen were also ex- 
amined from the vicinity of Josephstaal. A typical label from Pullen 1188 (Alocasia 
brancifolia), gives the collection site as 1/4 mi south of Josephstaal airstrip. Although the 
Pullen numbers are from comparable lowland habitats, they were collected slightly out- 
side the project area and are mentioned only incidentally in the following summaries. In 
any event, comparatively few Pullen specimens are present at the PNG National Her- 
barium (LAE); the NGF sheets comprise a decidedly larger set. 
Other botanists with collections from the Josephstaal tract include J.Womersley, 
B.S. Parris, and J.P. Croxall. The Parris and Croxall numbers are specialist pteridological 
collections made along the unimproved road to Josephstaal in 1980. Most of their fern 
records have been replicated by the surveys’ results. Womersley’s numbers from the 


a 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 5 


Wanuma area (NGF 48651-48678 from 600 to 950 m elevation in 1974) are substantially 
removed from the project site but are notable for being among the few sets taken from 
the Adelbert summit range. 

By far the most comprehensive specimen series for Josephstaal are the collections 
from the TNC-sponsored botanical surveys of 1998 and 1999. A total of 973 numbers 
were added to the national herbarium from these efforts; 62 from the 1998 ecological 
reconnaissance and 911 from the recently-concluded expedition. The combined ay 
increases the plant documentation for Josephstaal by nearly an order of magnitud 
what was previously available. Due to the surveys’ exclusive focus on he Sait 
interval below 400 m,the herbarium coverage for the JFMAA is now among the best for 
any lowland wilderness in Papuasia. 


METHODS 


The 1999 survey consisted of general exploration and collecting around three expedi- 
tion camps, primarily using established footpaths or following the secondary channels 
comprising the Guam drainage. The botanical collections were conducted by an inte- 
grated team consisting of M. Gorrez, W. Takeuchi, A. Towati, and J.Wiakabu. During the 
selection of specimens, deliberate attention was directed to groups usually spurned by 
botanists because of their inherent repellent qualities or other difficulties associated 
with their processing. Palms, aroids, stinging nettles, grasses, allen weeds, etc. were se- 
cured when suitable specimens were encountered, in contrast to the general reluctance 
for collecting such plants. Multiple gatherings of certain taxa were also made, when their 
significance was already apparent in the field, in order to allow evaluation of population 
variation. Survey protocols were consistent with the principal objective of developing a 
representative floristic profile of the project area within the allocated one-month period. 

Ethnobotanical polling was conducted separately by survey biologists J. Wiakabu 
and M.Gorrez, through group interviews with village elders. Names and uses of specific 
plants were recorded, with special attention paid to culturally sensitive taxa. All vouchers 
were field-pressed in 70% surgical alcohol and subsequently transported to the PNG 
Forest Research Institute (PNGFRI) for processing and determination. Materials for 
exsiccatae were often accompanied by bottled, carpological, and xylarium accessory 
collections when these were necessary for identification. 

The Lae National Herbarium (LAE) is the repository for first sets from the vouchers. 
Distribution of duplicate sets will follow LAE’s exchange sequence, on which Kew (kK), 
Rijksherbarium (L), and Harvard (A), are the principal receiving institutions. Residual sets 
will be allocated in conformity to preexisting agreements or in compliance with future 
TNC requirements. Whenever possible, soecimens were named using the current taxo- 
nomic revisions, or from a combination of authoritatively annotated sheets and original 
descriptions. Some collections could only be assigned with doubt to a species group or 
section. In a number of cases, submissions were made to international specialists. Taxa 
encountered only in sterile condition or otherwise not collectable, were enumerated as 
a sight record when the plant was known with certainty to the writer. 


6 SIDA 19(1) 


GENERAL DESCRIPTION OF THE VEGETATION 


Two principal forest-structural divisions (alluvial forest on riverine terraces, and foothill 
forest on well-drained slopes), were discerned during the initial 1998 reconnaissance 
The opportunity for detailed examination afforded by the 1999 survey, subsequently 
showed that many community types are included under these two formations. 

The margins of large streambeds in the Josephstaal area are marked by a distinctive 
riverine facies dominated by lianes, heliophytes, and rheophytic taxa. This edge commu- 
nity is generally absent from smaller streams with closed canopies. In swampy situations, 
the interior alluvial stands are typically species depauperate, Metroxylon-dominant, and 
with poor vertical development. On better-drained sifuivis the forest becomes more flo- 
ristically and structurally diverse, eventually forming a varied community with interlock- 
ing canopy layers and clear understories. There are intermediate communities appar- 
ently linked to diminishing rhizosphere stagnation. However in places with a linear series 
of staggered terraces, such intermediates are interpretable as a successional sequence 
resulting from progressive changes in streamcourse (Saxon, pers.comm.). Superimposed 
over the matrix of variation are smaller units in various stages of regrowth, which have 
been caused by large treefalls, attritional senescence, catastrophic storms, etc. While the 
alluvial forest is for the most part assigned to structural code 'Fri’in Hammermaster and 
Saunders (1995a), there is clearly a continuum of communities within this category. 

From examination of understory and subcanopy taxa, the forest on hillsides and 
ridgelines was initially regarded as homogeneous. Herbs and subarborescent plants 
appear to range through the foothill habitat without obvious distributional separations. 
However Weterings (pers.comm.) noted pronounced contrasts in canopy compositions 
between ridgelines and lower slopes during the independently-conducted timber as- 
sessment. His observation is supported by the known autecological patterns of arbores- 
cent genera, as for example the preference of dipterocarps for ridgecrest environments 
(Johns 1977; Paijmans 1976). Like the alluvial zone, there is very probably a fragmenta- 
tion of the foothill formation into subtypes, though the differentiation may be primarily 
reflected in overstory structure, while with the riverine communities, it is discernable in 
the near-ground compositions. Characterization of the hill forest communities will re- 
quire surveys of greater intensity than the one just concluded, employing a combina- 
tion of transects and random collecting. This suggestion is supported by recent findings 
from similar rapid-assessment surveys in other parts of Papuasia (e.g., Mack 1998). 

The Josephstaal foothill communities fall primarily under forest structural code’'Hm’' 
and are known to intergrade with alluvial formations (Hammermaster & Saunders 1995a: 
11).The ‘Hm’ category is the major merchantable forest unit in the existing JFMAA 
(Hammermaster & Saunders 1995b:SB 55-1 Bogia overlay). On the earlier classifications 
of Paijmans (1975) and Saunders (1993), the project sites are placed respectively under 
structural codes ‘FHm’ and ‘Hm.’ Due to similarities in terminology employed by each 
author, the different typing sytems yield comparable floristic descriptions. The Josephstaal 
tract is essentially a typical medium-crowned forest from low elevation environments. 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 7 


However the wide variation within this forest type obscures the commercial valuations 
assignable to specific subunits. 


ETHNOBOTANICAL COMPILATIONS 


The plant names provided by respondents are derived from the traditional Maia (Maya) 
language spoken by villagers within the project area. Clan elders Francis Muoimuado 
and Josef Sigagopa were principal sources for the information summarized in Appendi- 
ces 2 and 3. Ethnobotanical questioning usually occurred in the presence of a village 
audience, with the clan elders serving as a central authority or facilitator. A consensus 
was thus established. The group interviews provided a means for identifying the assets 
requiring protection from ecoforestry operations. Because the survey objectives were 
primarily taxonomic, ethnobotanical inquiry was subordinated to the main itinerary, and 
rapid appraisal procedures were adopted in place of rigorously analytic methods. All 
aspects of the ethnobotanical inquiry will require critical evaluation against established 
benchmark studies, when the present investigations are extended into other parts of 
the Adelbert Range. 

Despite these limitations, certain patterns are evident from the compilations. 
Unlike the nomenclature of formal science, with its simple and rigorously applied bino- 
mial protocol, the botanical classification employed by Josephstaal villagers is multifari- 
ous and idiosyncratic. Most Maian plant names appear to be descriptively based, and if 
translated will probably be seen as alluding to particular features of a plant, in the man- 
ner of pre-Linnean botany. Although it is obviously not a two-element nomenclature, 
Maian plant names can occasionally exhibit functional resemblances to a binomial sys- 
tem.In such instances ‘generic’ units are usually indicated with a common designator at 
the front of a complete name, the ‘species’ then being specified with a qualifying phrase 
or word following the generic mark. Examples include: ‘warubu-nganam,’ ‘warubu- 
sopasop,’ and ‘warubu-taleba;’ corresponding respectively to Glochidion sp. aff. 
chondrocarpum, Dysoxylum brassii, and Dysoxylum pettigrewianum. Various taxa in 
Strobilanthes (Hemigraphis) are similarly designated as variants of ‘sagag;’ ie. ‘sagag- 
gosmun,'‘sagag-u-goga-umun,’ and ‘sagag-ugosum. The conventions are comparable 
to findings reported by Petir et al. (1998), from a study conducted in another part of the 
Adelbert Range. 

In most cases, similarities in gross appearance are apparently the major criteria for 
application of names. This is demonstrated by the woody shrubs Lepisanthes senegalensis 
(Sapindaceae), Ixora sp. sect. Hypsophyllum (Rubiaceae), and Phaleria coccinea 
(Thymelaeaceae), which are all identified as‘kibi-kibale’ despite obvious contrasts in their 
fertile aspect. The gross equivalence in habit and leaf form is seemingly sufficient for 
combining these taxa under one concept. The Maian ‘maberu’ is similarly applied to 
Cleistanthus sp. aff. papuanus, Erythrospermum candidum, and Rhyticaryum longifolium, 
showing again that names are assigned on the basis of superficial aspect rather than by 
awareness of specific structural distinctions. The rationale may be less clear however, in 


8 SIDA 19(1) 


cases where the members of a nomenclatural group have little in common even in 
regards to general form. Other than the fact that they all pea pe constructed 
ferns, Microsorum memobranifolium, Asplenium cf. affine, Lind ia, Bolbiti 

and Pleocnemia macrodonta, are obviously different plants, yet are ree at least in 
part, to the one name ‘lasa-lasa.’ The differences among the referents of ‘dagol-dagol’ are 
even more striking; including such diverse taxa as Asplenium cf. amboinense, Lindsaea 
obtusa, and Liparis condylobulbon. From the fact that these plants are so obviously differ- 
ent, the Maian classification surely is not founded on judgments of taxonomic equiva- 
lence in the Western sense, but must be proceeding from some other logic; possibly 
involving a principle of utility. 

In floristically rich environments such as are generally present in Papuasia, only a 
fraction of the botanical diversity will enter the cultural consciousness. Many plants re- 
corded by the recent surveys do not have a local (tokples) name; or at least the respon- 
dents were unable to provide one. Of those taxa which find tokples assignment, a very 
small percentage are actually of ethnographic significance (Appendix 3).When a plant is 
of high usage-value, it is accorded a unique designation and the phonetic root for that 
name is often not transferred to other plants.In general the converse relationship is also 
true; Maian names with heterogeneous group membership are generally composed of 
‘useless’ elements (e.g,,lasa-lasa’) for which there are no clearly defined 
plants of diverse appearance are placed together under common designation, the spe- 
cies involved are not of particular value,so from the traditional-cultural perspective there 
may be no imperative for distinguishing them anyway. Maian botany is arguably 
grounded on practical principles. As an information retrieval system, it expends efforts 
toward the identification of resources with specific cultural application, and tends to 
consign everything else to loosely defined sets. In cultures without a written tradition, 
such economy is probably necessary to restrict the amount of ethnological data to lim- 
its amenable to oral transmission and retention. While the Maian plant classification is 
typically artificial and inappropriate as an adjunct to formal floristic research, it appears 
to represent a system closely adapted to local interests and requirements. 

The fact that very different species are often placed under the same Maian name, 
will complicate TNC’s intentions to train forest stewards from traditional landowner 


groups. Especially in speciose families such as Sapindaceae and Annonaceae, where iden- 


applications.Where 


tifications are largely dependent on an understanding of reproductive structures, Maian 
botany will be unable to contend with many discriminations even at generic level. This 
discourages use of the Maian system as a basis for cross-cultural instruction, and neces- 
sitates introduction of Western concepts into the training process. 

The frequent lack of tokples specificity also argues against reliance on villagers for 
plant identifications in floristic enumerations.Such dependence would result in informa- 
tion loss and underestimates of diversity. There is no effective substitute for employment 
of high-resolution taxonomic concepts during botanical inventories; the commentary 
by Kartawinata (1990: 125) regarding the unreliability of local names, is especially relevant. 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 9 


DESCRIPTIONS OF NEW SPECIES 


BARRINGTONIACEAE 

Barringtonia josephstaalensis Jakeuchi, sp. nov. (Fig. 2). Tyee: PAPUA NEW GUINEA. Maan 
Province: Josephstaal FMA area, Guam River, alluvial forest on riverine terraces, between GPS 
coordinates 9497596 N, 280100 E, and 9496.322 N, 274.601 E,80 m,9 Aug 1999 (fl), W Takeuchi, 
J. Wiakabu, M. Gorrez, & A. Towati 13,796 (Hotoryre: LAE; isoryees: A, BRIT, K). 


Inter species Barringtoniae singularis ob folia lineari-loriformia, 47-84 cm longa, 16-38 mm lata, 
apice acutata, basi sensim angustata. 

Understory shrub; monoaxial or branched, to 2 m tall, entirely glabrous. Branchlets terete, 
apically and discontinuously fistulose, otherwise pithy, periderm crustaceous, exfoliating 
in flakes. Leaves spirally congested in terminal rosettes, blades herbaceous or fleshy, 
adaxially medium green, abaxially light green, obliquely ascending in the lower half, droop- 
ing in the upper half, linear or ligulate, 47-84 cm x 16-38 mm (200-310 x longer than 
broad), attenuate at both ends, margins serrulate, the serrulations with an antrorse pro- 
cess inserted on the leading edge and following the forward margin; venation pinnate, 
secondaries 50-65 pairs, 8-12 mm apart, arcuate, tbrochidodromous but anastomos- 
ing freely beyond the commissural loops, tertiary nervation conspicuously and bifacially 
areolate, prominulous, midribs prominent on both surfaces; petioles slender, adaxially 
plane, rounded beneath, proximally swollen, to ca. 9 cm length but obscurely distin- 
guished from the lamina and occasionally with the decurrent leaf base nearly reaching 
the stem; stipules linear-acuminate, typically 9-18 x 1.5-2.0 mm, falling early, costate, 
the rib excurrently prolonged to a filiform cauda./nflorescence cauligerous or ramigerous, 
cernuous, racemose, at times corymbiform, pauciflorous from a rachis 2.0-3.5 cm long; 
peduncular bracts stipuliform, to 14 x 1 mm, involute; bracteoles minute, linear, not or 
barely exceeding 1 mm length; pedicels 7-11 mm, articulated at the base. Flowers (mea- 
surements from rehydrated specimen) globose in bud, initially brownish-purple, later 
green and red-suffused; calyx tube turbinate, not angulate nor alate, the limb membrana- 
ceous, completely connate, at first enclosing the other parts, mucronulate or infrequently 
with an apical orifice, rupturing at anthesis into 2(-3) subequal lobes, these approxi- 
mately plinerved, parting nearly to the base, suborbicular, ca. 14 x 13 mm; petals 4, nar- 
rowly obovate to oblanceolate, to 24 mm long for buds nearing anthesis, concave, venose; 


androecium multiseriate, staminal column 13 x 4—5 mm,stamens inserted on the outer 
side, the lowermost stamens arising 8-9 mm from the base, anantherous, ca. 35 mm 
long, antheriferous stamens 15-20 mm long, crowded above the lower staminodial ones, 
the tube rim crowned by a fringing whorl of staminodes ca. 2-3 mm long; ovary (3-)4 
celled, ovules several per locule, apically inserted, pendulous, irregularly obovoid; style 
capillary, 28-32 mm long, exceeding the petals but remaining enfolded until loss of the 
corolla, thereafter persistent, basally dilated into a conical stylopodium 3 x 3 mm;stigma 


weakly capitate. Fruits unknown. 
Distribution and ecology.—Known thus far only from the Josephstaal tract. 
Barringtonia josephstaalensis is a facultative helophyte from seasonally flooded riverine 


SIDA 19(1) 


H 


{ 
' 
\y 
a 


L ry + 


C. Flower 


Fic. 2. Barri jaj h lensis Tak hi, sp.nov A Vegetative habit.B.R 


4.4 . J? of 1 orl tit er a | rf. 


of the staminal cylinder. Fe 


mm 


rtile stamens, outer 


y N.H.S Howcroft 


staminodes, and style removed for clarity. Scale bars: A-B, 5 cm;(, 5 


cay 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 11 


flats and the lower slopes of foothill forest. It is apparently not common, at least within 
the area covered by recent assessment. 
Etymology.—The new binomial commemorates the Josephstaal type locality. 


Pararypes: PAPUA NEW GUINEA. Mapanc Province: Josephstaal FMA area, Guam River near expedition 
mp 2, lower slopes of natural-growth foothill forest, near GPS coordinates 9497596 N, 280100 E, 
50-100 m, 13 Aug 1999 (fl), W. Takeuchi, J. Wiakabu, M. Gorrez, & A. Towati 13,973 (LAE, NY). 


The connate calyces clearly indicate membership in section Barringtonia, within which 
the new species is easily distinguished by its linear leaves. However it is not as certain 
whether B josephstaalensis is naturally branched; the type collection was made along a 
forest track so the branches may be the result of bayonnet reiteration. 

The monadelphous androecium is marked by the unusual presence of both an 
outer and inner staminodial whorl. Although the inner series is highly reduced, the out- 
ermost structures are conspicuously longer than the fertile stamens. In Payens’s (1967: 
164) revision, the st des of all species are always clearly vestigial and only disposed 
in adaxial whorls. The existence of outer-marginal staminodes and their prolongation in 
B.josephstaalensis, are thus highly unusual elements. Together with the linear leaves, the 
character combination for this species is unprecedented. Although the plant's appear- 
ance is very deviant, the racemiform inflorescence and apical insertion of ovules are 
otherwise consistent with Barringtonia. 

The new species is known by the Maian name ‘kun-joob,’and its bark is reportedly 
used to poison fish in the manner of Derris (Fabaceae). Ethnobotanical application of this 
sort had been reported previously for the sympatric Barringtonia calyptrocalyx var. mollis 
(Payens 1967: 212). The latter taxon is identified by the separate Maian name ‘gaira- 
malapta.’ 

Barringtonia josephstaalensis will key to fork 23 in Payens (1967: 180) before reach- 
ing an impasse. It can be accommodated by interposing the following couplet between 
the existing couplets 22 and 23: 


Blades 200-310 x longer than broad; lamina ligulate Barringtonia josephstaalensis 

chi 
Blades 1.75-31 x longer than broad; lamina various but not ligulate to couplet 23 
MELIACEAE 


Aglaia ch ealameall sp.nov. (Fig. 3). Tyee: PAPUA NEW GUINEA. Mapanc Province: Josephstaal 
ea, along track to UlaeaDe W of Bia Camp 1 (’Kumamdeber’), mature growth 
eon between GPS coordina 4560 N, 281407 E, and 950285 N, 28030 E,ca. 160 
m,1 Aug 1999 (fr, carpological), W Take re |. Wiakabu, M. Gorrez, & A. Towati 13,712 (HOLOTYPE: 
LAE; isotypes: A, BRIT, K, L). 


Species haec Aglaia subsessili P ll affinis sed fruc porcatu, loculis 2,denique 


semine in quoque loculo solitario. 


Subcanopy tree to 15 m height. Branchlets elactiferous, moderately robust,6-8 mm diam., 
the periderm weakly sulcate or irregularly cracking in brittle flakes, at first with an orange- 
brown indument of dimorphically stellate hairs, the larger hairs appressed or obliquely 


12 SIDA 19(1) 


Fic. 3. Aglaia saxonii Takeuchi, sp. nov. A. F g B. Fruit, side view. C. Fruit, proximal polar view. D. Fruit, 
cross-section showing t ds and tral | Scale bars: A, 5 cm; B—C, 5 cm; D, 4. cm. Drawn from the type by 
N.H.S. Howcroft. 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 13 


patent, sometimes crisped, hyaline, arms 10 or more, acicular-setiform, ca. 1 mm long; 
minor hairs compact, rays coarse, congested, 0.1-0.2 mm long; stem surfaces early gla- 
brescent and then entirely glabrous below the leaf spray. Leaves imparipinnate, 3-5 ju- 
gate, spiral, terminally congested, sessile, 41-57 x 42-70 cm at maturity, rugose, herba- 
ceous or papery, adaxially opaque dark green, abaxially medium to light green, upper 
surfaces with hairs restricted to a costal channel and resembling the large hairs on ra- 
chis and branchlets, undersurfaces pustulate, indument lax, the abaxial hairs usually 0.5— 
1.0 mm diam., following veins, intermixing with smaller stelliform scales especially on 
the midrib, dark glandular pits bifacially scattered; leaf rachis with indumentum like the 
branchlets; leaflets opposite, decrescent, heteromorphous, the terminal one ob! olate 
basipetally elliptic-oblong then ovate-orbicular, the proximal pair ie and 
amplexicaulous, auricles ca. 1.5 cm diam.,subapical leaflets often the largest, oblanceolate, 
19.5-33 x 6.0-11.5 cm,shortly acuminate at the apex, basally cuneate; venation pinnate, 
inconsistently camptodromous or (brochidodromous), secondaries in 16-25 pairs on 
major leaflets, 5—7 pairs on small leaflets, diverging 45-75° from the midrib then gradu- 
ally arcuate toward the margin, partial intersecondary veins frequently present, tertiary 
nerves scalariform or not, reticulum coarsely areolate, veins impressed on upper sur- 
faces, the midrib immersed, beneath with all veins raised; petiolule absent or the leaflets 
subsessile and costae swollen at the insertion to rachis. Inflorescence unknown. 
Infructescence axillary, emerging from foliate nodes, rachis 6-11 x 5 mm, with hairs like 
the branchlets, bracteate. Fruits indehiscent, solitary, rarely two together, obovoid or glo- 
bose-oblongoid, to 66 x 48 mm; style semi-persistent, stellately hairy at the base, gla- 
brous above; exocarp completely obscured by dense tomentum, the vesture initially 
orange-brown, later reddish-brown, mealy to the naked eye, only with magnification 
discernable as thickened stellate tufts; developing fruits stipitate, the sepals foliaceous, 
disintegrating, adhering to the exocarp, covered by appressed scales with pale setiform 
rays resembling cystoliths; pericarp woody, indurate, odorous, 6-7 mm thick, locules 2, 
each cell monospermous; seed surface distinctly sinuate in transection. 

Distribution and ecology.—Aglaia saxonii is known only from the subcanopy of ad- 
vanced growth forest at Josephstaal, where it is locally common on foothill slopes. All 
populations were seen in submature or ripe fruit, suggesting that the species may ex- 
hibit big bang flowering. 

The plant's stature and distinctive features make it a conspicuous component of 
the Josephstaal vegetation. | Its Suse eptibility to proposed | gging ationsis u inknown, 
but as a fairly tall tree species, A. saxonii would probably oy adversely affected by selec- 
tive forest removal. 

Etymology.—lt is a pleasure to name the new species after Dr. Earl Saxon, the Asia- 
Pacific regional ecologist for the Nature Conservancy (TNC) and the project's senior sci- 
entific investigator. 


Pararypes: PAPUA NEW GUINEA. Manan Province: Josephstaal FMA area, along track to Morasapa W of 
expedition Camp 1 (‘Kumamdeber’), mature growth foothill forest, near GPS coordinates 9504560 
N, 281407 E, 160 m, 29 Jul 1999 (fr), W. Takeuchi, J. Wiakabu, M. Gorrez, & A. Towati 13,462 (CANB, LAE); 


14 SIDA 19(1) 


Josephstaal FMA area, between expedition Camp 1 (‘Kumamdeber’) at GPS coordinates 9504560 
N, 281407 E and Manag Wara at GPS coordinates 950629 N, 28052 E, ca. 160 m,5 Aug 1999 (fr), W. 
Takeuchi, J. Wiakabu, M. Gorrez, & A. Towati 13,765 (K, LAE, NY). 

The sessile leaves of A. saxonii immediately distinguish it from all the Papuasian conge- 
ners. The thickly woody pericarp is also atypical. As noted by Pannell (1992: 11) Aglaia 
species generally have brittle to coriaceous pericarps, but the fruits of A. saxonil are obvi- 
ously lignified and required considerable effort to section with a hacksaw. 

The novelty’s affinity is to A. subsessilis of Borneo, but the latter clearly differs in its 
larger infructescences, thin pericarp, and unilocular-monospermous fruits. 

Aglaia saxonii will key to fork 136 in Pannell (1992: 56-57). It can then be integrated 
to the existing treatment by inserting the following couplet in place of the present cou- 
plet 136 (ibid): 

Leaves sessile; leaflets sessile or pulvinate Aglaia saxonii lakeuchi 
Leaves petiolate; leaflets clearly petiolulate to the existing fork 136 


RUBIACEAE 
Calycosia mamosei Iakeuchi,sp.nov. (Fig. 4). Tyee: PAPUA NEW GUINEA.Mapanc PROVINCE: Josephstaal 
A area, streambed flowing to SW of expedition Camp 1 (‘Kumamde mature growth 
foothill forest with occasional landslip communities, near GPS pee ee ale N, 281407 
160 m, 28 Jul 1999 (fl, fr), W. Takeuchi, J. Wiakabu, M. Gorrez, & A. Towati 13,404 (Hotoryee: LAE; 
isorypes: A, BRIT, CANB, K, L, NY). 
Species haec ab C. kajewskio Merr.& Perry stipulis minorik d 32 mm longis),laminis subter 
furfuraceo-tomentosis indutis, denique nervis lateralibus palicielibus (18-24-jugis) differt. 


Understory shrub, 2.5—3.0 m tall. Branchlets plagiotropic, foliated only near the ends, twigs 
terete in vivo, compressed when dried, furfuraceous at nodes, the scales primarily in axils, 
crowded, reddish-brown and setiform, resembling colleters, other stem surfaces 
puberulous or glabrescent, internodes usually 2-5 cm long. Leaves decussate, elliptic or 
oblanceolate, 23-38 x 5.0-9.8 cm, apically with a short acumen to 1.5 cm long, margins 
entire, base attenuate and equal; venation regularly pinnatiform, camptodromous, sec- 
ondaries 18-24 pairs, arcuate, the central ones diverging 55—60° from the midrib, major 
veins raised above, more prominent below, reticulations prominulous on both surfaces; 
blades fleshy, adaxially epedue very dark green, abaxially pale green, frequently 
discolorous on drying: grayish-green or olivaceous above and brownish-red underneath; 
upper sides glabrous, minutely tuberculate (?cystoliths), undersides furfuraceous on 
principal veins, otherwise appressedly scalelike-hairy on the remaining surface; petioles 
2-5 cm long, puberulent, adaxially channelled or plane, rounded beneath; stipules acumi- 
nate, 22-32 x 9-14 mm, basally connate for ca. 1/4 the overall length, caducous, often 
disintegrating irregularly and leaving a scarious residue, externally pilosulous or glabrous, 
adaxially furfuraceous at the base. kes cau generally monocephaiic, 
terminating branchlets, infrequently al subapical axils but then depauperate, heads 
sessile, hemispherical or depressedly globose, 40-55 mm diam. when fully developed; 
receptacle discoid, densely shaggy; bracts herbaceous, dull, pink to orange, numerous, 
crowded, costate and with anastomosing venation, occasionally marked by linear cys- 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 15 


Ss 


SSS 
Ter SSA 
eS 
PEED 


why 


Fic. 4. Calycosi j Takeuchi, sp.nov. A. Fertile branchlet. B. Flower from side. C. Corolla limb; one lobe removed. D. 
Fl bud wi btending bracteole. E. Submature fruit. F. Pyrenes in cross-section. Scale bars: A, 5 cm; B—E, 5 mm; F, 
3 mm. Drawn from the type by N.H.S. Howcroft. 


16 SIDA 19(1) 


toliths, the outer involucral bracts largest, lanceolate to ovate-orbicular, 20-28 x 14-27 
mm, usually deciduous before fruit set, internal bracts persistent, highly variable in shape 
and size: linear, elliptic, oblong, to broadly oblanceolate, 11.5-26 x 1.5-10 mm, lanate on 
margins and base; floral bractecles oblanceolate, induplicative, bearded with a central 
line of hyaline filaments. Flowers (rehydrated measurements) obscured by the bracts, 
glabrous on all exterior surfaces, pedicels 1.0-1.5 mm long, pilose; calyx synsepalous, 
infundibular, 5.0-5.3 x 2.5-3.0 mm, lobes 2 or 3, obtuse, equal or not, 0.8-2.5 mm long, 
tube adaxially pilose; corolla valvate, pentamerous, cylindrical, 12.5-15.0 x 3.0 mm, lobes 
acute, 2.1-2.5 mm long, inner tube pilose for 2-3 mm below the throat; stamens 5, in- 
cluded, inserted 2 mm below the sinuses, anthers linear-sagittate, dorsifixed, 1.7-1.8 mm 
long, filaments 2.0—2.1 mm, provided with indument like the corolla throat; ovary bilocu- 
lar, completely inferior; stigma 2-fid, lobes oblongish, 1 rm, fimbriate, style 8 x 0.2 mm, 
glabrous, filiform, simple; disk coarsely rugose. Fruits 8-sulcate or smooth, obovoid, 8-9 x 


4—5 mm, exocarp opaquely yellow-orange and glabrous; pyrenes 2, planoconvex, dor- 
sally somewhat irregular but not clearly ridged, the commissural face with two linear 
invaginations into each seed, albumen lacking ruminations. 

Distribution and ecology.—Calycosia mamosei is a shade-adapted species of mature 
forest understories. Numerous individuals were seen on the latest survey, particularly in 
the elevational interval from ca. 80-200 m.The plant favors well-drained substrates but 
is also occasionally found on seasonally flooded ground. 

The new species is endemic to PNG‘s northern (Mamose) region, being represented 
by collections from Madang, East Sepik, and West Sepik Provinces. Although abundant 
at Josephstaal, C. mamosei is probably uncommon in its other localities of occurrence. 

Etymology.—The epithet reflects the plant's presently known range. 


Other specimens examined: PAPUA NEW GUINEA. West Sepik P B i Subprovince, / 
2 km N of Bewani, about 40 km SSW of Vanimo, lowland interest lat. 3° 01'S, long. 141° 10' E, 1 

m, 28 Aug 1982 (fl), Wiakabu et al.in LAE 73,773 (L, LAE). East Sepik Province: Angoram, primary 
rainforest, lat. 4° 04'S, long. 144° 04' E, 25 Jul 1985 (f1, fr), L. Harkink 2 (K,L, LAE). Madang Province: 
Bogia Subprovince, Tanvid River, inland of Malolo Hotel, regrowth forest in moderately swampy 
area, lat.4° 45'S, long. 145° 24’E, 200 m, 18 May 1986 (fl), O.G. Gideon & RJ. Johns in LAE 57,325 (L,LAE); 
Josephstaal FMA area, Guam River near expedition Camp 2, lower slopes of natural-growth foothill 
forest, GPS coordinates 9497596 N, 280100 E, ca.50 m, 11 Aug 1999 (fr), W. Takeuchi, J. Wiakabu, M. 
Gorrez, & A. Towati 13,877 (K, LAE); Josephstaal FMA area, Guam River, low ridge above expedition 
Camp 3,5 of Dumadum village, natural-growth foothill forest, GPS coordinates 9498679 N, 284829 
E,ca.80 m,21 Aug 1999 (fr), W Takeuchi, J. Wiakabu, M. Gorrez, & A. Towati 14,215 (LAE). 


Calycosia is distinguished by large leaves, relatively long infundibular calyces, and a capi- 
tate inflorescence surrounded by numerous bracts. It was earlier regarded as ranging 
from Samoa to the Solomons (Darwin 1979: 38-9). The genus has certain similarities to 
Cephaelis,and also approaches Psychotria condensata under Sohmer's broad concept of 
that genus. The Josephstaal type keys closest to Calycosia in Darwin (1979: 34-35). 
Calycosia usually has a regularly 5-lobed calyx but in C mamosei it is 2(-3) lobed. 
The calyx limb and inflorescence bracts are also reportedly deciduous (ibid: 38-39), but 
they are subpersistent in the new species. Calycosia mamosei is otherwise similar to the 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 7 


assigned genus on other characters, particularly with respect to the long-tubular calyx 
and the capitate, numerously bracteate inflorescence. 

The only other species of Papuasian Calycosia is the Solomon endemic C. kajewskii 
Merr. & Perry, from which the new species can be readily separated by a host of charac- 
ters, the most obvious of these being differences in indument, stipule size, and number 
of lateral veins. 


RUBIACEAE 
Psychotria mayana Takeuchi,sp.nov.(Fig.5). Tyee: PAPUA NEW GUINEA. Mapanc Province:Josephstaal 
MA area, Guam River near expedition Camp 2, lower slopes of natural-growth foothill for- 
est, GPS coordinates 9497596 N, 280100 E, 50-100 m, 12 Aug 1999 (fr), W. Takeuchi, J. Wiakabu 
M. Gorrez, & A. Towati 13,940 (HototyPe: LAE; isorypes: A, BRIT, K, L). 


Species haec ab P melanocarpae Merr.& Perry fructibus albis differt. 


Branched understory shrub, or subarborescent to 5 m height. Branchlets terete, 3.0-5.5 
mm diam., glabrescent, subapically smooth and green, on exsiccatae collapsing and com- 
pressed, fuscous. Leaves fleshy, rugose, adaxially very dark green and glabrous, abaxially 
medium green, lamina discolorous with drying:on both sides orange-brown to rufescent, 
rarely olivaceous, underleaf indument subappressed on costae, otherwise mostly patent, 
occasionally simple, more typically stellately branched or in stelliform fascicles; blades 
oblanceolate, 22-33 x 6-12.5 cm when mature, apex shortly acuminate, at most 
|; venation pinnate, upper surfaces inconsistently im- 
mersed-rugose, manifestly prominent beneath, secondaries equispaced, 12-24, on the 
large laminae always exceeding 15 pairs, straight, diverging at ca.45—60° from the mid- 
rib, supramedially arcuate toward the margin whether or not with closing commissural 
loops, tertiaries subscalariform, obliquely directed at the midrib; domatia absent; peti- 
oles 2-5 cm, adaxially plane, convex beneath, glabrescent; stipules valvate, caducous, 
lanceolate to ovate, 14-20 x 4-10 mm, bifurcately cleft, each lobe 5-9 mm aristate, ex- 
ternally marked by medial ridges insensibly confluent with the aristae, coarsely shaggy, 
inner surfaces densely appressed-hairy. Flowers unknown. /nfructescence strictly termi- 
nal, to 11.5 cm long, ebracteate, paniculiform, ramifications verticillately developed 
through 2-3 orders, the ultimate rachillae cymose, peduncle 1.5-4.0 cm, cernuous, all 
axial surfaces entirely white, with a reddish-brown vestiture of papillate or subulate hairs, 
these mostly spreading, often crisped. Drupes globose or obovoid, 9-10 x 8-9 mm, sub- 
sessile, nitid green, opaquely white when ripe, exocarp glabrescent but with lax hairs 
persisting at the apex and base; calyx coarsely puberulent. Pyrenes 2, equal or not, 
planoconvex, lacking dorsal crests;endosperm with a central lumen, ruminate by irregu- 
lar transversal folds. 

Distribution and ecology.—Psychotria mayana is a small tree growing in stands with 
multistoried canopy. |t is apparently restricted to the Josephstaal area and was collected 
only from the foothill zone. 

Etymology.—lt is a pleasure to name the new species after Maya Gorrez,a biologist 
currently serving with the Washington office of the Nature Conservancy. 


subcuspidate, base attenuate, 


cl 


SIDA 19(1) 


y 


7 


, 


B. Drupe in cross-section. Scale bars: A.5cm:B.5mm 


p.nov. A.F 


r, 
7 


SL 
ne 


ao 
ee 


— 


eatin S 


Fic. 5. Psychotria mayana Takeuchi, s 
Drawn from the type by N.H.S. Howcroft. 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 19 


Paratypes: PAPUA NEW GUINEA. Mapana Province: Josephstaal FMA area, along trail to Morasapa W of 
expedition Camp 1 (‘Kumamdeber’), mature growth foothill forest, near GPS coordinates 9504560 
N, 281407 E, 160 m, 30 Jul 1999 (fr), W. Takeuchi, J. Wiakabu, M. Gorrez, & A. Towati 13,585 (BRIT, LAE); 
Josephstaal FMA area, Guam River near expedition Camp 2, lower slopes of natural-growth foothill 
forest, GPS coordinates 9497596 N, 280100 E, ca.50-100 m, 12 Aug 1999 (fr), W. Takeuchi, J. Wiakabu 
M. Gorrez, & A. Towati 13,944 (K, LAE, NY). 


Among Papuasian representatives of Psychotria, the underleaf hairs assembled in stelli- 
form fascicles (or also basally branching) is a character shared only by P melanocarpa 
Merr. & Perry. The large fruits to ca. 10 mm diameter and the biaristate stipules are also 
characteristic of both species. As suggested by its epithet however, P melanocarpa has 
black fruits, while in P mayana the entire infructescence is white. Psychotria melanocarpa 
is presently known only from Western Province, and P mayana only from Madang Prov- 
ince. They are apparently geographically-separated sister species from opposite sides of 
the Dividing Ranges. 

Psychotria mayana will key to couplet 87 (Sohmer 1988: 24) and to a group of 5 
species consisting of P sphaerothyrsa, P kaniensis, P multicostata, P dolichantha, and P 
ramadecumbens, but is not related to those species. Because Pmelanocarpa differs from 
P mayana in fruit color, the two are far apart on the Sohmer key and their relationship is 
thus not immediately apparent. 

The new plant can be incorporated into the existing treatment by interposing the 
following couplet between forks 86 and 87 (ibid: 24) and then continuing on with the 
same decision train: 


Underleaf indument of stelliform hairs; stipules clearly biaristate Psychotria mayana 
Takeuchi 


eft, but not aristate 


Underleaf glabrous, or pubescent 
to fork 87 


DISTRIBUTIONAL RECORDS AND OTHER NOTEWORTHY COLLECTIONS 


APOCYNACEAE 

Rauvolfia moluccana Markgraf; coll. 14262. Hendrian and Middleton (1999: 457) cite 
three specimens from the Bismarck Archipelago in their recent revision.Most specimens 
originate from Indonesia; the species being rarely recorded in the eastern half of New 
Guinea. Lae Herbarium previously had only one mainland collection in Rauvolfia. 
ARACEAE 

Alocasia lancifolia Engl.; colls. 13852, 14097, 14216. A common aroid species, but not 
previously recorded for Madang Province (Hay & Wise 1991:522). 


— 


Homalomena magna A. Hay; coll. 73849. Previously known with certainty only from W. 
Sepik Province, in the Vanimo and Amanab areas (Hay 1999: 51). 

The species is readily identified by the reddish spathes and dimerous male flowers 
(ibid: 53); characters exhibited by the Josephstaal voucher. 


20 SIDA 19(1) 


COSTACEAE 

Tapeinochilos recurvatum K. Schum.; coll. 13700. The plant is a highly restricted en- 
demic, previously known from ten specimens obtained in the Gogol and South Naru 
drainages near Madang. Clear-cut logging is presently endangering the survival of his- 
torical populations (Gideon 1998: 325). The Josephstaal provenance represents the only 
occurrence whose habitat is not under immediate threat. 


Tapeinochilos sp. nov.; coll. 13743. Gideon (1998: 291) regarded this species as an 
undescribed Tapeinochilos endemic to the central part of northern New Guinea. It was 
previously known from four locations. The Josephstaal population extends the range 
significantly eastwards and is the fifth documented provenance for the novelty. 

The Adelbert foothills around Madang township had earlier been thought to con- 
tain only T. hollrungii, T. pubescens, and T. recurvatum. The discovery of an undescribed 
Tapeinochilos from accessible terrain is yet another indication of the comparatively un- 
explored status of the Adelbert Range. 


EUPHORBIACEAE 

Cleistanthus sp., aff. ?papuanus (Laut.) Jabl; coll. 73672.Possibly new. The collection will 
not key on any combination of characters using Airy Shaw (1980:58-9).It much resembles 
C. insignis in aspect, but the fruits are strigose and the vegetative indument is different 
from that of the latter species. 


Glochidion chondrocarpum Airy Shaw, or aff; coll. 13697. Previously known only from 
several specimens obtained at Mt. Bosavi in southern PNG (Airy Shaw 1978: 372-73). 
Possibly rare. Now recorded on the northern side of the mainland. 

The Josephstaal collection is similar to G. chondrocarpum, but is ramiflorous, unlike 
any of the species in the G. chondrocarpum-decorum-rugulosum group. The survey 
voucher is also vegetatively similar to glaucous, large-leaved species like G. chlamydogyne. 
In its ramiflory, the Josephstaal species seems to form a connection between all the 
preceding taxa and the Strictly cauliflorous G. beehlerii, and could represent a new spe- 
cies. However the paratypes to G. chondrocarpum (i.e., Jacobs 9107 & 9107A) have fruits in 
dense clusters emerging near the main stem, suggesting a transition to cauliflory. Some 
lumping may eventually become necessary in this complex of species with similar fa- 
cies, so it is prudent to preliminarily assign the name of the closest taxon to the present 
collection. 


FLACOURTIACEAE 

Casearia erythrocarpa Sleumer; coll. /3487.Originally known only from the type speci- 
men collected on the Fly River (Sleumer 1954: 87) but more recently discovered in the 
Oomsis-Gabensis areas near Lae (i.e., Henty in NGF 16501,and Takeuchi 7114). 

The species has not been reported in the literature since the time of the Flora 
Malesiana revision (ibid). Its habit as an understory shrub should ordinarily make the 
plant easy to find, so the scarcity of herbarium specimens probably reflects actual rarity 
rather than simple undercollecting. With discovery of the Josephstaal population, the 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 21 


species’ known distribution consists of three disjunct stations with (probably) low fre- 
quencies at each locality. 


ICACINACEAE 

Rhyticaryum novoguineense (Warburg) Sleumer; colls. 13947, 14192. Rhyticaryum 
novoguineense is easily distinguished from congeners by the paniculate inflorescences. 
The plant is restricted to Madang and Morobe Provinces, and at the time of the latest 
revision (Sleumer 1971) was represented in herbaria by three specimens. LAE has only 
two sheets of this taxon. 


PROTEACEAE 
Helicia affinis Sleumer; coll. 13997. An arborescent species known only from lowland 
environments in Madang Province. As a restricted endemic, H. affinis is the kind of plant 
likely to be endangered by introduction of logging operations to the management area. 
The expedition voucher is a fruiting collection and thus cannot be keyed on exist- 
ing treatments (Foreman 1976, 1995; Sleumer 1955). |t resembles both H. /atifolia and H. 
finisterrae in aspect, but the appressedly puberulous underleafis more similar to H./atifolia. 
The surface scrape on the drupe is conspicuously purple, a feature exhibited by several 
Papuasian congeners, though not previously noted for this particular species. 


RUBIACEAE 
Psychotria dipteropoda Laut. & K. Schum.; colls. 73831, 13869, 14045, 14200. Psychotria 
dipteropoda had not been seen for nearly a century until its rediscovery during the TNC 
surveys. The type collection was obtained in the Gogol drainage in 1890, but was subse- 
quently lost during the WWII destruction of Berlin Herbarium. The most recent of the 
surviving collections was obtained in 1907, even though many botanists have visited 
and collected from mie eae S former localities. 

Psychotria dip a occurs only in tall-growth stands beneath intact canopy, on 
the alluvial flats sdiacen? to flowing streams. This kind or Pane is likely to be highly sus- 


ceptible to anthropogenic disturbance; firstly, because riverine borders are environments 


easily altered by human entry into wilderness habitats,and secondly, because its consis- 
tent association with advanced growth shows this plant does not flourish in seral situa- 
tions. Though not a rheophyte, P dipteropoda is apparently adapted to conditions in the 
seasonal surge zone on river verges. The Gogol and Ramu drainages have been seriously 
impacted by habitat alteration since the early 1900s, and this is almost certainly the 
cause for the plant’s disappearance from its historical range. 

Although the Josephstaal colonies represent the only known occurrences of the 
species, it is moderately frequent within the Guam drainage. Most sightings were of ster- 
ile individuals, but could still be identified because of the Gidea blades and the plant's 
consistent association with a narrowly defined habitat. 

The ripe fruit on P dipteropoda is unexpectedly yellow. Fruits of P talasensis have a 
yellow or orange phase, but the drupe matures to a conventional red (Sohmer 1988: 
278), while in P dipteropoda yellow is apparently the ripe color. 


22 SIDA 19(1) 


Psychotria sp. nov.; coll. 13574. An undescribed monocaulous species with a marked 
resemblance to Maschalodesme, except that the fruit has two pyrenes. 


Psychotria sp. ?nov.; colls. 13457, 13756. A vining species. The lianous Psychotria are 
unrevised for Papuasia, having been last treated by Valeton (1927). 

The Josephstaal collections are unusual for their extremely membranous leaves and 
the oblong cystoliths d ly marking all surfaces, including the inflorescence. The flow- 
ers are small, sessile, and glomerulate on lax rachides. Although possibly new, the status 
of this plant can be definitely established only through revision of the vining taxa. 


Versteegia grandifolia Valeton; coll. 13405. A rare species from the pachycaul alliance, 
previously represented by three specimens from West Irian.In their synopsis of the ge- 
nus, Ridsdale et al. (1972: 340) had specified the plant's distribution only as ‘mainland 
New Guinea.’ Collection 13405 keys directly to the above binomial and conforms pre- 
cisely to Valeton’s (1911: tab LXXIII) plate. Although Lae Herbarium has no specimens of 
this species for comparison, the taxon’s distinctive characteristics permit identification 
from the literature. 

The Josephstaal plants were identified as ‘wanam-barewa’ by village respondents. 
When the stems are used as a planting implement, they are said to increase crop yields 
(Wiakabu and Gorrez, field notes). 

Josephstaal villagers are also aware of the distinction between this species and the 
more common V.cauliflora, as indicated by their conferral of the different name’waipa'to 
the latter.In this particular instance, traditional knowledge conforms to formal taxonomic 
concepts in Western science. The belief that V.grandifolia is connected to increased crop 
yields is possibly related to its more robust habit in comparison to V. cauliflora. It can be 
surmised that the oversized leaves from the grandifolia facies have become associated 
with a special capacity for growth, which is then transferred when the larger species is 
used as a planting tool. 


RUTACEAE 

Wenzelia dolichophylla (Laut.& K.Schum.) Tanaka; colls. 13594, 13623.Wenzelia is amem- 
ber of the subfamily Aurantioideae and remains imperfectly understood despite the 
potential horticultural value of its alliance. Only six collections of W. dolichophylla were 
known at the time of the last published commentary on these plants (Stone 1985). 

Swingle (1967) established two subgenera but was unable to assign W.dolichophylla 
into either one due to lack of adequate material. In the ripe fruiting specimens now at 
hand from Josephstaal, the thin seeds with irregular hyaline margins make it clear that 
W. dolichophylla belongs to his subgenus Papualimo. Stone (1985: 214-15) did not pro- 
vide a subgeneric affinity for W. dolichophylla and expressed doubt whether Swingle’s 
seed characters were sufficient for recognition of subgenera. 

The Josephstaal plants have wide leaves with prominent and anastomosing sec- 
ondaries similar to those from the Sepik populations. However the bicolorous blades 
with divaricate laterals otherwise agree with features more generally characteristic of W. 
dolichophylla.The red hesperidium (salmon-pink on 13594 and pinkish-red on 13623) is 


= 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 23 


distinctive; other aurantioid taxa are usually orange-yellow. Since most collections of W 
dolichophylla originate from the Ramu-Gogol drainage, it is very likely that the survey 
vouchers are correctly placed under this binomial. 


Zanthoxylum conspersipunctatum Merr. & Perry; coll. 13636. The tree is a montane 
species from elevations above 1500 m (Hartley 1966: 205) and is primarily known from 
the Highlands Provinces. The expedition gathering is a first record for Madang Province. 
More significantly, the elevation of collection at ca. 160 m is anomalous and represents a 
significant extension of the species’ vertical range. 

The Josephstaal voucher has a number of atypical characters which initially ob- 
scured its generic identity. At the time of collection, spines were not visible on the 
branchlets, the leaflets were epunctate, and the foliage showed no evidence of resinous 
content. /n sicco, spiculate excrescences only became evident after collapse of the 
branchlet. The taxonomic concept for 2. conspersipunctatum sensu Hartley is that of a 
polymorphic complex, since the species consists of numerous distinctive forms which 
cannot be assembled into discretely repeating units. Although the Josephstaal speci- 
men keys to Z conspersipunctatum, it does not match any conspecific LAE sheet in ap- 
pearance, so its status is uncertain. 


STILAGINACEAE 
Antidesma katikii Airy Shaw; coll. 73729. The species is represented in herbaria by few 
collections, having been discovered only fairly recently (i.¢., 1968 by Katik in NGF 32762). 
All gatherings have originated in the Ramu-Gogol basins. During the 1995 Bismarck Mts. 
expedition, large populations were recorded and documented near the 600 m level and 
the species did not appear to be as rare as the small number of collections might sug- 
gest (Takeuchi 1999a: 763).With its discovery at Josephstaal, the distribution of A. katikii 
now extends across both sides of the Ramu drainage and the plant is certainly more 
common than previously supposed, though remaining endemic to Madang Province. 
The recency of its discovery, and the uncertainties over the conservation status of 
A. katikii, are circumstances applicable to many other Papuasian taxa. This situation is a 
natural outcome of the uneven state of floristic exploration in PNG. Botanical collecting 
in Madang Province has been heavily focused on the Gogol and Ramu basins, yet the 
Josephstaal populations are within a mere half-day walk from the principal coastal high- 
way. Clearly, there is much work remaining to be done in the floristic documentation 
even of accessible areas. 


TILIACEAE 

Microcos sp. ?nov.; colls. 13469, 13562, 13732, 13830, 14104. The Josephstaal specimens 
apparently represent a new species distinguished by a bilayered indument of erect simple 
hairs with smaller stellate hairs underneath (most Papuasian taxa are lepidote). The plant 
has been collected in several of the north coast districts and possibly also from the Gulf 
region of PNG. Although undescribed, this is attributable to taxonomic neglect of the 
genus rather than to biological scarcity. 


24 SIDA 19(1) 


ZINGIBERACEAE 

Etlingera sp. ?nov. (series Polyanthae); coll. 73985. A nomenclatural conspectus of the 
genus by R.M.Smith (1986) made many transfers from Achasma, Geanthus, and Nicolaea. 
Most of the Papuasian species were treated by Valeton (1913, 1914) as Geanthus, and 
were depicted with excellent illustrations in early issues of Nova Guinea. 

The flowers of the Josephstaal collection are unlike those from any species for- 
merly included in Geanthus sensu Valeton. Obvious points of distinction are the unusual 
subfoliaceous wings at the top of the staminal column and the flat paired nectaries at 
the style base. 


THE JOSEPHSTAAL FLORA IN OVERVIEW 


Although the Gogol basin near Josephstaal represents one of PNG's better-explored 
localities, its vegetation has been extensively degraded in recent decades by clear-cut 
logging. During the severe drought of 1941,a major part of the Gogol was destroyed by 
fire Johns 1986: 351) so it is reasonable to assume that a significant but unknown part 
of the former flora has already been eliminated, especially when past events are consid- 
ered in light of the alterations occurring today. 

Josephstaal is the largest lowland wilderness remaining near Gogol, and not unex- 
pectedly contains many taxa recorded from the latter locality. The records for Psychotria 
dipteropoda, Rhyticaryum novoguineense,and Tapeinochilos recurvatum, exemplify the simi- 
larities between the Josephstaal and Gogol floras. These similarities can be understood 
in terms of new evidence showing that distributions of New Guinea endemics are corre- 


lated to geohistorical phases of terrane accretion (Balgooy et al. 1996: 201 and fig. 16).On 
the basis of the geological relationship, additional linkages can be anticipated between 
Josephstaal and the adjacent Gogol-Ramu drainages, because the localities were all part 


of the same accretional phase in New Guinea's paleohistory. //ligera novoguineensis is 
thus more likely to be rediscovered at Josephstaal than from the now-disturbed histori- 
cal habitats in the Madang-Ramu area. The species is still known from only three collec- 
tions dating to ca. 1900 (Croft 1981: 201; Duyfjes 1996: 759). With further exploration of 
the project tract, other narrow endemics like Cynometra katikii might also be found. There 
is a distinct possibility that Lauterbachia will finally be recovered within the JFMAA. 
From compilation of completed revisions in the Flora Malesiana series, Balgooy et 
al.(1996:198) concluded that the number of endemic species from northern New Guinea 
is higher than from other Malesian areas. It follows from their conclusion, that explora- 
tion of poorly surveyed parts of the northern sector is likely to uncover substantial num- 
bers of previously unknown taxa. The discoveries from the recent TNC surveys are con- 
sistent with the Adelbert's position in the NE quarter of the Island and the high endemicity 
associated with northern New Guinea as a whole (ibid). There are general grounds for 
predicting that future exploration will yield additional discoveries, especially since the 
higher elevations at Josephstaal have not been examined. Most Papuasian endemics 
are montane species, though from the past emphasis on montane exploration, it is ap- 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 25 


parent that the higher-elevation percentages are overstated in relation to the lowland 
component (ibid: 200; Conn 1994: 125, 128). The discoveries within the project tract are 
partly a consequence of the poor attention historically devoted to lowland environments. 

Because many endemic taxa are shared between the Gogol and Josephstaal locali- 
ties, it is natural to ask why the new plants had not been found during previous surveys 
of the Gogol and Ramu drainages. The novelties are visually conspicuous and with at- 
tributes which should ordinarily have ensured earlier discovery. If the new taxa had in- 
deed once ranged into similar habitats in the Gogol-Ramu basins, they were probably 
eliminated there by the environmental upsets of the 1900s, otherwise they would have 
already entered the botanical record. Circumstances are consistent with the premise 
that population extinctions are occurring in the modern period without the popula- 
tions ever being detected. 

The TNC surveys point to a possibility that Josephstaal environments are refugia for 
remnant populations extirpated from other parts of their range. Future work should be 
deliberately structured in ways to evaluate this presumed status. However the percep- 
tions arising from the recent surveys also need to be weighed against the inadequacy of 
existing information on the New Guinea flora. Despite its status as a center for biotic 
diversification, PNG has the dubious distinction of being one of Malesia’s worst surveyed 
nations (Stevens 1989: 127). On a regional comparison, only the Celebes and Sumatra 
have comparably low collection densities (ibid). While it is generally conceded that cer- 
tain mountainous areas are hotspots for floristic endemism, the low-elevation centers 
are not easy to identify.When documentation coverage is so incomplete, it is difficult to 
be sure if patterns determined by current surveys are real. It may be just as plausible to 
argue that presumed connections between Josephstaal and adjacent areas might be 
overturned, if more information were available on the surrounding region. 

A total of 139 families, 445 genera, and 730 distinct morphospecies, have been col- 
lected at Josephstaal (Appendix 1). An unknown proportion of the flora remains un- 
documented. The 41 alien species (exclusive of cultivated plants) recorded by the sur- 
veys represent 5.6 % of the checklist. For a Papuasian wilderness tract, this is a 
comparatively high count, which can be attributed to Josephstaal’s proximity to provin- 
cial population centers and the resulting exposure to anthropogenic influence.The pres- 
ence of dirt roads into the project sites has no doubt facilitated entry of introduced 
plants.An instructive contrast can be drawn between these alien elements at Josephstaal 
and those from more isolated sites accessible only by air travel. Remote landlocked ar- 
eas in the Lakekamu basin and Bismarck-Ramu Range were recently assessed on the 
same protocols (including attention to weeds) as the Josephstaal inventory;and with an 
equivalent search investment of one-month duration.These latter evaluations produced 
an alien naturalized count of 8 species (1.2% of the total) for Lakekamu (Reich 1998; 
Takeuchi & Kulang eeu and 9 species (1.5 %) for Bismarck-Ramu (Takeuchi 1999a). At 
Crater Mt.a more intensive survey found 24 alien species comprising 1.8% of all recorded 
taxa (Takeuchi 1999b). These figures suggest that collection of alien plants can be prof- 


26 SIDA 19(1) 


itably integrated into future schedules as an independent means for determining the 
relative quality of evaluated habitats. Investigators have an understandable preference 
for focusing on indigens during site assessments, but the adventives may actually pro- 
vide a more practical indicator of habitat preservation and isolation. 

None of the weeds recorded at Josephstaal poses any threat to the environment, 
with most of the introductions consisting of benign herbs restricted to repetitively dis- 
turbed ground (Piper aduncum excepted). The presence of anthropogenous plants i an 
unwanted condition in any wilderness habitat, but alien occurrences are inevitable when 
ever an area is relatively easy to enter. Within PNG, conservation easements and land 
units comparable to Josephstaal are usually very isolated environments, and thus logis- 
tically difficult to botanize. The accessibility of Josephstaal's high-value habitats is a marked 
contrast to these other venues. Although proximity to urban centers Causes greater ex- 
posure to unwanted factors, it can also foster scientific research and community-based 
development, because of the comparative ease of operations afforded by convenient 
access. The combination of biodiverse wilderness with low cost logistics, will enable con- 
sideration and implementation of a wide range of planning alternatives for the project 
tract. 


APPENDIX 1 
LIST OF PLANT TAXA FROM JOSEPHSTAAL 


Voucher source for occurrence record: NGF = collections from the New Guinea Force 
series, P&C = B.S. Parris and J.P. Croxall, T = W. Takeuchi, J.Wiakabu, M.Gorrez, and A. Towati, 
T&S = W.Takeuchi and E. Saxon, sn = sin numéro (without number), SR = sight record of 
taxon known to the project botanist. Other collectors indicated by name. Determina- 
tions by WT unless otherwise noted. 
FERNS AND FERN ALLIES ATHYRIACEAE 
Callipteris Bees (Lamk) Bory; T 13605, 13843, 
Ne 
diantum philippense L., T 13436, 14254 Callipteris en (Blume) J. Smith; 7 13919 
ASPLENIACEAE Ca 55D) aff. spinulosa (Blume) J. Smith; P&C 
Asplenium affine Swartz, affine atum group’ 840 
(cf. Copeland 1949: 220-221, Sledge 1962: Diplazium sp.; 1 13662 
408); 1 13490 BLECHNACEAE 
Asplenium cf. amboinense Willd., ‘amboinense- — Stenochlaena milnei Underw.. T 13891 
papuanum group’ (cf. Copeland 1949: 212); 
T 13649, 13745, 13893 CYATHEACEAE 
Cyathea sp; 1 14118 


Asplenium cuneatum Lamk, ‘affine-cuneatum 
255 


group’; 7 1425 DAVALLIACEAE 
Asplenium nidus L. var. nidus; T 13943 Davallia solida (Forst.) Swartz; SR, occasional 
Asplenium phyllitidus Don ssp. malesicum throughout area 
Holttum; 7 13590 Davallia sp., section Humata; SR, sterile, 
Asplenium subemarginatum Rosenst.; 1 13928 throughout area 
\splenium um Forst. 1 14181 Davallia sp.; KJ. White in NGF 10285, cited in NGF 


Bee duvillae (Bory) C. Chr; T 13450, 13800, files but not found at LAE 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


ee 
Dennstaedtia scandens (Blume) T.Moore;T 14072 
eee: speluncae (L.) T. Moore; T 14107 


EQUISETACEAE 
mum Desf. ssp. debile 
(Vauch.) Hauke; T 13873, 14242 


GLEICHENIACEAE 
Dicranopteris linearis (Burm.f) Underw.;SR, burn 
near Roumirap 


HYMENOPHYLLACEAE 
Cephalomanes atrovirens Presl; T 13546, 13903, 
14170 


LINDSAEA GROUP 

Lindsaea obtusa J. Smith; T 13540, 13894, 13904, 
13935, 14180 

Lindsaea tenuifolia Blume; T 13489, 13529, 13924, 
14182 


Sphenomeris retusa (Cav.) Maxon; 7 14138 


LOMARIOPSIDACEAE 
Bolbitis heteroclita (Presl) Ching; T 13883 
Bolbitis quoyana (Gaud.) Ching; T 13589, 13657 
Bolbitis cf.quoyana (Gaud.) Ching; T 13488, 13615 
Lomagramma cf.sinuata C.Chr,, closer to sinuata 
than L. melanolepis vAV.R.; T 13865, cf. P&C 
61 


3 
Lomariopsis kingii (Copel.) Holttum; T 13665 


LYCOPODIACEAE 

Huperzia cf. squarrosa (Forst. f) Trevisan; T 13693 

Palhinhaea cernua (L.) Vasc. & Franco; SR, foothill 
forest 


MARATTIACEAE 
sare pes (Forst.) Hoffm.;SR, Guam River 
OLEA 


A 
a. biserrata (Swartz) Schott; T 14078 
Nephrolepis (close to) biserrata (Swartz) Schott; 
62 


OPHIOGLOSSACEAE 
Helminthostachys zeylanica (L eae T 14080 
Ophioglossum nudicaule L.f; 7 142 


Ophioglossum pendulum L.; a foothill 
forest 


POLYPODIACEAE 
Aglaomorpha drynarioides (Hooker) Roos; T 
13872 


Aglaomorpha heraclea (Kunze) Copel.; T 14010 

Drynaria sparsisora (Desv.) T. Moore; T 13978 

Goniophlebium percussum (Cav.) Wagner & 
Grether; T 14281 

Lemmaphyllum accedens (Blume) Donk; T 13435, 
13611 


27 
Microsorum linguiforme (Mett.) Copel.; T 13651, 
13900 


aa membranifolium (R. Br.) Ching; T 
13431, 13996 

Microsorum papuanum (Baker) Parris; T 13975 

Microsorum punctatum (L.) Copel. SR, alluvial 
orest 

Platycerium wandae Racib.;SR, alluvial forest and 


p 
Pyrrosia lanceolata (L.) Farwell; T 14018 
Pyrrosia princeps (Mett.) Morton; T 13515, 13660, 
13983 


PSILOTACEAE 
Psilotum nudum (L.) Palisot de Beauvois; 7 14000 


PTERIDACEAE 

Pteris ensiformis Burm.f. 7 14253 

Pteris gardneri (Fée) Hooker; T 14099 

Pteris ligulata Gaud.; T 13626, T 13953, also P&C 
8398 


Pteris pacifica Hieron.; P&C 8387 

Pteris cf. torricelliana Christ, ‘P excelsa Gaud. fa- 
cies’; T 14256 

Pteris tripartita Swartz; T 13650, 14039 

Pteris warburgil Christ; 7 13632, also P&C 8385 


SCHIZAEACEAE 
lywooditnm circi t 


(Burm. f.) Swartz; T 13493, 


14204 
Lygodium dimorphum Copel.;T 14081 
Schizaea dichotoma (L.) Sm.;T 13528 
Tae erie 
Selaginella cf. longiciliata Hieron.; T 14146 
Selagineltla cf. veluti ina Cesati; is 13539 
Selaginella sp., aff V.ANV.R.: 7 13537, 
13906 
TECTARIA GROUP 
Pleocnemia macrodonta (Fée) Holttum; T 13492, 
13684 
Tectaria bamleriana (Rosenst.) C. Chr; T 14002 


Tectaria sp., aff. ?teratocarpa (Alderw.) C. a T 
14079 


THELYPTERIDACEAE 

Plesioneuron tuberculatum (Cesati) Holttum; T 
13633 

Pneumatopteris sogerensis (Gepp) Holttum; T 
13604 

sige a aaa sp., aff. keysseriana (Rosenst.) 

Holtturn; T 13882 

Pronephrium micropinnatum Holttum; T 13412, 

also P&C 8395 


28 


Sphaerostephanos acrostichoides (Desv.) 
Holttum; T 14259 

Sphaerostephanos (closest to) acrostichoides 
(Desv.) Holttum; 7 13664 

Sphaerostephanos ses ianus (Baker) Holttum; 7 
13416, also P&C 

see ie invisus (Forst. f.) Holttum; 7 
14001, 14144 

ae pilososquamatus (v.A.v.R.) 
Holttum; 7 13690 

ees hanos unitus (L.) Holttum var. 
mucronatus (Christ) Holttum; 7 14709 

VITTARIACEAE 

Antrophyum oo eat Port) Kaulf., 
‘callifolium- plex’ (cf. Holttum 


1954:605);T 1 13902 
Vittaria elongata Swartz var. elongata; T 13656, 
14017 


GYMNOSPERMS 


CYCADACEAE 

Cycas soo ana Laut.; SR, ridge near 
Wagadab 

GNETACEAE 

Gnetum costatum K. Schum.; T 13725, 13964, 
14106 


Gnetum gnemon LT 13 

Gnetum gnemonoides an T 13439 

Gnetum sp.; KJ. White in NGF 1 ea cited in her- 
barium log but not found a 


PODOCARPACEAE 

Podocarpus rumphii Blume; KJ. White in NGF 
10293, det. D.de Laubenfels 

Podocarpus cf.rumphii Blume; T 13492 


DICOTS 


ACANTHACEAE 

Blechum brownei Juss. 1 14051 

Calycacanthus magnusianum K. Schum; T&S 
13086, 1 13574 

Dicliptera papuana Warburg; T 13859 

Graptophyllum pictum (L.) Griff; 7&S 13070, 13084, 
T 13720 

Hulemacanthus (Lindau) 
Bremek.; 1 1373 

Hypoestes floribunda R. Br. var.neoguineensis R.M. 

544 


novoguineensis 
0 


Justicia gendarussa Burm. f; 7 13655 

Justicia sp; 1 13429, 13434, 14203 

Lepidagathis cf. royenii Bremek.; T&S 13079 

Odontonema cuspidatum (Nees) Kuntze; T 14735 

Pseuderanthemum he cf.'variabile group’ sensu 
Barker (1986: 146-156); 1 13678 


SIDA 19(1) 


Ptyssiglottis pubisepala (Lindau) B. Hansen; T 
13413, 13658, also Pullen 1067, det. B. Hansen 

Ruellia sp. (Leptosiphonium); I 13427, 13738 

Rungia sp., aff. ?klossii S. Moore; T 14272 


CAE aaa | 


(Nees) F-Vill. facies; T 13889, 14147 
Strobilanthes sensu lato (Hemigraphis sp.); T 
13453, 13583, 13587, 13679 


ACTINIDIACEAE 


Saurauia ferta Warburg; T 13981 

Saurauia oe jana Diels, gee T 14285 

Saurauia ?stichophle bia 
Diels: 714 17 


ALANGIACEAE 

Alangium villosum (Blume) Wangerin ssp. 
ferrugineum (C.T. White) Bloembergen; 7&S 
13103, T 13674, 13828 


AMARANTHACEAE 

Achyranthes bidentata Blume; T 13898 

Amaranthus dubi eee T 13874 

Celosia argentea L.; SR, Guam R. streambed and 
margins 

ANACARDIACEAE 

Buchanania macrocarpa Laut.; 1 13598 

Dracontomelon lenticulatum Wilkinson; T 14012 
also KJ, White in NGF 10312, det.D.Frodin 

Mangifera minor Blume; SR, Guam alluvial forest 

Rhus taitensis Guillemin; KJ. White in NGF 10309, 
det. Ding Hou; also SR at Cam 

Semecarp us brachystachys Merr.& Perry; 7 13740 


sera poten PIU T 13832, 14171, 14250 
K.Schum,; T 1 a 
ANNONACEAE 
Cananga odorata Hooker f.& Thoms.; SR, Guam 
iver 


Cyathocalyx papuanus Diels, or aff; 7 13507 
Cyathocalyx sp. ?nov.; 1 13458 
Goniothalamus aruensis Scheffer; KJ. White in 
8, det. K. Salleh 
Goniothalamus cf.aruensis Scheffer; T&S 13093 
Goniothalamus cf. imbricatus eta T 13475, 
13639, 13840, 13948, 13950, 1 
Goniothalamus sp.; T&S 13058, T 1 ee 
Haplostichanthus longirostris ee van 
Heusden; 7 13835, 14233 
Petalolophus sp., aff. megalopus K. Schum.,; T 
4,13769 


Popowia sp.,aff.| 
13773 

Popowia sp.; 1 13748 

ae sp., aff. ?versteegii (Diels) Merr; T 


(Blume) Endl. 7 13447, 


3500 
ene sp.A; 7 13751, 13857, 14061, 14070 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


Pseuduvaria sp.B; T&S 13054,T 13423, 13733, 13856 

Pseuduvaria sp. C.; T 13507 

cf. Xylopia sp.; T&S 13064, also KJ. White in NGF 
10272 


APOCYNACEAE 
ee scholars ( J R. Br, SR, eu ee 
um Hemsl.; 7 
Cerbera floribunda K. Schane E135 
Melodinus cf.acutus (Markgraf) Na 7 7 13494 
Melodinus forbesii Fawc.; T 1430 
Nei pets citrodorum (Laut.& on Fosb. 
& Sach.; 1 13907, 14207, also KJ. White in NGF 
HOt det, F, Markgraf 
a (Teijsm.& Binn.) Miq.; T 13998 
Parsonsia curvisepala | Schum; KJ. White in NGF 
10318, det. P|. Forster 
Parsonsia lata Markgraf; T 14297 
Parsonsia oligantha (K. a ) DJ.Middleton; T 
14123, 14263 
Rauvolfia moluccana Markgraf; T 14262 
Tabernaemontana aurantiaca Gaud.; T 13510, 
13802 


Tabernaemontana orientalis R. Br. T 13887 

Tabernaemontana pandacaqui Lam., sensu 
Forster (1992:528-529): T&S 13080,T 14205 

Voacanga grandifolia (Mig.) Rolfe; T 14290 


ARALIACEAE 

Gastonia spectabilis (Harms) Philipson; 7 13498 

Mackinlaya celebica (Harms) Philipson; T 13570 

“ee boerlagei (Warburg) Philipson; T 
1427 

Bae novoguineense ena Becc.; SR, 
disturbed areas near Roumira 

Osmoxylon sessiliflorum (Laut.) PrilseGet 713912 

Osmoxylon (closest to) Soe (Laut.) 

Philipson; T&S 10353-B,T 13836 

Schefflera sp.; = A ae in overstory of 
foothill for 

ARISTOLOCHIACEAE 

Pararistolochia schlechteri (Laut.) M.J. Parsons; T 
13926 

pierre at 

(dets. by Forster 


— 


unless otherwise 


P.l. 
indi cat 

Weenies curassavica L.; T 14245, det. WT 

Hoya anulata Schltr; T 14284 

Hoya pottsii Traill; T 13536, 14289 

Hoya sp.; KJ. White in NGF 10323 

Marsdenia velutina R.Br; T 14229 

genera indet.; 7 13786, 14124 

ASTERACEA 

Ageratum ie Lj 713718 

Bidens pilosa L.var.minor (Blume) Sherff; 7 14137 


29 


R/ fake 


1a Martelli; 7 13419, 13612, 13791 
Blumea riparia (Blume) DC.; T 13669 
Blumea riparia (Blume) DC. f. riparia; KJ. White in 
319, det. J. Koster 
Cosmos caudatus HBK; T&S 13073 
Crassocephalum crepidioides (Benth.) S. Moore; T 
13648 


Eclipta prostrata (L.) LT 13793, 13861, 14088 

Emilia sonchifolia (L.) DC. var. javanica (Burm,) 
tfeld; 114244 

Erecht itites valerianifolia (W If) D 


\D 15 
Mikania cordata (Burm.f) B.L. or ae villosa 


Synedrella nodiflora (L.) Gaertner; T 1 oe 14236 


ae arborea Ham. var. mollissima (Ridl.) 
-T&S 13085, 7 14261 
es 7 ae o .) DC, T 13809 


jaa 
ae tt T 13790 
BARRINGTONIACEAE 
Barringtonia apiculata Laut.; Robbins 1667 
Barringtonia calyptrocalyx K. al var. mollis 
Laut; T&S 13067, T 13567, 135 
Barri ingtonia josephstaclensis eee T 13796, 


jae papuana Knuth; KJ. White in NGF 
10250, det. K. Kartawinata 


BEGONIACEAE 
Begonia papuana Warburg; T 13417, 13543, 13545, 
13884 


ie feild Warburg; T 14420, 14422 

BIGN 

ee cf pendron (Andr.) Steen.; T 14246, 
et. K 


eas  dendoph la (Blume) K. Schum.; T 


13766, 1 
BIXACEAE 
Bixa orellana L.; SR, cultivated 


BOMBACACEAE 
Bombax aie L. SR, foothill forest 


BORAGINACEAE 
Tournefortia sarmentosa Lamk; T 14193 


BURSERACEAE 

Canarium acutifolium (DC) Merr.var.acutifolium; 
Kd. . Whit ite in pig 

A.Gray; 7&8 13109,T 14234, also 
KS, ‘White ir in NGF 10325 

Haplolobus floribundus (K. Schum.) HJ. Lam; K/L. 
White in NGF 10275, cited in Leiden determi- 
nation lists but specimen not found at LAE 


30 


CAESALPINIACEAE 

Bauhinia ampla Span.; 

Caesalpinia bonduc (L.) ee ss 

Caesalpinia sumatrana Roxb.; ne across river 
from Cam 

Cassia alata L.; 1 13797 

Intsia bijuga (Colebr.) Kuntze; T 13692 


2 Ralfa: 


14126 

Maniltoa cynometroides Merr. & Perry; KJ. White 
in NGF 12095, det.B. Verdcourt 

Maniltoa plurijuga Merr.& Perry; T 13984 

sea rosea (K. Schum.) Meeuwen; KJ. White 

10226, 10301, det.M. van Meeuw 

Mani ee schefferi K.Schum.& Hollrung:T 13571 
13783, 14011 

CARICACEAE 

Carica papaya L.; SR, ?naturalized 


CARYOPHYLLACEAE 
Drymaria cordata (L.) Willd. ex Roem & Schult.; 
am R 


CECROPIACEAE 
Poikilospermum amboinense Zipp. ex Miq.; F 


, 


— MUTT! 


U x 
q.; | 14041, ‘also KJ. White in NGF 10244 


CELASTRACEAE 
Salacia plies Schum.,; T 14240 


CHLORA ACEAE 
ee dante (Buch.-Ham.) Verdcourt; 7 
13752 


CLUSIACEAE 

Calophyllum sp.;SR, scattered sightings of ster- 
ile individuals ee ut area 

Garcinia cf.celebica LT 14 

Garcinia dulcis (Roxb.) ape T 13 

Garcinia hollrungii Laut.; KJ. Tee NGF 103173, 
de evens 

Gainic naka Laut., or a KJ. White in NGF 
10267, det. PF. Steven 

Garcinia maluensis ne 
13571 

Garcinia sp.; KJ. White in NGF 10257 


COMBRETACEAE 
Terminalia complanata K. Schum.; T 13991, also 

KJ. White in NGF 10266, det. MJ.E. Coode 
Terminalia impediens Coode; T 13951, 13952 
CONVOLVULACEAE 
Ipomoea congesta R.Br; T 13722 
Merremia peltata (L.) Merr; [13726 

lina riedeli (Oliv.) Oostroom; T 14235 


T&S 13083, T 13477, 


vP 


SIDA 19(1) 


CUCURBITACEAE 

Alsomitra macrocarpa (Blume) Roem.; SR, Camp 
3, cf. Pullen 1096 

Cucurbita sp, T1387 

Trichosant Ze sp., longiflora-bracteata group (cf. 
Harms 1925: 159); 7 1397 

Zehneria ene (Blume) Miq,.;7 13609, 13954 

DATISCACEAE 

Octomeles sumatrana Miq,.; SR, very common 
emergent in alluvial zone 


DICHAPETALACEAE 

siaeann sessiliflorum Leenh.,;T 13851, 14065 

DILLENIACEAE 

Dillenia sete ifolia ye Martelli ex Dur. & 
J 13050,T 14 

Tetracera nordtiana FVM; 3905 

DIE TEROSAREASE AE 

rth.) Blume; T 14208 


EBENACEAE 
Diospyros papuana Valeton ex Bakh,; T 13577, 
13915 


Diospyros pulchra Bakh.; K.J. White in NGF 10261 
Diospyros rostrata (Merr.) Bakh.; 7 13764, det. K. 
Damas 


oe 
eratium ieee T 13671, 13946 
amplifolius a KJ. White in NGF 
10256, det. M.J.E.Coo 
Elaeocarpus sphaericus eal K. Schum.; SR, 
amp 2 
Sloanea cel Baker f., 7 14063 


EUPHORBIACEA 

Acalypha grandis penth: T 14303 

Acalypha hellwigii Rial oar . ee mollis 
(Warburg) K.Schum. & L 

Actephila lindleyi (Steud.) Mes iba T 13668 

Aporosa (probably) papuana Pax & Hoffm,; 7 
13890 


Ela q 


Breynia cernua (Poir.) Muell. Arg,; 7 13614, 13723, 


Bridelia macrocarpa Airy Shaw; KJ. White in NGF 
10280, det. J.R. Croft 
a sp., aff. ?papuanus (Laut.) Jabl.; 7 


oa ie lean (L.) Blume var. 
moluccanum (Decne) Muell. Arg.; T 13746, 
14300 

Codiaeum sp.; T 13742, 14035, det.K. Damas 

Drypetes longifolia (Blume) Pax & Hoffm.; KJ. 
White in NGF 10313A 

Endospermum moluccanum (Teijsm. & Binn.) 
Kurz; 1 13863, 13980 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


Euphorbia oo L; 7 14089 
Euphorbia hirta LT 13798 
Euphorbia pumas ee ex Hassk. var. 
acuminata JJ.Sm.; 7 
Fahrenheitia sp. ?nov., T baie 
Glochidion chondrocarpum Airy Shaw, or aff; T 
9] 


Glochidion granulare Airy Shaw; T 13557, 13734, 
13966 

Glochidion lobocarpum (Benth.) Bailey; T&S 
13092- 

Glochidion novoguineense K. Schum.; T 14125, 


4155 
— ies to) perakense Hooker f. var. su- 
pra-axillar (Benth.) — Air Shaw, 
ee -perakense complex’ (cf. Airy 
Shaw 1975:125, 1980: 106-107); T 14073 
Macaranga aleuritoides Fv.M.;T 14160-B 
Macaranga fallacina Pax & Hoffm.; / 1 13541 
Macarang a denia Pax & Hoff; i le 
inN 4, this staminate may 
be eae with T 14105, M, ee 
andulosa Warburg var. 


Macaranga subpeltata Laut.& K.Schum.;T 14105 
pete sp.; KJ. White in NGF 10260 
Manihot esculenta Crantz; T 13785 
Melonoep Hee ele (Reinw. ex Blume 
Reichb f.& Zoll.; T 14036 
Cay ae a (Warburg) K 
um.; 7 13956, 14071 
Omphalea queenslandiae FM. Bailey; T 13572, 


ra 


1417 
Phyllanthus rubriflorus J.J.Sm.; 7 13688, 13752 
Phyllanthus urinaria LT 14 
Pimelodendro ee Hassk.; 7 13923 
Ricinus communis LT 14 
EUPOMATIACEAE 
Beatie ie R. Br; SR, occasional in foot- 
hill for 


Ans pales Tha ssp. pulchellus; 1 13724 
jum mu T 13799 
osema pubescens Benth.; 1 13805, 13814 
eee pallida gay a 14015, 14140 
Crotalaria retusa L.; T 
ia 5 koolgbberah E . iy ssp. koolgibberah; 


ee 


ae ormocarpoides DC.; T&S 13090, T 
14296 


Desmodium umbellatum (L.) DC; T 14129 
Flemingia macrophylla (Willd.) Merr; T 14132 
Flemingia strobilifera (L.) R. Br.ex Aiton f; 114265 
Inocarpus fagifer (Parkinson) Fosb.; T 13560, 


31 


13914, KJ. White in NGF 10284, det. P. van 
R 


Inocarpus sp. ?nov., ‘rubidus' morphotype fide 
Verdcourt (1979: 304-305); T 13823, 14210 

Mucuna cyanosperma K. Schum.; T 13829 

Mucuna novoguineensis Scheffer; T 14174 

Phaseolus lunatus L.; 7 14257 

Phylacium bracteosum Belly ;1 13909 

Pterocarpus indicus Willd R frequent sightings 
throughout area 

Pueraria phaseoloides (Roxb.) B 

th.) Baker; T 14069 

Pueraria phaseoloides (Roxb.) Benth. var. 
phaseoloides; T 1424 

Pueraria pulcherrima (Koorders) Koorders- 
Schumacher; T&S 13072, T 13838 


FLACOURTIACEAE 
Casearia erythrocarpa Sleumer; T 13481 
asearia macrantha Gilg, or aff; T 13517, 13619 
Casearia sp. ?nov.; T 14150, 14173 
Erythrospermum candidum (Becc.) Becc.; T&S 
13060, T 13495, also KJ. White in NGF 10229 


var. javanica 


Flacourtia inermis Roxb.; T 13502, 14176 
ae cal foetidum (Roxb.) Benth.; SR, alluvial 
for owers present on 7 1394 
osmelo php (Turcz.) Benth.;T 13922, 14046, 
o K.J.White in NGF 10281 
gi ae Reinw.; 1 13646 


GENTIANACEAE 
canon tenuis Blume; T 13524, 13911 


GESN AE 

Gytond batt Warburg; T 13989 

Cyrtandra s ection Fe T 13731, 
ar 13888, 13901, 13958, 14. 

Cyrtandra sp., between si ee 
and Loxanthe; T 13433, 

Rhynchoglossum papuae Be T 13613, 14148 

HERNAND qaanan 

BU sae uam R., fruits and 
lea cond at eee ee 

ICACINAC 

eae laxiflora (Miers) Howard; T 13588, 
14199 


oe scandens Becc.; T 14276, cf. Pullen 


al DiI A\ Cc] 
xe 


GF 10298, det. J. Womersley 
oe i trys dorae Moeser; T&S 13055,T 13945 
hyticaryum longifolium K. Schum. & Laut, T 
3537 


KJ. White 


| epee 7 VATA \cl 


113947, 14192 


32 


LAMIACEAE 

Faradaya splendida Fv.M.; 1 14054 
Hyptis capitata Jacq.; 1 13663, 13774 
Ocimum gratissimum LT 13813 


LAURACEAE 

Actinodaphne sp., (possibly) nitida Teschner; SR, 
foothill forest 

Alseodaphne archboldiana (Allen) Kosterm,; .C. 

ders 946 

Cryptocarya as Blume; T 13486, 13599, 
13753, 1418 

Cryptocarya ne (Oken) Kosterm.; SR, 

Wagadab transect 

Cryptocarya multinervis Teschner; T 14282 

Cryptocarya weinlandii K. Schum.; KJ. White in 
NGF 10243, 10306, dets. A. Kostermans 

Endiandra grandiflora Teschner; KJ. White in NGF 
10269, det. A. Kostermans 

shes magnilimba Kosterm.; T 14114, KJ. 

in NGF 10270 (type), 10293, det. A. 

ae 

Endiandra squarrosa Kosterm.; KJ. White in NGF 
10252, 10307, also R.G. Robbins 1625 

Litsea sp.,‘L.calophyllantha K.Schum. facies’; SR, 
foothill forest 

ane indet.; 1 13763 


LEEA 

Lea (closer to) eo Panne Laut.,'coryphantha- 

(cf. Ridsdale 1974: 78-79): 
T 13581, 13624, 13982 

Leea heterodoxa K.Schum.& Laut.; 1 13446, 13452 

Leea indica (Burm.f.) Merr.; 7 13421, 13959 

leea aiaaens Miq.; T 14032, 14201 

LOGA 

Pa cei pie Thunb, 713 

Fagraea elliptica Roxb.; SR, foothil forest around 


4 


amp 
Fagraea racemosa Jack ex Wall; SR, alluvial for- 
along Guam R. 

Geniostoma rupestre J.R. & G. Forst. (closest to) 
var. rupestre; T 13484, 13538, 13593 

Neuburgia corynocarpa (A. Gray) Leenh. var. 
corynocarpa; T ee also KJ. White in NGF 
10299, det.B. Con 

Neuburgia ae Leenh.; 7 13696, 14029 

LORANTHACEAE 

Amyema olla _ Schum 
seemeniana; T 142 

Decaisnina holl ungll a Schum.) Barlow; SR, hill 


.) Danser ssp. 


orest 
Dendrophthoe curvata (Blume) Miq.; SR, near 
Roumirap 


SIDA 19(1) 


LYTHRACEAE 

Lagerstroemia cf. piriformis Koehne; T&S 13100 

MAGNOLIACEAE 

Elmerrillia tsiampaca (L.) Dandy ssp. tsiampaca 
tsiampaca; SR, foothill forest 


eee, 
Hibi oldianus 5. SR, from lepidote 
pene on note foothill forest 
Hibiscus Borss.; 1 13509 


Hibiscus tiliaceus L.; SR, transect at Wagadab 
Sida rhombi ifolia L. le rhombifolia, T 14014 


Thes sicalyx Borss.; T 13739, also KJ. White 
in NGF 1 10297, det. P Fryxell 
MELASTOMATAC 


Medi inil la musofo ae & K.Schum, 1 14708 
r ogn., ‘musofo-triplinervia 
p (cf. Mansfeld 1925: 116);7 13418, 14228 
Medinilla ae - gaia Baker f,; 7 13566, 
13617 
Memecyl on cadre oe T&S 13052 
Memecylo aff. ?papua err. & Perry, 
‘excelsum- Beale te ee (cf. Merrill 
Perry 1943: 439); T&S 13052 
Otanthera bracteata Korth.; 7 13968, 14271 
MELIACEAE 
Aglaia agglomerata Mert. & Perry, small fruited 
form; T 13685 
Aglaia argentea Blume; T&S 13071 
Aglaia cuspidata C.DC.;T 13918, also KJ. White in 


ae ee Harms; T 13470, 13584, 13694, 


Gi es ee Harms; 1 1397 
Aglaia ee (Fv.M.) Harms; a 13092-A, I 
1368 


Aglaia saxonii Takeuchi, T 13462, 13712, 13765 


Aglaia sp., aff. Merr.& Perry; T 13833, 
14102 
Aglaia sp.; T&S 13] 
Aphanamixis hya (Wall.) R.N. Parker; T&S 
13110, T 13501, ae aes 
hisochet qa is (Mig.) C.DC.; KJ. White in 


3,det.D.Ma bber ley 
Chisocheton lasiocarpus (Miq.) Valeton; T&S 13112 
Chisocheton pohlianus Harms; | 13424, 13474, 
oe 13485, 13999 
brassii Merr. & Perry; T 13715 
Bee excelsum Blume; T 14301 
Dysoxylum latifolium Benth, T 14098 
Dysoxylum pettigrewianum F.M. Bailey; T 13601, 
13784, 14220,also KJ. White in NGF 10320, det. 
D. Mabberle 


Pee 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 

Dysoxylum sparsiflorum Mabberley; T 13556, 
13806, 13827, 14056, 1 

Dysoxylum variabile Harms; T 14298 

Dysoxylum sp.; T 14306 


MENISPERMACEAE 
Arcangelisia flava (L.) Merr.; T 13761 
hlaenandra ovata Miq,.; T 13519, 13777 
Parabaena tuberculata Becc.; 7 14 
es end Sp., ee tumefacta 
group (cf. Forman 1986: 173); T 140 
Tinomiscium petiolare Hooker f.&Thoms.; 1 14047 


MIMOSACEAE 
Archidendron aruense (Warburg) de Wit; T 13526, 
13603, 14021, 14191, also KJ. White in NGF 


10286 
es cf. aruense (Warburg) de Wit; T&S 


fechidencon bellum Harms; T 13707 

n lucyi Fv.M.; es 
ai Biles (L.) Merr:T 13607 
Leucaena leucocephala (Lamk) de Wit; T 14143 
Mimosa diplotricha C. Wright ex Sauvalle var. 


dica L SR, G 
Paraserianthes Pa ee 
sp. falcataria; T 1373 


MONIMIACEAE 

Steganthera dentata (Valeton) Kaneh.& Hatus,; 7 
13934, 13939 

Steganthera hirsuta (Warburg) Perkins; T 13670 

Steganthera hospitans (Becc.) Kaneh. & Hatus. 
T&S 13095, 7 13822 

MOR 

Antares s ane K. Schum.; T&S 13099, T 
13 

ee communis J.R. & G. Forst.; T 13848, 
14160-A 


mR. 
4 .) Nielsen, closest to 


Artocarpus vriesianus Miq. var. refractus (Becc.) 
Jarrett; T 14006 

Ficus ampelas Burm.f, T 13727 

Ficus arbuscula Laut.& K.Schum,; SR, Guam R. 

Ficus bernaysii ae T 13713, 14219 

Ficus botryocarpa Miq. var. subalbidoramea 
(Elmer) eas 13634, 14225 

Ficus comitis King; T 

Ficus congesta oe T 13600, 14164 

Ficus conocephalifolia Ridl.; T 13597 

Ficus copiosa Steud.; T 14043 

Ficus crassiramea Miq. var. patellifera (Warburg) 
Corner; T 14212 

Ficus dammaropsis Diels var. obtusa Corner: T 
13913 


Ficus erythrosperma Miq.; T 13108 
Ficus gul Laut. & K.Schum.; T 14134 


Ficus hesperidiiformis King; T 13979, 14278 

Ficus hystricicarpa alee 71 13520, 13792 

Ficus mollior Benth.; T 

Ficus odoardi King; Se) 1391 

ae pachyrrachis Laut.& K. Schum.; KJ. White in 


Fic ee Laut.& K.Schum.; 7 13967, 13971 
Ficus polyantha le T&S 13103,T 14161 
Ficus primaria Corner; KJ. White in NGF 10274, det. 

.Corner 
Ficus primaria Corner, or aff; T 14 
Ficus pungens Reinw.ex Blume; T 13880, 14116 
Ficus subcuneata Miq.; T&S 13107,T 14189 
Ficus subulata Blume; T 13818, 13995, 14266, also 
KJ. White in NGF 10273, det.E. Corner 
Ce wassa Roxb.; T 13549, 13760 
cus sp. A, oe i key; | 14020 
ie sp.B;T 14 
a ee venenosus (Zoll.& Mor.) Becc. ssp. 
papuanus (Becc.) Jarrett; 7 14059 
Prainea papuana Becc.; KJ. White in NGF 10230, 
det. by KJ.White 


bt a reccas 
ocomia macrocoma ee oe Wilde ssp. 
prainii (King) de Wilde; T 14 
Gymnacranthera Arete ake: f. & 
Thoms.) Warburg var. zippeliana (Miq.) R. 
Schouten; SR, Wagadab transect in hill for- 


est 

Horsfieldia basifissa de Wilde; T 13762, also KJ. 
White in NGF 10242 (type) 

Horsfieldia hellwigii (Warburg) Warburg; T&S 
13065 (sterile) 

Horsfieldia hellwigii (Warburg) Warburg var. 
brachycarpa de Wilde;K.J.White in NGF 10258 

Horsfieldia laevigata (Blume) Warburg cf. var. 
novobritannica (J. Sincl.) de Wilde; KJ. White 
in NGF 10263 

Horsfieldia cf. pulverulenta Warburg; T&S 13101 
(sterile 

Horsfieldia sepikensis Markgraf; KJ. White in NGF 

10237, fide de Wilde's (1985: 81) synonymy 
Horsfiel Hasubiils (Miq.) Warburg;T 13787, 14050 
Horsfieldia subtilis (Miq.) Warburg var. subtilis; T 
13445, 13508, 14183 

Horsfieldic ia ie eae eet KJ. White 
in NGF 10262, det. J. Sin 

Myristica eee ana Wars T&S 13078, T 
14295, also KJ.White in NGF 10259 

Myristica cylindrocarpa J. Sincl.; KJ. White in NGF 

88, det. J. Sinclair, conf. D.F 
acai Tae 


eed 


Myristica fissiflora de Wilde ssr 


t 


34 


Myristica lancifolia Poir. ssp. lancifolia; T 13497, 
, 13824, also KJ. White in NGF 10235, 


10265 

oe subalulata Miq. var. subalulata; 1 13644, 
also K.J. White in NGF 10251 

Myristica tristis Warl Durg, OF aff; T 13487, 13770 


MYRSINACEAE 

Ardisia ese Schum.; T 134617 

Conandrium polyanthum (Laut.& K.Schum.) Mez; 
T&S 13106 


Maesa rufo-villosa Mez; T&S 13091,T 14198 


MYRTACEAE 

Decaspermum bracteatum (Roxb.) AJ. Scott var. 
bracteatum: T 13819 

Decaspermum neurophyllum Laut. & K. Schum.; 
&S 13087 


Octamyrtus behrmannii Diels; T1471 

Syzygium aeoranthum (Diels) Merr. & Perry; T 
13666, 13858 

Syzygium cf, amplum Hartley & Perry; T 13430, 
1422 

Syzygium buettnerianum (K. Schum.) Niedenzu; 


SR, Camp 3 

Syzygium coalitum (Greves) Hartley & Perry; T 
14142 
gium hylophilum (Laut. & K.Schum.) Merr. & 
Perry, or aff: T 13564, ee - en 1094 as cf. 
hy at lum, det. 1.G. Har 


Merr.& P ee T 13 

Syzvoil um m madangens ae & aa a White 
in NGF 10300 (t 

Syzygium nutans (K. a) Merr. & Perry; T 
13661, 14003 

Syzygium pteropodum (Laut. & K.Schum.) Merr. 
& Perry; T 13855, also KJ. White in NGF 10264, 
det. T.G. Hartley 

Syzygium trachyanthum (Diels) Merr. & Perry; T 


pd um tri ivene Merr. & Perry; 7 14710 
Sy a err.& Perry; KJ. White 
NGF 10245 e dev TG rtle 

ea ium Ee aff tne (Diels) Merr. & 
Perry; 1 1345 

Syzygi um <p bee aff. megistophyllum Merr. & 
Perry; T&S 13068 

Syzygium sp.;T 13437, 14214,probably conspecific 

Syzygium sp.; KJ. White in NGF 10239, cited in 
herbarium log but not found at LAE 


NYCTAGINACEAE 

Pisonia longirostris Teijsm.& Binn.; 7 135617, 
13837, 14040 

Pisonia muelleriana Warburg; | 13667 

Pisonia umbellifera (.R. Forst.) Seemann; Pu 
1063, det.D. Frodin 


13565, 


oO 


SIDA 19(1) 


OCHNACEAE 

Schuurmansia henningsii K. Schum. SR, 
occasional throughout area 

OLACACEAE 

Anacolosa cf papuana Schellenb., T 13868, det.K. 
Damas 


OLEACEAE 

Chionanthus ramiflorus Roxb.; T 14308 

ONAGRACEAE 

Ludwigia octovalvis 

OPILIACEAE 

Opilia amentacea Roxb.; T 13735 

OXALIDACEAE 

Averrhoa bilimbi L.; T 13547, 13931 

PASSIFLORACEAE 

Adenia heterophylla (Blume) Koorders; SR, near 
gadab 


(Jacq.) Raven; 7 14025 


PIPERACEAE 
Piper aduncum L.;SR,Guam R.and foothill forest 


iper cf.caninum Blume; T 13810 
Piper celtidiforme Opiz; T 14030, 14058, 14292 
— ee yum (Rumph,) lL; 7 13575, 13942, 


i cane iper Pennant; 7 14016 
Piper (por eae macropiper pee : 4122 


p; 


Piper majusculum Blume; T 13704, 1 
Piper mestonii iFM, Bailey; T 13801 
Piper plagiophyllum K.Schum. & Laut; 7 13610 
Piper pseudoamboinense C.DC; 1 13552, 14024 
Piper cf. pseudoamboinense C.DC.; 1 14084 
Piper pullibaccum Trelease; 1 13677, 139/70, 14166 
Piper versteegli C. DC; SR, Guam R 
PITTOSPORACEAE 
Pittosporum ferrugineum Aiton f. ssp. laxiflorum 
de; 1 14294 
Pittosporum sinuatum Blume; KJ. White in NGF 
10238, det. R. Schodde 
Pittosporum sinuatum Blume var. sinuatum; T 
441, 135471 


yo s 


POLYGALACEAE 
Eriandra fragrans van Royen & Steen,; 1 13455, det. 


.Damas 
Xanthophyllum papuanum Whitm.ex Meijden; 7 
14154, also KJ. White in NGF 10292, det. T.C. 
Whitmore 
PROTEACEAE 
Finschia chloroxantha Diels var. macrocarpa 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


oa KJ. White in NGF 10322, det. KJ. 


Whi 
Helicia ee Sleumer; T 13602, 13997 


RHAMNACEAE 

Alphitonia excelsa (Fenzl) Reiss. ex Endl., sensu 
Schirarend (1995: 308-311);SR, common in 
regrowth 

Gouania cf javanica Miq.;T 13721 

Ziziphus tae ae Hatus.; SR, infre- 
quent in hill for 

Ziziphus ee i T 13938 
RHIZOPHORACEAE 
troch iHlaris Bl R throughout area 


ROSACEAE 
Prunus dolichobotrys (K. Schum. & Laut.) 
2 


s L. var. discolor (Blume) 
Kalkman; 7 13842, 14111 


RUBIACEAE 

Airosperma psychotrioides Laut. & K. Schum,; T 
13449, 13504, 13969 

Amaracarpus grandifolius Valeton, or aff.; T&S 
13082, ee 13 ee 

i ., aff. ‘attenuatus-heteropus 

but re those aan cf. Merrill and 
ae (1946: 221) group 1;7 13456 

Amaracarpus sp.,aff. i ngifolius Valeton; T 134 
13682, 13 932 

Amaracarpus sp., Be, and Perry's (1946: 221) 
group 2;T 14273, 14277 

?Amaracarpus spp.; 1 13767, 

Calycosia mamosei Tea : fone 13877, 
14215 


Canthium sp.; T 13438, 13744, also KJ. White in 
NGF 10249, det. C. Ridsdale 

Hedyotis sp., cf. f auricularia L., or H. lapeyrousit 
DC.;T 1341 

Hydnophytu ect Becc,; T 14005, 14283 

lxora sp.,section Hypsophyllum; T 13728, 13972 

— ee chlorocarpus K. Schum.; T 13480, 


cf, Neonauclea sp.; KJ. White in NGF 10243 
Porterandia sp.; T 14133, also KJ. White in NGF 
10311 


Psychotria amplithyrsa Valeton; T 13411, 14076 

cone dipteropoda Laut.& K.Schum,;7 13831, 
13869, 14045, 14200 

Psychotria sey var. leptothyrsa; T&S 
13062,T 13476, 1 

ory mayana Saeera 13585, 13940, 

eee a Bartl.ex DC; T 13521, 


Psychotria mirlabost a & K. Schum.) 
Valeton; T&S 13098,T 
ies ‘micrococca oe & K. Schum.) 
Valeton; T&S 13088 
Psychotria olivacea Valeton; SR, foothill forest 
Psychotria phaeochlamys (Laut. & K. Schum.) 
Valeton; T 13625, 13631, 13781, 13937 
Psychotria sp., aff. micralabastra (Laut. & K 
Schum.) Valeton; T 13629, 13920, 14085 
Psychotria sp.nov.; 1 13514 
pele sp. (vining, possibly nov.); T 134517, 
3756 


Pree sp., ‘decora Val., or sphaerocarpa K. 
Schum. facies; the genus is now recognized 
only for the neotropics (cf. Puff & Wong: 
1993: 29);T 13516 

‘Randia'sp.,cf.or aff.schumanniana Merr.& Perry 
(R. speciosa K. Schum.); KJ. White in NGF 

10236, 10302, dets. C. Ridsdale 

Saprosma subrepandum (K. Schum. & Laut.) 
Valeton; T 13930, 14169 

Spermacoce assurgens Ruiz & Pavon.; | 14153 

Tarenna gulcheriana (K.Schum. )Valeton; T 13483, 
13749, 13788, 13864 

Timonius timon (Spreng.) Merr.var.timon;T 141 

Timonius sp., aff. densiflorus Valeton; T 14270 

Trukia sp., Oe dryadum (S. Moore) Fosb.; T&S 
13089 


Uncaria bernaysii Fv.M.; T 14249 
Uncaria lanosa Wall.; T 14127 
aie SP. i me og 


Most enc pachyclados (K.Schum.) Melch 
achyclados; T 1 pe also KJ. White in 

NGF 03 05, det. S. Darw 

Morinda bracteata Roxb.;K. Write in NGF 10232 

Morinda umbellata L. var. papuana Valeton; T 
13789, 13921 
njendna ylind q 
3817 

Mussaenda scratchleyi Wernh.; SR, Guam 

Mycetia javanica (Blume) Reinw. ex Korth.; T 

44, 13616 
Nauclea orientalis L.; SR, Guam R. 


pa Burck; T 13627, 13706, 


se oe hum. & Laut.) Valeton; 
TI 13460 3949 
ese de ig ie T 13405 


RUTACEAE 
aes haters R.& G. Forst,; T 13608, also KJ. 
ein NGF 10278, 10279 
ba amara Blanco var. amara; [&$ 13069, T 
13448 


Melicope sp., cf. M. burttiana Stone or M. 
grandifolia B.L. Burtt; T 13936 

Micromelum minutum (Forst.f.) Wight & Walker- 
Amott; 7 13811 


36 
Wenzelia dolichophylla (Laut.& K.Schum.) Tanaka; 
13594, 13623 


Zanthoxylum conspersipunctatum Merr. & Perry; 
T 13636 


SABIACE 

Meliosma_ pinnata  (Roxb.) Ss 
macrophylla (Merr.) Beus.; SR, ool forest 

eee Blume; T 13897, 14165 


SANTALACEA 

Scleropyrum  pateetn (Laut. & K. Schum.) 
Pilger; T 14052 

SAPINDACEAE 

Allophylus cobbe (L.) Raeuschel; SR, alluvial forest 

Arytera sp., aff. litoralis Blume, ‘litoralis complex’ 
(cf. Turner 1994:474); T 13472, 13596 

Cardiospermum halicacabum L.; SR, infrequent in 
regrowt 

Cupaniopsis macropetala jy . 3465, 13591 

Dictyoneura obtusa Blume; 

Seance obliquinervis fad, : ; 13741 


Guioa comesperma Radlk. 
Gua rig ic a Radlk., or va cd luscula complex’ 
elzen 1994: 993); 7 ee 


om 


Lepi ae ee ensis cae ieee : 13683, 

13699, 13933, 13941, also KJ. White in NGF 
et. D.Frodin 

Pometia pi innata Forst.; T 14275 

SAPOTACEAE 

Pouteria maclayana (F.v.M.) Baehni; 1 14287 

SCROPHULARIACEAE 

Limnophila rugosa (Roth) Merr.; 1 13895 

Lindernia anagallis (Burm. f.) Pennell; T 
13794-A 


Lindernia crustacea (L.) Fv.M.; 1 13794-B, 14151 


SOLANACEAE 

Capsicum annuum L. var. annuum; SR, 
cultivated 

Nicotiana tabacum Lt o ae 


Physalis minima ale 114 
Solanum torvum sie a 3807, 14157 


STERCULIACEAE 

Se ia bartramia (L.) Merr; | 14707 

Klei pita L.; SR, along Guam R. 

‘Mele id na (Houtt.) Stapf; T 14293 

Pterocymbium beccarii K.Schum. T&S 13097, also 
Pullen 1071 (cited by P. van Royen 1964: 26) 
but possibly a KJ.White coll, specimen not 


seen 
Sterculia (closest to) ampla Baker f, 7 13768, 14251 


SIDA 19(1) 


Sterculia edelfeltii Fv.M.; KJ. White in NGF 10321, 
oyen 
Sterculia schumanniana (Laut.) Mildbr; T 13466, 


STILAGINACEAE 
Antidesma katikii Airy Shaw; T 13729 
Antidesma rhynchophyllum K.Schum.; T 14033 


THYMELAEACEAE 
Phaleria coccinea (Gaud.) Fv.M,; T 13499, 13503, 
13506, 13782, 14137, 14175, 14202, 14231, 


TILIACEAE 
— argentata Burrett; KJ. White in NGF 
4, det.P. van Royen 
Microcos — argentata Burrett; KJ. White in NGF 
10271, label reads ‘same as 10234' 
Microcos sp. ?nov.; T 13469, 13562, 13732, 13830, 
14104, also KJ. White in NGF 10231 
Microcos sp.; KJ. White in NGF 10231, 
ferent from previous spp. 
Trichospermum tripixis (K. Schum.) Kosterm.; 7 
14156 


10271, dif- 


Triumfetta rhomboidea Jacq.; 1 14091 
ULMACEAE 
Celtis latifolia (Blume) Planch.,; T&S 13102,T 13987 


URTICACEAE 

cf. Boehmeria platyphylla D.Don; T 14087 
Cypholophus cf. nummularis Winkler; T 14049 
Dendrocnide cf. corallodesme (Laut.) Chew; T 


1402 
Dendrocnide cordata (Warburg ex Winkler) 
Chew; IT 13992 
Dendrocnide nervosa (Winkler) Chew; T&S 13105 
Dendrocnide schlechteri (Winkler Chew; T 14067 
Dendrocnide ternatensis (Mi q.) Chew; T 13803 
Flatostema cf. beccarii Schroeter; 7 14055 
Elatostema macrophyllum Brongn,; T 14274 
Elatostema ne Warburg, or aff; 7 
13676, 1389 
Elatostema sp., "aft macrophyllum Brongn.; T 
414, 14053 


a 


ee sp.; | 14258 

Laportea decumana (Roxb.) Wedd,; T 13698 

Pee cf. capitellata (Poir.) Chew; T 13841, 
/ 


4121 
othocnide repanda (Blume) Blume; T 14073, 
14083 
Pipturus argenteus (Forst.f) Wedd; 1 13630, 13779, 
94 
lai act hirta (Blume) Hassk,; T 14130, 14267 


aff. (Forst.) Wedd.; 1 14190 
nee pibeeen. (Blume) Blume; T 14031 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


VERBENACEAE 
Callicarpa cumingiana aah i 7 13963 
ee longifolia Lamk, TT 

aut K.Schum,; T 


13425, 13653 

wae ana cayennensis (Rich.) M. Vahl; T 

08, 14034 

Te ssmenniodendro bogoriense bal T&S 
13114, KJ. White in NGF 1 , aet, J. 
on in NGF 1034 o 

Vitex cofassus Reinw. ex Blume; T 14086 

VIOLACEAE 

Rinorea pen (Korth.) O.K; T&S 13063, T 13554, 
137 

VITACEAE 

Cayratia geniculata (Blume) Gagn.; T 13559 

— Tene (Thunb.) Gagn.; T&S 13077, T 


a ee ana DC.; KJ. White in NGF 10317 
Tetrastigma lauterbachianum Gilg; T 13686, 14223 


WINTERACEAE 
Zygogynum sp.; 1 14195 


MONOCOTS 


AGAVACEAE 
Cordyline fruticosa (L.) A. Chev.; T 13565, 14280 


AMARYLLIDACEAE 

Crinum asiaticum LT 13844 

Proiphys amboinensis (L.) Herbert; T&S 13081, T 
14264 


ARACEAE 

Aglaonema marantifolium Blume; T 13534 

Alocasia aequiloba N.E. Br; T 13595, oo 13892, 
also KJ. White in NGF 10276, - Hay 

Alocasia brancifolia (Schott) A 3573, also 
KJ.White in NGF 10277, and Pullen ran dets. 


A. Hay 
Alocasia cf hollrungii Engl; T 13717 
Alocasia lancifolia Engl.; T 13852, 14097, 14216 
Alocasia lauterbachiana (Engl.) A. Hay; T 13846, 
14713 


a galbra F.M. Bailey; SR, foothill 
for 


Ameypephaluspaccrifl ius (Dennst.) Nicolson; 
SR, road to Roumira 
Colocasia esculenta (L.) sais T 13640 
a cuspidispathum Alderw.; T 13917 
also KJ. White in NGF 10241, det. A. Hay 
ee cf, macrotum Becc.ex Engl.; 7 13705 
Holochlamys beccarii Engl.; T 13478, 13850, 14218 
Homalomena magna A. Hay; T 13849 
Homalomena cf.magna A. Hay; T 13402 


othos Bale ie > ae T 13675 
roto rumphii S 

haphidophora hoe nen T 13879, 14007 
maphiopha versteegil Engl.& Krause; T 13701, 
KJ. White in ee aoe 0, det. D. Nicolson 
Bhai ee Sp ls 
Schismatoglottis sp. A; 1340) 13876, 14103 
Schi ae a sp.B; 7 1363 
gen | White in ceo 10246 


ARECACEAE 

Areca catechu L.; SR, cultivated 

Areca macrocalyx Zipp. ex Blume; T&S 13056, T 
13638 


Areca cf. macrocalyx Zipp.ex Blume; T 13464 

Brassiophoenix schumannii (Becc.) Essig; 7 13513 

Calamus hollrungii Becc.; SR, infrequent near ex- 
pedition Camps 2 and 3 

Calamus humboldtianus Becc.; T 13512-B 

Calamus schlechterianus Becc.; T Ee 

Calyptrocalyx albertisianus Becc.; T 1364 

la ie hollrungii gee ue 1305 : 713401 

hroughout project 


area 
Eon nucifera L.; SR, eae ee 


tor (Be 


GAle/3522 
a fees cfmicrospadi Bee) Bure 1382 


Licuala cf. lauterbachii Dammer & K.Schum.,; T&S 
13171 

Metroxylon sagu Rottb.; SR, common in alluvial 
f 


ores 
Orania macropetala Laut.& K.Schum.; T&S 13057, 
T 13643 
Ptychococcus sp., ‘elatus-paradoxus group’ (cf. 
Essig 1977:19); T 14252 


COMMELINACEA 
oe cane Hassk.; T 13578, 


ilema vitiense Seem.; T 138 
Anetiema Sp., aff. ?humil fee T 13860 
Aaa sp.; 7 13530, 14082 
cf hasskarl i Rao; T 13408, 13621 
ce thyrsiflora (Blume) Steud.; 7 13618 
genus indet., but probably Floscopa scandens 
Lour; SR, Guam R. 


COSTACEAE 

Costus speciosus (Koen.) J. Smith; 7 13702, 14095 
Tapeinochilos eer K.Schum.; 7 go 
Tapeinochilos recurvatum K.Schum.; 7 13700 
Tapeinochilos sp. nov.; 1 13743 


CYPERACEAE 
Cyperus diffusus Vahl var. diffusus; T 13870 


38 


Cyperus kyllingia Endl. T 13881, 14141 
Fimbristylis dichotoma (L.) Vahl ssp. dichotoma; T 
038 


Mapania macrocephala (Gaud.) K. Schum. ssp. 
macrocephala; T 13750, also Pullen 1095, det. 
D. Simpson 

Scleria polycarpa Boeck.; T 13719, also Mills s.n. 
(March 20, 1961) 

DIOSCOREACEAE 

Dioscorea esculenta (Lour.) Burk. SR, cultivated 
_ area 


DRACAENACEAE 
eee cps ia Roxb, 7 13555 
FLAGELLARIACEA 
ipl indica L,; a Guam River 
HELICONIACEAE 

Heliconia ae W.J. Kress; 1&% 13094, 1 13400 
eee. 
C itulata (Lour,) Kuntze; 7 13628 


LILIACEAE 
Dianella ensifolia (L.) DC; T 13442 


MARANTACEAE 

Cominsia gigantea (Scheff) K.Schum, T 13579 
Cominsia cf. minor Valeton; T 1384 

— cannaeformis (Forst.f.) K.Schum.; T 13473, 


ae cf. macrocephalum K.Schum,; T 13468, 
13496 


Phrynium pedunculatum Warburg, or aff; T&S 
13053-A,T 13568, 13878, 13988, 14209 
Phrynium sp. T 13443, 14197 


MUSACEAE 

Musa banksii Fv.M.; T 13642, 14004 

Musa schizocarpa Simmonds; SR, scattered 
throughout area 


ORCHIDACEAE 

(dets. by N.H.S. Howcroft unless otherwise indi- 
cated 

ulbophyllum es Schltr, 
Micromonanthe); T 

Bulbophyllum sp.; TI 36nd coll.) 

Blume; T 14057, 


(section 


Corymborki TA VERA 
140/77 
Dendrobium macrophyllum A. Rich,; TT 3825 


Grammatophyllum papua 

Habenaria chloroleuca Schltr; 7 13525 

— - ongifolia Blume, sl; 7 13527, det. L. 
Jus 


Liparis cay ee bon Reichb f; T 13457, 14094 
Nervilia sp.; KJ. White in NGF 10240,det.A.Dockrill 


SIDA 19(1) 


Oeceoclades pulchra (Thouars) Cribb & Cements; 
T 14075 

Peristylus ?papuana JJ.Sm.;1 13927 

Pholidota imbricata Hooker; T 14008 
Plocoglottis cf. moluccana Schitr; T 13908, 14066 

ted ottis plicata oa ue ssp. puberula N.H.S. 

owcroft; T 13403 

aa disticha oe 7: en 14168 

Vrydagzynea cf. rivularis Schltr; T 13886 

pea aware Be (Fv.M.) Cribb; T&S 13051 

genus indet 


PANDA 
hejeeto a = 13407, 13778, currently under 
by K.-L. Huyn 
ee angiensis Kaneh., or aff; T 13847 
Pandanus cf. cernuifolius Merr. & Perry, ‘beccarii- 
cernuifolius group (cf. Merr.& Perry 1939: 180); 


<< 


T 14222 
Pandanus lustrorum Stone, or aff.; 7 14224 


POA 
Ap adam mutica L.; T&S 13076, also KJ. White in NGF 
303 


ea compressus (Swartz) Beauv.; 7 14145 
a microcephala (Pilger) Holttum;T 13558, 
13622 


Brachiaria mutica (Forsk.) Stapf; SR, Guam R. 
bridge 
Centotheca latifolia (Osb.) Trin. T 13532 
Chrysopogon aci ie SR,Roumirap 
Coix lachryma-jobi LT 
Cyrtococcum accrescens fae Stapf; T 13896 
Dactyloctenium aegyptium (L.) oo T 14023 
Fleusine indica (L.) aoe 714 
Bailey) ae T 14304 
ee banksii R. He ny det. kK. Damas 
Leptaspis urceolata (Roxb) R.Br; T 13467, 13533 
Selo splay (L.) PR Beauv, 7 13955 
nicu osum Roxb.; 1 13548 
Penta, ene Schumach.,; T 13780 
Pogonatherum paniceum (Lamk) Da T 14152 
709 


g 
= 
Y 
= 
C 
a 
u& 


bth 


Benth.; 7 13 
altata Lf,; 


olytoca 

Rottboellia eee 

h ffici “SR, cultivated 
Saccharum robustum Srandes : Jeswiet ex 

Grassl:SR thickets alon 
Beers i (Blanco) vee Pullen | 17, 

et. R. Holtt 
Setaria pa Imifolia Koenig) Stapf; 7 13957 
Sorghum nitidum (Vahl) Pers.; KJ. White in NGF 

10304 


Sorghum propinquum (Kunth) Hitchc.; 1&S 13074 
Themeda arguens (L.) Hack.; T&S 13075 
SMILACACEAE 

Smilax . australis R.Br; 7 13569 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


ZINGIBERACEAE 
rcuma cf. australasica Hooker f.,‘petiolata 
group (cf. Valeton 1918: 10); T 13703 
Etlingera dekockii (Valeton) R.M. Smith; 7 713535 
Etlingera sp. ?nov. (series Polyanthae); T 13985 
Hornstedtia scottiana (Fv.M.) K.Schum.,; 7 142177 


APPEND 


39 


Pleuranthodium . erate (Valeton) R.M. 
Smith; T 13406, 

Riedelia ee acon T 13853, 14128, 
14213 

Riedelia longifolia Valeton; T 13771 

Riedelia macrantha K.Schum.,; T 13816 

Riedelia sp.; T 14269 


IX 2 


THE SCIENTIFIC EQUIVALENTS 
OF MAIAN (TOKPLES) PLANT NAMES 


Traditional names are listed with the corresponding binomials determined from the 
vouchers. Spellings are phonetic. Several taxa are apparently represented by orthographic 
variants in the Maian language (cf.compilation following main listing).This situation may 
be due to the lack of a written grammar for the tokples, and the resulting absence of a 


medium for its standardization. 


Maian Name Scientific Binomial Family 
ah-maap Micromelum minutum (Forst.f.) Wight & Rutaceae 
Walker-Ar nott 
ah-mo-rap Piper cf.can Piperaceae 
ah-nah-sahr Decespermum eae (Roxb.) AJ.Scott var. Myrtaceae 
ah-rap a ee ve a-jobi L Poaceae 
ai-ke-kav Erec Hae Sees a (Wolf) DC. Asterac 
akakarap-nevermbup — Asplenium subemarginatum Rosenst. Aspleniaceae 
ambo-dera-namb Zizi ite Gane Laut Rhamnacea 
mbu Dendrocnide ternatensis (Miq.) Chew Urticaceae 
amora-kamorap BIDE decumanum Buia) Es Piperaceae 
amungcurcuri p (Laut. & K. Schum.) Apocynaceae 
Fosb. & Sach. 
anganange-woganamb _ Tetracera nordtiana Fv. Dilleniaceae 
angkuma nee Ae leto ex Bakh. Ebenaceae 
ngo-leb omalom Hay Araceae 
arab pe heter sok oxa Cate .& Lau Leeacea 
aramitap Steganthera hospitans (Becc.) ae & Hatus. Monimiaceae 
arap Pimel sce amboinicum Hassk. Euphorbiaceae 
arep ammatae Sp. Rubiacea 
aru tris Teijsm. & Binn. Nyctaginaceae 
atep oe tis S eae 
ve-na Steganthera dentata (Valeton) a & Hatus. Monimiaceae 
babagalum pal acarpus ae ae He Rubiacea 
babagalum tria mayana Rubiaceae 
babagalum Pychatno ee ae (ae & K.Schum.) Rubiaceae 
babagalum San sp., aff. micralabastra (Laut. & K. Schum.) Rubiaceae 
Valeton 
badidir Elattostachys obliquinervis Radlk. Sapindaceae 
bago-bagot Dianella ensifolia (L.) Liliaceae 
bailalum St th hi (Warburg) Perkins Monimiaceae 
bailarum ndet. Orchidacea 
bal-sivar Merremia peltata (L.) Merr. Convolvulaceae 


bimur-nganam 
o-rap 
boa-boak 
boagalum 
bodog 
bogang-dap 
boge-namb 
bogo-namb 


buasum 
buburat 
buko-bukop 
buku 


uku 
buku-wabado 
bulubul 


da-da-da-dag 


daveh-veh 
dawab 
dawaba-sivar 
dide-lolol 
diga 

dodor 


duaram 
duat-murukun 
ese- 
esg-ese-ya 

eve 

eve 

ga-tsurup 


abin 
gaga-bumer 
gaga-bumer 


peleene nidus ie var. nidus 
ene ean Benth. 


Intrirtic armeNntel 


intsia bijuga Cole 
Solanum torvum 
oe 


| t 


oo 
Airosperma psychotrioi 


eet ss ial Gi gq 
oode 


IPECTET!S 


ver nga iirabell ata L. var. papuana Valeton 
Pararistolochia schl id (Laut.) M.J. Parsons 
Th Bor 


espesia fissicalyx Bors 
Codiaeum s 
Archidondr, 


n belli nm 


ms 
Eq HSeTLLM FOAMOSISSIMUM Desf. ssp. debile 


(Vauc 


Aglaia ood a Harms 
k 


Aglaia saxonii Takeuc 


maa jepidopetal a Harms 
a Miq.v 


(Elmer) Comer 


aut. 
Syzygium sp., aff. goniopterum (Diels) Merr.& Perry Myrtaceae 
Asplenium cf amboinense Willd Aspleni 
ries obtusa J. Smith 
Liparis condyl oy bon Reichb f. 
Gouania cf. javanica Mi 
Horsfieldia subtilis (Miq.) Warburg var. subtilis 
Ficus dammaropsi Ss Diels var. obtusa Corner 


Begonia f se 
ia nee Coo 
Alangium villosum (Blume) 


=> 


Termina 


Rhyticarvijm 


Garcinia dulcis (Roxb.) Kurz 
Entada phaseoloides (L.) Merr 
Abrus nade lus Thwaites ssp. pulchellus 


Ficus wa oxb. 
Satine aruense (\ 


hanno cf.macrotum pice ex Engl. 
Aglaia agglomerata Merr. & Perry, or aff. 
ae 


deaaimpertle K. . 
Phyllanthus rubriflor 


J.Sm. 
Inocarpus fagifer A es eee ) Fosb. 
Inocarpus (close to) ‘rubidus’ morphotype fide 
Ver 


oe phaeochlamys (Laut. & K. Schum.) 


ifolia (Sc ao) 


P| 
Bag Wore Christ 
Pothos rumphii Schott 


Rhaphidophora versteegii Engler & Krause 


tah 
uron tuberculatum (C 


SIDA 19(1) 


Urticaceae 


Combretaceae 
Rubiaceae 
Aristolochiaceae 
Malvaceae 
Euphorbiaceae 
Mi 

Equisetaceae 


Meliaceae 


Clusiaceae 


Fabaceae 


Rubiaceae 


Thelypteridaceae 
Pteridaceae 
Araceae 

Araceae 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


gaira-malapta 


gilagal 

giliba 
giligelum 
giligelum-sivar 
go-idi 

guragor 


kasiwar-gili-gilib 
kasiwar-gili-giliba 


kasuar dadi 


kasuar-mudu-mado 
kawa 


ker-ker-kanamb 
ker-ker-kanamb 
kibai 


kibaip 
kibi-kibale 


Barringtonia calyptrocalyx K.Schum. var. mollis 
aut, 


Laportea cece ee ) Wedd. 
puana W.J. Kress 


ottis s 2 
ania macrocephala (Gaud.) K.Schum. ssp. 


pitulat 


PCOS 
Cur 


Kuntze 


(Lour) 
Cu ee cf. ausiaae) ica Hooker 


Psychotria (po 
ii FV.M 


a VM. 
oo falcataria 
falcatari 


ossibly new vining sp.) 


(L.) Nielsen cf. ssp. 


rene sparsiflorum Mabberley 


Piper mestonii F.M. Baile 
Ptyssiglottis pubisepala 


(Lindau) B. Hansen 


vee sp., aff. tenuipedicellata Baker f. 


Tropiai _ di Micha Schltr. 
Ge 


a (K.Schum 


.& Laut.) Valeton 


iostom ma sete R.& - Forst. 


g 


Metre sou 


y 

nthus sen ca la (Lindau) Bremek. 
oa is hetrocla (Presl) C 
Poe ne fruticosa (L.) A 
Brass 


f.) Merr 
lisa ecu aS f.) K.Schum. 
ium p m (La 


t.& K.Schum.) 


OLR 
= 
a3 2 
< 


en a i (Becc.) Essig 
Br. 


ipomoea congesta R 


Phryn ium pe | f 
Phryniu 
Piper caninum Bl 


Warburg, or aff. 


me 
Macaranga fallacina Pax & Hoffm. 


| 


Dp. qd 
GLY! Ufii CUUIC 


Schizostachyum lim 


Reir VV, 


(Blanco) Merr. 


Etlingera dekockii (Valeton) R.M. Smith 
eae os Blume ssp. puberula 


Le ia’sp., ee Val., 
.facies’ 


or sphaerocarpa 


Cerbera flo eee K.Schum. 
K. 


Schum. 


Amischotolype mollissima Hassk. 


omalom 


— 


ena i magna A. Hay 


Xora sp., section Hypsophyllum 


Barringtoniaceae 


Urticaceae 
Heliconiaceae 
Araceae 
Cyperaceae 


Hypoxidaceae 
Zingiberaceae 
biaceae 


Mimosaceae 


Meliaceae 


Melastomataceae 
Rubiaceae 
Orchidaceae 
Loganiaceae 
Thymelaeaceae 


Myrtaceae 


Acanthac 
Lo ee 
Agavaceae 
Arecaceae 
Convolvulaceae 


Euphorbiaceae 
Flacourtiaceae 
Poaceae 
Zingiberaceae 
Orchidaceae 


Rubiaceae 


Apocynaceae 


Rubiaceae 


42 


kibi-kibale 

kibi-kibale 

kidi-kidi 

kivi-kiva 

kobos-susul 

kobou-susul 
ita 


kolaiv-nganam 
komekelak 
korang-korang 
kututal 
kuvu-kuv 
kwarikioari 
lago-lagod 
lala-lala 

lam 


lasa-lasa 


lasa-lasa 


mamba-mambap 
mamba-mambap 


man-duroop 
man-gab 


man-trep 


Lep isanthes senegalensis (Poir.) Leenh. 
ern coccinea (Gaud.) F.v.M. 

f Pleuranthodium Sp., ?trichocalyx (Valeton) 
R.M. Smith 


Geniostoma ri Ipestre JR. & G Forst 


Myristica subalulata Miq. var. subalulata 
Horsfieldia subtil is (Miq.) Warburg var. subtilis 
Sleumer 

Glochi dion granulare Airy Shaw 

Petalolophus sp., aff. megalopus K.Schum. 


Sterculia schumanniana (Laut.) Mildbr. 


Pseuduvaria sp., aff. ?versteegil (Diels) Merr. 
Sphaerostephanos arfakianus (Baker) Holttum 
a: polystachya (Wall.) R.N. Parker 
Valeton 

Flatostema novoguineense ee or aff. 


immanemoan ro 


~ 


a enl ium phyllitidus Don mh ‘malesicum 
ttum 


sane genicu ata (Blume) Gagn. 

Euphorbia hirt 

aaa ium cf. a Swartz,‘affine-cuneatum 

rou 

Bolbitis quoyana (Gaud.) Ching 

Lindsaea tenuifolia Blume 

Microsorum membranifolium (R. Br.) Ching 

Pleocnemia macrodonta (Fée) Holttum 

seteeaipus sll ae Merr. & Perry 
K.Schum. 

Popowi wia sp., aff. pisocarpa (Blume) End. 

Harpullia crustacea Radlk. 

Callipteris prolifera (Lamk) Bory 

Lomagramma cf. sinuata C. Chr. 

Stenochlaena milnei Underwood 

Trichosanthes sp.,‘longiflora-bracteata group’ 

Holochlamys beccarii iEngl. 

Riedelia liligula Valeton 
a ia macrantha K.Schum. 

Osmelia philippina (Turcz.) Benth 

Cleistanthus sp., aff. ?papuanus (Laut.) Jabl. 

Exythrospermum avila (Becc.) Becc. 

chum. & Laut. 


Ficus odoardi K 

Grammatophyllum papuanum J.J.Sm. 

Pyrrosia eee seas Morton 

Garcinia maluensis Laut. 

Syzygium trach a nthum (Diels) Merr. & Perry 

Osmoxylon sessiliflorum (Laut.) Philipson 

eee (closest to) sessiliflorum (Laut. 
ilips 

Docuiont cf. moluccana Schitr. 

Alocasia lancifolia Engl. 

Anacolosa cf. papuana Schellenb. 


LS 


SIDA 19(1) 


Sapindaceae 
Thymelaeaceae 
Zingiberaceae 


Loganiaceae 
Myristicaceae 


n 
Thelypteridaceae 
Meliaceae 

ubiaceae 
Urticaceae 


Aspleniaceae 
Vitaceae 
Euphorbiaceae 


Aspleniaceae 


Peele Anis 


Lomariopsidaceae 
Blechnaceae 
Cucurbitaceae 


Araliaceae 


Orchidaceae 
Araceae 
Olacaceae 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


manda-peb 
mandurup 
mane-kav 
mane-mane-kav 
mane-mane-kav 


mansu-borobor 


marap 
mariap-tobitobi 
mariap-tobitobi 


maruruma 
masa-wun-bandep 
mavanda-ngamb 
mekukum 
migim 

moimoit 
mondi-mi 


monia-kiui-kiva 
monia-nasag 
mor 
mora-morava 
mouko 


moyab- pooh 


muania-kivikiva 
muara-muarav 
muat-upot-ugarum 


muat-upot-ugarum 
mugum 


muonia-kivikiva 
musus 

mutu-ngomb 
nanggu-nanggu-nam 


ngabu-ngabu 
nganam idir idir 


ngapar-nganam 
ngaparu-pot 


Selaginella sp., aff. ?hieronymiana v.AV.R. 
indet. 
Areca cf.macrocalyx Zipp.ex Blume 
Lasianthus chlorocarpus K.Schum. 
Medusanthera laxiflora (Miers) Howard 
ieee MaHOGAVE a ex Blume 

ae E. Br. 


a eceias vterboctan ae JA 
(Diels) vee & Perry 
ygium longipes Merr. & une 
ee Miva inense _ DC. 
Piper pullibaccum Treleas 


Steganthera dentata (Valeton) Kaneh. & Hatus. 


Ficus bernaysii King 
Ficus conocephalifolia Ridl. 
Wenzelia dolichophylla 
Cyperus diffusus Vahl var. diffusus 
Pseudobotrys ae sl 

Hoya pottsii FM.B 
Cyrtandra sp., Sone oo phon 
oeHe polycarpa Boe 

rpa 1A. Gray) Leenh. 


2 


Averrhoa bilimbi L 
Cryptocarya laevigata Blume 
{nocarpus fagifer (Parkinson) Fosb. 
Piper ae Blume 
vm icrocos Sp. ?nov. 
Comin sia gigantea (Scheffer) K.Schum. 
Phrynium cf.macrocephalum K.Schum. 
Phrynium sp., aff. macrocephalum K.Schum. 
Alocasia cf. hollrungii Engl 
Piper decumanum (Rumph),) L. 
Hedyotis sp., cf. H. auricularia L., or 
H. ees DC. 
Justic 
ee spectabi ilis (Harms) Philipson 
Melodinus cf. acutus (Markgraf) Markgraf 
aie eat i. rck 

d t.& K.Schum. 
Graptophyi um m pictum ‘i 5 Grif 

L. 


uiloba 


jen s mollior B 


a llanthes aL Hemigraphis primulifolia (Nees) 


F.Vill. facies 
Amaracarpus grandifolius Valeton 
Lunasia amara Blanco var.amara 
Syzygium : amplum Hartley & Perry 
Syzygium 


eae conspersipunctatum Merr. & Perry 


Psychotria leptothyrsa Miq. var. leptothyrsa 


Medinilla triplinervia Cogn.,'mufoso-triplinervia 


group 
Amaracarpus sp., aff. longifolius Valeton 
Sphaerostephanos pilososquamatus (v.AV.R.) 
Holttum 


(Laut. & K. Schum.) Tanaka 


Fen 
idac 


Piperaceae 
Rubiaceae 


Acanthaceae 


Acanthaceae 


Rubiaceae 


U 
Melastomataceae 


Rubiaceae 
Thelypteridaceae 


44 


NQauNn-Nasag 


ngayom-nasag 
ngoku 
ngumusinam 


ninara- NauMuU 


pah-tooey 
pake-koal 
pale-palel 


palisar 
pat dagol dagol 


sagag-gosmun 
sagag-mum 


sagag-u- ie umun 


sagag-ug 
5 


Sa 

sakamb 

sakam 
sakas-sakamb 
sakomb 

samangi manggib 
sana ngamb 


sang-guab 


sauga-sivar 
Sauia 
say-ri-keep 
say-veb 


Aglaia sapindina (Fv.M.) Harms 

Helicia affinis Sleumer 

Geniostoma rupestre J.R. & G. Forst. 
Psychotria membranifolia Bartl.ex DC. 
Psychotria cf. ee Bartl.ex DC. 
Ficus odoardi Ki 

Pueraria pucherina ee 
Koorders- Schum 


tha 


Gilg, or aff. 
eee eomee (Kort th.) OK. 
Rhaphidophora eee Schott 
eerie oleae 

Psy eT ne 
Syzygil m hy ie philu im (Laut. & K.Schum.) 
Mert i. Perry 


pa Burck 
Ficus congesta Roxb. 
Codiaeum variegatum (L.) Blume 
Dysoxylum sparsiflorum Mabberley 


m) 
Mackinlaya ae ica oars Philipson 
Selaginella velu a a sati 


Ocimum gra 
Harpullia i an 
Cryptocarya laevigata Blume 
egonia papuana Warburg 
Begonia cf. papuana Warburg 
goni Warburg 
Calycosia mamosei lakeuchi 
Cyrtandra sp., section Centrosiphon 
Flatostema sp., aff. macrophyllum Brongn. 
Cyrtandra sp., section Centrosiphon 


wx 


Strobilanthes s.|.(Hemigraphis sp. 
Myristica lancifolia Poir. ne ere 
Myristica tristis Warburg, o 
Amaracarpus Sp., aff. a Valeton 
Dicliptera papuana Warburg 
Eclipta prostrata (L.) L. 
indet. 
Tabernaemontana orientalis R. Br. 
Pneumatopteris sp., aff. keysseriana (Rosenst.) 
olttum 
Goniothalamus imbricatus Scheffer 
Goniothalamus cf. imbricatus Scheffer 
Myristica lancifolia Poir. ssp. lancifolia 
Aceraiionn vedere nai) Schltr 
Opilia ar a Roxb. 
ne sarmentosum Roxb. 


Aneilema $0: aff, ?humile Warburg 


SIDA 19(1) 


Meliaceae 


Fabaceae 
Flacourtiaceae 
Violaceae 
Araceae 
Se ae 


Rubiac 
ee 


Rubiaceae 
Moraceae 
Euphorbiaceae 
Meliaceae 
Thelypteridaceae 
Polygalaceae 
Acanthaceae 
raliaceae 
Selaginellaceae 


Asteraceae 
indet. 


Thelypteridaceae 


Annonaceae 


eae 
Commelinaceae 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


awag 
sigirpa-nganam 
SINGI-siINggip 
singi-singgip 
singi-singgip 
SINGI-SINGGip 


Eom SeEAGIED ae Leenh. 
Cyr Sta 


AN ixis ail ) R.N. Parker 


ae cf. pohlianus eee 
Calycacanthus magnusianum K Schum. 
Bey nem 2 Roxb. 


Ae laia eget Mert. - Perry 


aus oe rostris Teijsm. & Binn. 
. sple cf.a amboinense Willd. 


Polytocha magpie aie 
Sch 


va raalalalalil 


Cephaloma trovi ie 
Lindsaea obtusa J. Smith 

Lindsaea cf. obtusa J. Smith 
Lindsaea tenuifolia ae 


Amaracarpus Sp., aff.'attenuatus 
Amaracarpus sp., aff. ey Valeton 
Goniothalamus cf. imbricatus Scheffer 


Omphalea cueenslandie F.M. Bailey 


ee 


Elaeocarpacea 


ty eases ee 


Lindsaea group 


Annonaceae 
Euphorbiaceae 


sob-barewa Psychotria sp.n Rubiaceae 
sob-takevam ee ae tis cf. quoyana wa oe on Lomariopsidaceae 
sopi-sebip abaceae 
sretu-ngomb N itrod on & K.Schum.) Apocynaceae 
Fosb. & Sach 

staylki Calopogonium mucunoides D ace 
sumbu-wadab Elatostema novoguineense Weibits or aff. Urticaceae 

mure Strobilanthes s.|.(Hemigraphis sp.) canthaceae 
ta-kup Ficus pungens Reinw. ex ne oracea 
ta-wop Cominsia . ss ONnOr Valeton Marantaceae 
tagle Antia .Schum. race 
tagle hoe ps Burm Moraceae 
tah-bop hrynium pedun ulatum Warburg, or aff. Marantaceae 

i-na eae dubius ae Amaranthaceae 
take-takel Aglaia saxonii Takeuch Meliaceae 
take-takepa Arytera sp., aff. litoralis plumevitorts complex’ Sapindaceae 
takevam Tectaria menyanthides (Presl) Copeland Tectaria group 
taleb Cupaniopsis macropetala Ra Sapindaceae 
tatar-ulalat Podocarpus cf.rumphii Blum Podocarpaceae 
televa-nganam Pseuderanthemum 5$p., cf. vvariabile group’sensu. Acanthaceae 

Barker 
ga rania macropetala Laut.& K.Schum. Arecaceae 

timaiyag-nav ou ape yx a lberti lee Becc. Arecaceae 
timber-digeep P Kaneh., or aff. Pandanaceae 
titirigi Licuala beccariana ae Arecaceae 


titirigi Licuala cf. beccariana Furtado Arecaceae 
tuar Callicarpa longifolia Lamk Verbenaceae 


tuar 
tukum-avang 
tuturat 


uaia 

uduat-murukun 

ugag 

ulengkuduv 

umbol-lap 

umbol-menyap 

undu-beb 

urat 

urat 

urem dagur 

usiman-kekerup 
-tib 


voa-baga-bungam 
vogerdak 


vogo-gon-dab 
vokor 


Wana-barewa 
wanam-barewa 
wanclap 


wandarumep 
ang-gep 


wanga udial 


warubu-sopasop 
warubu-taleba 


SIDA 19(1) 


Mycetia javanica (Blume) Reinw. ex Korth. Rubiaceae 
Medinilla sp., aff. tenuipedicellata Baker f. Melastomataceae 
Stachytarpheta cayennensis (Rich.) M.Vahl Verbenaceae 
lapeingemon ane aurantiaca Gaud. Apocynaceae 
nciq neni iloha N. E. Br. 
Mussaenda cylindrocarpa Burck Rubiaceae 
Calyptrocalyx ee fi ti Arecaceae 
Sabia pauciflora Blum Sabiaceae 
Tabernae onan ere Gaud. Apocynaceae 
Peristylus ee J.Sm Orchidaceae 
Cai my sp. Rubiaceae 
nh. Loganiaceae 
Hibiscus elliptici ifolius Borss. Malvaceae 
Diospyros alee Mere ex Bakh. benaceae 
Diplora d'urvillaei (Bory) C Aspleniaceae 
Ase rasleie! ype mollissima ieee Commelinaceae 
ndrobium macrophyllum ee Orchidace 
a ipteris pro ie (Lamk) B Athyriac 
Pleocnemia macrodonta (Fée) Holttum Tectaria group 
peel: sogerensis (Gepp) Holttum Thelypteridaceae 
Leucosyke cf. capitellata (Poir.) Chew Urticaceae 


Cryptocarya laevigata eu 
ieee age 


Haplostichanthus ae (Scheffer) Ann 

van Heusden 

Pseuduvaria An ace 

SUED Anam a macocea Lau acardiaceae 
L. var. ee or (Blume) Kalkman Rosaceae 

Gipaniopsi macropetala Radlk. apindaceae 


Leptaspis urceolata (Roxb.) R. Br. Poaceae 


Ficus subulata Blume ac 

uliflora (K. Schum. & Laut.) Valeton — Rubiaceae 
Antidesma katikii Airy Shaw eee 
Cucurbita ucurbitaceae 
Psychotria sp.nov ee 
Versteegia folia Valet Rubiaceae 
Melicope sp., - . burttiana one or Rutaceae 
M. grandifolia 
Cyrtosperma Cu ee hum Alder 
Saprosma subrepandum (K.Schum. ads Rubiaceae 

Valet 

ee alyx sp. ?nO Annonaceae 
Tarenna gulcheriana (K. Schum. ) Valeton Rubiaceae 


Lepisan nthes sene galensis (P oir.) Leenh 

Breynia cernua (Poir.) Muell. Arg. 

Crinum asiaticum 

AL cf. reti ae (Forst.) Kaulf. 

Microsorum linguiforme (Mett.) Copel. 

Poikilospermum amboinense Zipp. ex Miq. 
C. 


Ag laomorpha drynarioides (Hooker) Roos Polypodiaceae 


Glochidion chondrocarpum Airy Shaw, or aff. Euphorbiaceae 
Dysoxylum brassii Merr. & Perr Meliaceae 
Dysoxylum pettigrewianum F.M. Bailey Meliaceae 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 47 


wasagep Psychotria sp., aff. micralabastra (Laut. & K. Rubiaceae 
.) Valeton 

wasimi Diplora d'urvill aei (Bory) C. Chr Aspleniaceae 
wat-uduat Maniltoa schefferi K.Schum. & Hollrung Caesalpiniaceae 
wat-ukauel Smilax cf. australis R. Br. Smilacaceae 
wat-urimap Cyathocalyx papuanus Diels, or aff. Annonaceae 
wat-virimav Arytera sp., aff. litoralis Blume, “ioral complex’ Sapindaceae 
wata-katok Freycinetia sp. Pandanaceae 
wato-karok Pothos papuanus Becc. ex 7 a 

watulam Psychotria lex DC. Rubiac 
wedem-lalaut Cephalomanes atroviren a res s Hymenophyaceae 
wegem-lalaut pupelzia et eigualios a (Forst.f.) Trevisan Lycopodiac 
widasag iry Shaw Sic 
wingam oe Gael Blume Anacardiaceae 
wo-mamb Aporosa cf. papuana Pax & Hoffm. Euphorbiaceae 
wo-roon-botop Pipturus argenteus (Forst. f) Wedd. Urticaceae 
wo-sarep Psychotria mayana Takeuchi Rubiaceae 
wo-sayep Calycosia mamosei Takeuchi Rubiaceae 
wo-wop Callipteris spinulosa (Blume) J.Smith Athyriaceae 
wonkibung Cry tocarya weinlandii K.Schum. Lauraceae 
wuka-wukap aeane kyllingia Endl. C 
wumbu-ngam Endospermum moluccanum (Teijsm. & Binn.) Kurz Euphorbiaceae 
wung-wanam Artocarpus communis J.R. & G. Forst. Moraceae 
wungo-bunyam Wedelia biflora (L.) DC. Asteraceae 
yag-ikikav Blumea arfakiana Martelli Asteraceae 
yag-mara-marav oe pseueciaieiail Warburg Begoniaceae 
yag-sawea lerodend calyx Laut. & K.Schum Verbenaceae 
yag-tauita ee gtilcheriana (K. Schum.) Valeton Rubiaceae 
yavera-ukum Fagraea ceilanica Thunb. Loganiaceae 


Taxa represented by Maian orthographic variants: Aceratium ledermannii Schltr, sangu-mutup 
sigirpa-nganam; Aglaia lepidopetala Harms, buku, buku-wabado; Aglaia saxonii Takeuchi, buku, take- 
takel;Alocasia aequiloba N.E. ae Maas gap, moyab-pooh, uduat-murukun; Amaracarpus grandifolius 
Valeton, babagalum ; acarpus sp., aff. longifolius Valeton, ngapar-nganam, sakamb, sivar- 
viav; Amischotol lype mollissima feeat keiki, vap; Aphanamixis agente (Wall.) R.N. Parker, korang- 
korang, seger; Arytera sp., aff. litoralis Blume, take-takepa, wat-virimav; Averrhoa bilimbi L., monia- 
kiui-kiva, sepi-sepib; Begonia papuana Warburg, damu- se rubu-rubu; Begonia pseudolateralis 
Warburg, rubu-rubu, yag-mara-marav; Callipteris prolifera (Lam.) Bory, ma-bairap, yee Calycosia 
mamosei Takeuchi, rubu-rubu, wo-sayep; Cephalomanes atrovirens Pres|,singi singgip,W m-lal 
Cryptocarya laevigata Blume, monia- nasag, mets oem -nasaq, vinisa; Cu ] Radlk., 
taleba, wadi h., angkumamb, usiman- kekerup;D Caer aei 
(Bon) C. Chr, vah- tib, wasimi; Dysoxylum sparsiflorum eee, i- “Pap, pale Ficus odoardi King, 
maboramb, ngoku; Garcinia maluensis Laut., da-da-da-dag, ma a rupestre JR. & G. 
Forst.,ivang-glu, kivi-kiva, ngawar-dodol:G lochidion granulare Airy shaw, koita, widasag; Holochlamys 
beccarii Engl., kolaiv, ma-nem-gab; Horsfieldia subtilis (Mig.) Warb. var. subtilis, dalulup, kobou-susul; 
Inocarpus fagifer (Parkinson) Fosberg, eve, mor; Lepisanthes senegalensis rou) Leenh., kibi-kibale, se- 
bip, wange-warap; Lindsaea obtusa J. Smith, dago-dagol, lhe sineg)p Lindsaea tenuifolia Blume, 
lasa-lasa, singi-singgip; Mussaenda cylindrocarpa Burck, mum-nganam, pah-tooey, ugag; Myristica 
lancifolia Poir.subsp./ancifolia, sagua, sang-guab; Phaleria coccinea (Gaud.) F.v.M. a eee! 
Phrynium pedunculatum Warburg, or aff, kamasasak, tah-bop; Phyllanthus rubriflorus J.J. Sm., 
ese-ya, seger-buga; Piper decumanum (Rumph.) L.,amora- kamorap, muara-mMuarav; pes aigenteus 
(Forst. f.) Wedd., bial, wo-roon-botop; Pisonia longirostris Teijsm. & Binn., ar »p; Pleocnemia 
macrodonta (Fée) Holttum, lasa-lasa, vebum; Psychotria See aie aut 2 K ih mumbutakut 
osag-rep; Psychotria mayana Takeuchi, babagalum, wo-sarep; Psychotria membranifolia Bartl. ex DC., 
ngayom-nasag, watulam; Psychotria phaeochlamys (Laut.& K.Schum.) Valeton, babagalum, ga-tsurup; 


48 SIDA 19(1) 


Psychot rid Sp. nov., s0b- barewa, wanambarewa; Rinorea horneri (Korth.) O.K., ninara-umu, sekera- 
& Hatus.,ave-namb,marap; Jabernaemontana aurantiaca 
Gau d, uaia, umbol-menyap; Tarenna gtlcheriana (K. Schum.) Valeton, wange-abab, yag-tauita; 
Terminalia impediens Coode, bogang-dap, danga-namb; Versteegia cauliflora (K. Schum. & Laut.) 
Valeton, idi-muyat, waipa. 


APPENDIX 2A 
THE SCIENTIFIC EQUIVALENTS OF MAIAN (TOKPLES) 
PLANT NAMES (ALPHABETICAL BY GENUS) 


Maian Name 
dide-lolol 
sangu-mutup 
Sigirpa-nganam 


Scientific Binomial 


Abrus pulchellus Thwaites ssp. pulchellus 
Ace iy lod, ny Schltr. 


Aceratitim ledermannii Schlt tr. 


Family 


Fabaceae 
Elaeocarpaceae 


oro-momb ped uu sta Blume Amaranthaceae 
USUS Asteracea 
seger-nanam Agiaia gogiorea er & Perr Meliaceae 
dzam Agl rr. & Perry, or aff. Meliaceae 
warang-gap Aglaia cuspidata C. De Meliaceae 
buku Agl aid lepidopetal a Harms Meliaceae 
buku-wabado Aglaia lepidopetala Harms Meliaceae 
ngaun-nasag Aglaia sapindina (F.v.M.) Harms Meliaceae 
buku Aglaia saxonii Takeuchi Meliaceae 
take-takel nae cc ao eliacea 
Warap-tep ides (Hooker) Roos Polypodiaceae 
boagalum Airosperma psychotrioides Laut.& K.Schum. Rubiaceae 
davaru-guar Ala IgiuM villosum (Blum €) Wa angerin Alangiaceae 
mang-gap Al ia aequiloba NE. Br. Aracea 
moyab-pooh Alocasia aequiloba We Araceae 
uduat-murukun Alocasia dequilo ba Araceae 
gabin a brancifolia (eho A. Hay Araceae 
muania-kivikiva eile Araceae 
man-gab ees lancifoli Araceae 
Alocasia ce (Engl.) A. Hay ac 


mang-gap 
babagalum 
n 


Amaracarpus grandi he \ | iaceae 
sivar-Vviav Amaracarpus sp., aff.‘attenuatus-heteropus group’ Rubiaceae 
ngapar-nganam Amaracarpus sp., aff. longifolus Valeton iaceae 
sakamb Amaracarpus sp., aff. longifolius Valeton Rubiaceae 
sivar-viav Amaracarpus i aff. longifolius Valeton Rubiaceae 
arep Amaracarpu Rubiacea 
tai-namb Amaranthus fee Thell. Amaranthacea 
keiki Amischotolype mollissima Hassk. Commelinaceae 
va Amischotolype mollissima Hassk. Commelinacea 
man-trep Anacolosa cf. a Schellenb. Olacaceae 
ker-ker-kanamb Aneilema vitiens en Commelinaceae 
Say-ve Aneilema sp., aff. humile oe Commelinaceae 
tagle Antiaropsis decipiens K. et cea 
wakup Antidesma katikii Airy Sha Stilaginaceae 

ara-te Antrophyum cf. aan (Forst.) Kaulf. Vittariaceae 
korang-korang Aphanamixis polystachya (Wall.) R.N. Parker Meliac 
er NM a cia ale (Wall.) R.N. Parker Meliaceae 
wo-mamb Aporosa cf. papuana Pax & Hoffm. Euphorbiaceae 
dodor jenennene aruense (Warburg) de Wit Mimosaceae 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


buburat 


wung-wanam 
take-takepa 
wat-virimav 
lasa-lasa 


dago-dagol 
sibi-namb 

baratep 

lago-lagod 
akakarap-nevermbup 
monia oe i-kiva 

sepi-se 

aiebe 

gaira-malapta 


yag-mara-marav 


ulengkuduv 

urat 

koita 

ninara-naumu 
lala-lala 

bee-en 
singi-singgip 
wedem-lalaut 
kasuar-mudu-mado 


yag-sawea 


idendron bellum Harms 
Arisa imperi ialis K.Schum. 
Areca macrocalyx Zipp. ex Blume 
Areca cf.macrocalyx Zipp.ex Blume 
Artocarpus communis J.R. & G.Forst. 


Arytera sp., aff. litoralis Blume, ‘litoralis complex’ 
Arytera sp., aff. litoralis Blume, ‘litoralis complex’ 
sae ium cf. affine Swartz,affine-cuneatum 


gro 
erate cf.amboinense Willd. 
Asplenium cf.amboinense Willd. 
pla ium nidus L. var. nidus 
ple itidus Don ssp. malesicum 


eNHIM phy i 


tum 
Asplenium oe um Rosenst. 
ieee bilimbi L 
flied bilimbi L 
mbusa mi icrocephala eee Holttum 


ae calyptrocaly ar. molli 
Laut. 
Begonia par Warburg 
nia | Warbur g 
Begonia cf.p 


ia cf. papu 
Begonia pseudolateralis Warburg 


ee itis quoya we au hing 
Bolbitis cf. quoyana (Gaud,) Chi 
rassiophoenix schumannii (Becc.) Essig 


— 


Biya cernua (Poir.) Mu - # 


aniaq macro arpa 


ean humboldtianus se 


Callipteris prolifera (Lamk) Bory 

meat prolifera (Lamk) Bory 

Cal Was ris EDI rele a 
SV 


NILIIM IMILICLIN 


alo 
| ycacan nthus magnusianum K. Schum. 
alycosia mamosei Eee 
lycosia mamosei lakeuc 

alyptrocalyx albertisianus oe 
Calyptrocalyx hollrungii Becc. 
Canthium sp. 

Casearia erythrocarpa Sleumer 
recite macrantha Gilg, or aff. 

ratia lata (Blume) Gagn. 


Ca 
Ca 
Ca 
Ca 


— 


Chisocheton pohlianus Harms 
Chisocheton cf. pohlianus Harms 


Sasa a oP all hia oy ) Jabl. 
t.& K.Schum 


Mimosaceae 


Sapindaceae 
Sapindaceae 
Aspleniaceae 


Aspleniaceae 


Aspleniaceae 


Aspleniaceae 


ae 
Barringtoniaceae 


Begoniaceae 


Lomariopsidaceae 
Lomariopsidaceae 
Lomariopsidaceae 


Flacourtiaceae 


ae 
Hymenophyllaceae 
Hymenophyllaceae 


Euphorbiaceae 
Verbenaceae 


50 SIDA 19(1) 
pale-palel Codiaeum varlegatum (L.) Blume Euphorbiaceae 
Codiaeum s Euphorbiaceae 
ah-rap Coix oaceae 
sagag-mum eee id esculenta (L. ) Schott Araceae 
mua-muadi (Scheffer) K.Schum. Marantaceae 
ta-wop a ominsia cf.minor Valeton Marantaceae 
kalagid Cordyline fruticosa (L.) A.Chev. Agavaceae 
keiki Costus speciosus ae Smith Costacea 
Wwapa-ruap Crinum asiaticum Amaryllidaceae 
monia-nasag Cryptocarya ee Blume Lauraceae 
rubu-gem-nasag Cryptocarya laevigata Blume Lauraceae 
vinisa La Viaa laevigata Blume Lauraceae 
wonkibung eee weinlandii K.Schum, Lauracea 
walo Cucurbita Cucurbitaceae 
taleb Cupani oe cee a Radik. Sapinda 
wadi-diri uPanOP SI is rps a Radlk. Sapindacea 
iliba G ur.) Kuntze Hypoxidaceae 
giligelum Cur rcuma cf. australasica Hooker f. Zingiberaceae 
wat-uri Panes papuanus Diels, or aff. Annonaceae 
wanga udial athocalyx sp. ?nov. Annonaceae 
masa-wun-bandep oye diss Vahl var. diffusus Cyperaceae 
wuk p Cyperus kyllingia Endl. Cyperaceae 
migim Cyrtandra sp., Ae Centrosiphon Gesneriaceae 
rubu-rubu Cyrtandra sp., section Centrosiphon Gesneriaceae 
sa-kam i as section Centrosiphon Gesneriaceae 
-ve um accrescens (Trin.) Stapf Poaceae 
wandarum ea eae eatin cee Aaeses 
duat-murukun Cyrtosperma cf.macrotum Becc.ex Engl. aceae 
ah-nah-sahr Decaspermut ae teatum (Roxb.) A.J. Scott ae 
var. bracteatum 
varatep Dendrobium macrophyllum A. Reich. Orchidaceae 
ambup Dendrocnide ternatensis (Miq.) Chew Urticaceae 
bago-bagot Dianella ensifolia (L.) DC Liliaceae 
voa-baga-bungam Dichapetalum sessiliflorum Leenh. Dichapetalaceae 
sakamb Dicliptera papuana Warburg Acanthaceae 
angkumamb Diospyros papuana Valeton ex Bakh Ebenacea 
usiman-kekerup Diospyros papuana Valeton ex Bakh Ebenacea 
vah-tib Diplora d'urvillaei (Bory) C. Chr Aspleniaceae 
wasimi Diplora d'urvillaei (Bory) C. Chr. Aspleniaceae 
kadimu eS cannaeformis (Forst. f) K.Schum. Marantaceae 
uaram ss s—sSS=rtatce'A Roxb. Dracaenaceae 
warubu-sopasop Dysoxylum brassii oe & Per Meliacea 
warubu-taleba Dysoxylum pettigrewianum FM. Bailey Meliaceae 
i-pap Dysoxylum sparst hott m ae bberley Meliaceae 
Dysoxylum sparsiflorum Mabberley eliacea 
sakas-sakamb Eclipta prostrata (L.) L. eraceae 
kuvu-kuv ma novoguineense Warburg, or aff. Urticaceae 
sumbu-wadab Flatostema novoguineense Warburg, or aff. Urticaceae 
rubu-rubu Elatostema sp., aff. macrophyllum Brongn. Urticaceae 
badidir cD ae Sai Radlk. indacea 
wumbu-ngam Oe ijsm.& Binn.) Kurz Euphorbiaceae 
dawaba-sivar rad phased odes i ) Mer imosaceae 


buko-bukop m Dest ssp. debile Equisetaceae 


en Hauke 


ai-ke-kav Erechtites valerianifolia (Wolf) DC Asteraceae 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


pat rigid Friandra fragrans van Royen & Steen. 
maber rythrospermum candidum (Becc.) Becc. 
asia gili-gilib ae dekockii (Valeton) R.M. Smith 
uphorbia hirta L. 
nae Fagraea ceilanica Thunb. 
seger-nganam Fahrenheitia sp. ?nov. 
tagle Ficus ampelas Burm.f. 
mariap-tobitobi Ficus bernaysii Kin 
bulubul Ficus ee Mia. var. subalbidoramea 
(Elmer) C 
pake-koal Ficus con a oxb. 
mariap-tobitobi Ficus conocephao ia Ridl. 
dambotan da psis ae var. obtusa Corner 
keimang Ficus a ani Warburg 
mutu-ngomb Ficus mollio 
maboramb Ficus odoardi King 
ngoku oe odoardi Kin 
ta-kup cus pungens ide ex Blume 
wah-ran-gab subulata Blum 
diga Ficus wassa Roxb. 
seger-buga Flacourtia inermis Roxb. 
wata-katok Freycinetia s 
dawab Garcinia dulc is (Roxb) Kurz 
da-da-da-dag Garcinia Gales Laut. 
magule Garcinia maluensis Laut. 
mugum Gastonia spectabilis (Harms) oe 
ivang-glu Geniostoma rupestre J.R.& G.Fo 
kivi-kiva Geniostoma rupestre J.R.& G. ae 
ngawar-dodol Geniostoma rupestre J.R. & G. Forst. 
warubu-nganam cecil chondrocarpum Airy Shaw, or aff. 
koita dion granulare Air 
widasag Gloctidio granulate Airy Shaw 
kawari m K.Schum. 
kwarikioari Gnetum gnemonoides Brongn. 
sang-guab Goniothalamus imbricatus Scheffer 
sang-guab Goniothalamus cf. imbricatus Scheffer 
sivar-wiav Goniothalamus cf. imbricatus Scheffer 
sivaru-guaru Goniothalamus sp. 
dai-dai-sivar Gouania cf. javanica Mi 
mago-ragor Grammatophyllum papuanum J.J. sm. 
muonia-kivikiva Graptophyllum ee (L.) Griff. 
vogerdak Haplostichanthus longirostris (Scheffer) 
ven aoe den 
lomal-nganam farpullia crustacea Radlk. 
rasapakay ampulla ramiflora Radlk. 
muat-upot-ugarum eae sp., Cf. H. auricularia L., or 
H. lapeyrousil DC. 
ngavisi Helicia affinis Sleumer 
gemea iconi 
urem dagur Hibiscus ellipticifolius Borss 
kolaiv olochlamys beccarti Eng| 
ma-nem-gab Holochlamys beccarii Engl 
ango-leb malomena magna A. Hay 
kibaip Homalom a at magna A. 
dalulup Horsfieldia lis (Miq.) Warburg var. subtilis 


Polygalaceae 


Moraceae 


Moraceae 
Moraceae 


Pandanaceae 


nnonaceae 
Sapindaceae 


Sapindaceae 
ubiaceae 


Prot 


Myristicaceae 


52 


kobou-susul 
e 


muat-upot-ugarum 

gawok 

mane-mane-kav 
abav 


wange-Wwadldp 
Wagu-wagum 


puale-pualel 
wat-uduat 
gilagal 

nganam idir idir 
idi-dir 
tukum-avang 


mane-mane-kav 
wanclap 


ah-maap 


wara-tep 
lasa-lasa 


Horsfieldia subtilis (Miq.) Warburg var. subtilis 
Hoya pottsii F.M. Baile 
Hulemacanthus novoguineensis (Lindau) Bremek. 
Huperzi a cf. squarrosa (Forst.f.) Trevisan 

t. 


indet. 


indet. 
el: fagifer (Parkinson) Fosb. 
Hoses fagifer (Parkinson) Fosb. 
Ino ie ee to) aa morphotype fide 


Ver 
ee ee (Colebr.) Kuntze 
oe congesta R.Br. 
Ixora sp., section Hypsophyllum 
‘a Cla Sp. 
ee eecurigna OND) Wedd. 
is K.Schum. 


Leea cf. arena Lau 

Leea heterodoxa K. aa. & Laut. 
Leea indica (Burm. f.) Merr. 
Lepisanthes senegalensis (Poir.) Leenh. 
Lepisanthes Poir.) Leenh. 
Lepisanthes senegalensis (Poir.) Leenh. 
Leptaspis urceolata (Roxb.) R. Br. 
Leucosyke cf. capitellata ee Chew 
Licuala beccariana Furta 

Licuala cf. beccariana Furtado 


Lygodium circinnatum (Burm. f) Swartz 
Macaranga giao Pax & Hoffm 
Harms) Philipson 
Maniltoa ‘schefferi K.Schum. & Hollrung 
Mapania macrocephala (Gaud.) K.Schum. 
ssp. macrocephala 
Medinilla triplinervia Cogn.,'mufoso-trip 

roup’ 
Medinilla sp., aff. tenuipedicellata Baker f. 
Medinilla sp., aff. tenuipedicellata Baker f. 
Medusanthera laxiflora (Miers) Howard 
Melicope sp., cf. M. burttiana Stone or 
M.grandifolia Burtt 
Melodinus cf. acutus oe Markaraf 
Aa au (L.) Mer 

Os sp. ?no 

Mi ixomelum minum (Forst. f) Wight & 
Walker-Arn 
Microsorum aun (Mett.) Copel. 
Microsorum membranifolium (R. Br.) Ching 


nervid 


SIDA 19(1) 


Myristicaceae 


Orchidaceae 
indet. 
Fabaceae 
Fabaceae 
Fabaceae 


Caesalpiniaceae 


Lomariopsidaceae 
Rutaceae 
Schizaeaceae 


Caesalpiniaceae 
Cyperaceae 


Melastomataceae 


Melastomataceae 
Melastomataceae 
Icacinaceae 


Apocynaceae 
Convolvulaceae 
Tiliaceae 
Rutaceae 


Polypodiaceae 
Polypodiaceae 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


boge-namb 
say-ri-keep 
go-idi 


mum-nganam 
pah-tooey 


u 
sang-quab 
kobos-susul 
sagua 
amungcurcurl 


sretu-ngomb 
mondi-minab 
ura 
quasi-kwas 
Sivila 


Sauga-sivar 
tibaga 


mabarara-dangamb 


mamba-mambap 
mamba-mambap 


timber-digeep 
kasapa 
sauiak 


bogo-namb 
guragor 


sopi-sebip 


kamora-kamorap 
ah-mo-rap 
amora-kamorap 


mara-marav 
bial 
wo-roon-botop 
aru 


Mennas Mine NaKG L. var. papuana Valeton 
K.Schum 


Masa! banksit Fu M, 


rck 
Mussaenda oi lindrocarpa ane 
Mussaenda cylindrocarpa Burck 
Meet ia aval lea peal nae ex Korth. 
oir. ssp. lancifolia 
Myristica ees ia | ssp. lancifolia 
Myristica subalulata Mig. var. subalulata 


Myristica tristis Warburg, 0 
Neisosperma citrodorum L aut. & K.Schum) 
Rose: & Sach. 

perma jtrodorum (Laut. & K.Schum.) 
Fosb. & Sach. 


(A. Gray) Leenh. 
Neuburgi a rumphi jana Leenh. 


laldalaiadiaaliiaa) 


onl ea queer lance F.M. Bailey 


etala Laut.& K.Schum. 
Osmelia ohilinding a Benth. 
Osmoxylon sessiliflorum (Laut.) Philipson 
Osmoxylon (closest to) sessiliflorum (Laut.) 
Philipson 
Pandanus angiensis Kaneh., or aff. 
Pangium edule Reinw 


Pararistolochia schlechteri (Laut.) M.J. Parsons 
Paraserianthes falcataria (L.) Nielsen cf. ssp. 
falcataria 

Peristylus guage JJ.Sm 

Petalolophus sp., aff megalopus K.Schum. 
Phaleria coccinea (Gaud.) F.v.M 


Phrynium cfm acrocephalu um 1K. Schum. 
Pg um pe ie Warburg, or aff. 
jum p ) Warburg, or aff. 
Prim sp aft oman K.Schum. 
Phryn 
Prylacium bracteosum Benn. 
hy JJ.Sm. 


Phy llanthy ithriflorns JJ. Sm. 


Pimelodendron amboinicum Hassk. 


y 
Piper pseudoamboinense C.DC. 
sae pu llibaccum ae se 


urus argenteu st. f) Wedd. 
Prptuusc agency (Forst f) Wedd. 
is Teijsm.& Binn. 


Rubiaceae 
F eae 


pocynaceae 
Apocynaceae 


Loganiaceae 


Euphorbiaceae 
Opiliaceae 
Arecaceae 
Flacourtiaceae 
Araliaceae 
Araliaceae 


Pandanaceae 
Flacourtiaceae 
Poaceae 
Aristolochiaceae 
Mimosaceae 


Orchidaceae 


Euphorbiaceae 
Euphorbiaceae 
Euphorbiaceae 


Nyctaginaceae 


54 


sesambop 
lasa-lasa 
bum 
gaga-bumer 
kidi-kidi 


man-duroop 
sana ngamb 


tatar-ulalat 
warang-gab 
sigawa 
lawalang wiab 
sivar-wiav 
wato-karok 


gaga 
pat dagol dagol 
televa-nganam 


mavanda-ngamb 
kolaiv-nganam 


babagalum 
ga-tsurup 


sob-barewa 
wana-barewa 
babagalum 


wasagep 


giligelum-sivar 
gaga-bumer 
idang io 
ngumusinam 


mago-ragor 
kasuar dadi 


ooh-rau-rap 
gagap 
maberu 


Pisonia longirostris Teijsm. & B 
a3 e€0C cnemia macr fog0 nta (Fée) alton 
ta (Fe 


f Son fubercuaiimn (Cesati) Holttum 


Pleuranthodium sp., ?trichocalyx (Valeton) 
Smith 


Plocoglottis cf. moluccana Schltr. 
Pneumatopteris sogerensis (Gepp) Holttum 
Pneumatopteris sp 
Holttum 
Podocarpus cf. rumphii Blume 
Poikilospermum amboinense Zipp. ex Mia. 
Polytocha macrophylla Benth. 

Popowia sp.,aff. pisocarpa (Blume) Endl. 
Popowia s 

Pothos papuanus Becc. ex Endl. 

Pothos rumphii Schott 

Pronephrium micropinnatum Holttum 


Pseuderanthemum sp.,cf.'variabile group’ sensu 
Barker 


Pseudobotrys dorae Moese 


Pseuduvaria sp., ai (Diels) Merr. 
Pseuduvaria 
Pseuduvaria sp. 
EsevaUMatiG sp. 
I litt Valeton 
ut.& K.Schum. 


cae a ia & K.Schum, 
Bes Ee Miq. var. leptothyrsa 
uchi 


Psychotria peed ee Bartl. ex DC. 
Psychotria membranifolia Bartl.ex DC. 
Psychotria cf membranifolia Bartl. ex DC. 


Psychotria phaeochlamys (Laut. & K.Schum. 
V 


aleton 


Psychotria phaeochlamys (Laut. & K. Schum.) 
Valeton 


Psychotria sp. nov. 

Psychotria sp.n 

oe sp aff micralabastra (Laut. & 
K.Schum.) Valeton 

Psychotria sp., aff. micralabastra 

(Laut.& K.Schum.) Valeto 

Psychotria (possibly new ae sp.) 

Pteris warburgii Christ 

Ptyssiglottis pubisepala (Lindau) B. Hansen 

Pueraria pulcherrima (Koorders) Koorders- 

Schumacher 

Pyrrosia princeps (Mett.) Morto 

Kia - ecora Val.,or Gece 

K.Schum 

ieee aia korthalsii Schott 

Si idophora versteegii Engler & Krause 
hyticaryum longifolium K.Schum. & Laut. 


aff. keysseriana (Rosenst 


SIDA 19(1) 


Nyctaginaceae 
Tectaria group 
Tectaria group 
Thelypteridaceae 
Zingiberaceae 


Orchidaceae 
Thelypteridaceae 
Thelypteridaceae 


Podocarpaceae 
Cecropiaceae 
P 


helypteridaceae 
Acanthaceae 


Icacinaceae 


Rubiaceae 
Rubiaceae 
Rubiaceae 
Rubiaceae 


Rubiaceae 


Rubiaceae 


Fabaceae 


Polypodiaceae 
Rubiaceae 


Araceae 
Araceae 
Icacinaceae 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 55 


daveh-veh Rhyticaryum novoguineense ee) Sleumer — Icacina 
ma-rab Riedelia grandiligula Valeto Zin see 
ma-rab Riedelia macrantha K. ane Zingiberaceae 
ninara-umu Rinorea horneri (Korth.) O.K. Violaceae 
sekera-nasag Rinorea horneri oe O.K Violaceae 
vorap Rub | var. discol or (Blume) 
Kalkman 
pat-sani-sani Ruellia sp. ae eee Acanthaceae 
umbol-lap ale flora Blu Sabiaceae 
wang-gep Sap ame (K.Schum. & Laut.) Rubiaceae 
ea 
atep Schismatoglottis sp. Araceae 
gibaiv Schismatoglottis sp Araceae 
kibaip eal S Araceae 
kasipul Schizostachyum tue ale Merr. Poaceae 
moimoit Scleria polycarpa Boec Cyperaceae 
pupun-lov Selaginella - ee = Selaginellaceae 
manda-peb Selaginella ee VAM. Selaginellaceae 
lawa-lawat Semecarpus a Merr. & Perry Anacardiaceae 
wingam Semecarpus forstenii Blume Anacardiaceae 
lawa-lawat Semecarpus magnificus K. Schum Anacardiaceae 
wat-ukauel Smilax cf. australis R. Smil ae 
O-r Solanum Solanaceae 
kasiwar-gili-giliba Spathoain plicata Blume ssp. puberula Orchidaceae 
.Howcroft 
komekelak oe arfakianus (Baker) Holttum Thelypteridaceae 
ngaparu-pot ene pilososquamatus (v.A.VR.) Thelypteridaceae 
olttum 
tuturat Stachytorphet eeataee (Rich.) M.Vahl Verbenaceae 
ave-namb Valeton) Kaneh. & Hatus. Monimiaceae 
marap Steganthera dentata Neleton) Kaneh. & Hatus. Monimiaceae 
bailalum picgantney mi meta ete) Perkins onimiaceae 
aramitap tegant ns (Becc.) Kaneh.& Hatus. Monimiaceae 
ma-bairap Stenochlaena milnei Underwood Blechnaceae 
kokam-tol culia schumanniana (Lau t.) Mildbr. Sterculiaceae 
nanggu-nanggu-nam ae eo s.1., Hemigraphis primulifolia (Nees) Acanthaceae 
Vill. 
Sagag-gosmun Strobi ae s.l. (Hemigraphis sp.) Acanthaceae 
Sagag-u-goga-umun Strobilanthes s.|.(Hemigraphis sp.) Acanthaceae 
sagag-ugosum Strobilanthes s.|. (Hemigraphis sp.) Acanthaceae 
sumure _weeiianaiies = MEI apis sp.) Acanthaceae 
mansu-borobor m (Diels) Merr. & Perry Myrtaceae 
navyia Syagum oe anu ae & Perry yrtaceae 
padada Syz m (Laut.& K.Schum.) Myrtaceae 
Merr, & Perr 
mansu-borobor Syzygium Dae Merr. & Per Myrtaceae 
kah-bik Syzygium pteropedum (Laut. & K.Schum.) Myrtaceae 
rr. & Per 
magule ae eae (Diels) Merr. & Per Myrta 
da-da-dag eile sp., aff. goniopterum (Diels) Merr. a Perry vee 
navyia Syzygium 
uaia ae aurantiaca Gaud. Apocynaceae 
umbol-menyap Tabernaemontana aurantiaca Gaud. Apocynaceae 
samangi manggib Tabernaemontana orientalis R. Br. Apocynaceae 
keiti Tapeinochilos recurvatum K.Schum. Costaceae 


56 SIDA 19(1) 
keiti Tapeinochilos sp. no Costaceae 
wange-abab Tarenna gu ee (K. Schum.) Valeton Rubiaceae 
yag-tauita Tarenna gtilcheriana (K.Schum,) Valeton Rubiaceae 
takevam Hera a AEA ides (Presl) Copeland Tectaria group 
bogang-dap Coode ombretaceae 
danga-namb lemingia 1 mpecens Coode Combretaceae 
anganange-woganamb v.M. Dilleniaceae 
od nee ee anum Gilg Vitaceae 
bon Thespesia fissicalyx Borss. Malvaceae 
ma-ka Trichosanthes sp.,'l ees group’ Cucurbitaceae 
ikikap panda disti Oe Sch Orchidaceae 
idi-muyat Versteegia caullfl ‘Schum. & Laut.) Valeton Rubiaceae 
waipa | gia cauliflora (K.Schum. & Laut.) Valeton — Rubiaceae 
wanam-barewa verte Spal ia Valeton Rubiaceae 
ker-ker-kanamb Vrydagzynea cf. rivularis Schltr. Orchidaceae 
wungo-bunya Wedelia biflord (L (L.) D Asteraceae 
maruruma Wenzelia dolichophylta a (Laut. & K.Schum.) Tanaka Rutaceae 
ngabu-kuruk Zanthoxylum conspersipunctatum Merr.& Perry — Rutaceae 
ambo-dera-namb Ziziphus djamuensis Laut. hamnaceae 


APPENDIX 3 
ETHNOBOTANICAL VALUE OF JOSEPHSTAAL PLANTS 


PLANTS WITH FOOD VALUE 


Amaranthus dubius Thell; leaves edible 
Artocarpus communis J.R. & G. Forst; seeds are 


eaten 

Bambusa microcephala (Pilger) Holttum; young 
shoots are eaten 

Buchanania macrocarpa Laut, the rotting wood 


ly to ae to promote Heir physical 


dev 
Callioter 5 ae inulosa 


(Blume) J.Smith;new leaves 
are edible 
ay ta sp.; fruits and leaves are edible 
plora d'urvillaei (Bory) C ek leaves are burnt 
and the ashes used as salt 
Fu dammaropsis Diels v var. obtusa Corner; fruit 
ee 
Ficus wassa Roxb, eaten as a vegeta 
Inocarpus rubidus morphotype fide ae 
seeds are edible 


episanthes senegalensis (Poir.) Leenh.; fruit is 
edible, fed especially to children to improve 
their growth 

Lomagramma cf.sinuata C.Chr, leaves are edible 

Melicope sp., cf. M. burttiana Stone or M. 
grandifolic ia 1B, L. Burtt; the plant is a source of 
edible leaf caves 

Fang edule 


aie edible 
| Pe pal 


wood is a good source of edible larvae 
Pneumatopteris sogerensis (Gepp) Holttum; 
young shoots or fronds eaten as a vegetable 
PREGrODIEy: sp., aff. rae ana (Rosenst.) 
| edible, cooked with meat 
Schismatoglott 5 SP. young leaves are edible 


leaves dic 


edible 
Terminalia impediens Coode; seed is edible 
Trichosanthes sp., ‘longiflora-bracteata group; 
fruit is edible 


MEDICINAL OR PSYCHOACTIVE PLANTS 


Alocasi a aequiloba N.E. Br; leaves used to treat 
pain from salat (stinging nettle) injuries 
lauterbachiana (Engl.) A. Hay; leaves 
sed to alleviate pain ot nettle stings 
jee a macrocalyx Zipp.e Blume; chewed as 
a substitute for buai eo 


ae 


sean Becc.; mature fruits 
substitute for buai 
Cassia alata Ct: ee to treat ringworm and skin 
diseases 
Cyperus kyllingia Endl; leaves are boiled and 
sed for body aches and diarrhea 


Calyptroca 


ia) 
oS 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 57 


Euphorbia hirta L; leaves boiled and the solu- — Psychotria membranifolia Bartl. ex DC.; roots 


ion is used to treat fever and cold symptoms mashed and mixed with coconut juice, given 
Licuala beccariana Furtado; mature nuts are to children to treat malaria and stomach 
wed as a buai substitute disorders 
Lunasia amara Blanco var.amara; young leaves —_Scleria polycarpa Boeck.; plant is cooked in bam- 
ee over fire and the juice squeezed onto oo and eaten to induce abortion during 
sore the early stages of pregnancy, cf. Mills s.n. 
Piper a caninum Blume; spikes and all other from Josephstaal 
parts chewed with buai Tabernaemontana orientalis R. Br; roots are 
Piper decumanum (Rumph,) L.; roots are chewed boiled and the solution consumed to pro- 
with buai mote aggression 


PLANTS USED IN CONSTRUCTION OR FOR MAKING IMPLEMENTS 


Aglaia cuspidata C.DC.; wood is used for mak- — Intsia es (Colebr.) Kuntze; trunks used for 
ing spears making garamuts, also a strong timber for 
Alangium villosum (Blume) Wangerin ssp. 
es (C.T.White) Bloembergen;poles — Licuala beccariana Furtado;leaves used as roof- 


sed for house rafters ing for bush shelters 
een ee schumannii (Becc.) Essig; planks = Macaranga fallacina Pax & Hoffm.; used in mak- 
house flooring, also made into ing rafters for houses 
implements for sharpening bows and ar- — Neuburgia corynocarpa (A. oo Leenh.; wood 
rows used in house construc 
Calamus humboldtianus Becc.; canes are split — Porterandia sp.; poles are a in making cas- 
and re as ropes for tying and fastening sowary traps 
(e.g. e and fence construction) Pseuduvaria sp.; used as timber poles during 
a macro asi. or aff; poles used as house construction 
digg mplem Psychotria ee Bartl. ex DC.; stems 
een ett . a acta eenedes used as a planting implement for mami 
for house buildin Ce esculenta) 
Dichapetalum sessiliflorum Leenh.; used as ties st sp. nov,; the wood is used for making 
and bindings in house construction digging sticks for planting yams and mami 
lis cannaeformis (Forst. f.) K. Schum.; stems (Dioscorea esculenta); crop yields are be- 
sed as rope | for house building tees to increase when this particular wood 
D | | for tool 
handles ;wood i is very stron Schi ead lima (Blanco) Merr.; used for 
Garcinia maluensis Laut.; Wee ae used asa makin ug po veuine cf.R. Pullen 1 
: eee oe or plant ng implement for — Versteegi ra (K.Schum, & Laut.) Valeton; 
mi (Dioscorea esculenta) wood used as cultivation too 
a“ ae lipo Borss.; bark is used as wall Yrteega sono Valeton; stem is used as a 
nels in ho digging implement for planting, it is a tradi- 


fiona pellertiat this increases yam yields 


PLANTS WITH CEREMONIAL, RITUALISTIC, OR SPIRITUAL APPLICATIONS 


Agl d Hooker) Roos; leaves seals taro shoots are dipped into the sap 

used in ritual ceremonies nd planted, said to increase yield 
Antrophyum cf. reticulatum (Forst.) Kaulf.; leaves ae hortensis J.R.& G.Forst.;juice extract used 
ford litional erfume bodies during sing-sings, cf. NGF 


bilas 102 

Dracaena angi HOXE me leaves cut — Ficus odoardi King; sap is rubbed on yam before 
wash chil- planting to increase growth 

to prevent crying and chase away spirits = Holochl/ [ i Engl 1 in magic rituals 

Entada phaseoloides (L.) Merr; sap collected in to increase abundance of game animals 


58 


Huperzia cf. estes (Forst. f.) Trevisan; plant 

ii se aa Cui rain clouds as - ecial 
ae e spoken to stop the ra 

Microsorum ingu forme (Mett,) cope! eaves 


dage  solts st (Laut & K.Schum.) Fosb. 
& - as 


yie 
eae a as decora- 
e bi ne in ye eae 


SIDA 19(1) 


Tabernaemontana lan ning fruits are 
d Christmas or 


Cc 
Ww 
i) 


fate a a orn R Hy Ha used 
in rituals to improve crop grow 

Tetracera en ana Fv.M.; water in rhe vine is 
used in black magic to inflict illness 

Trichosanthes sp.,longiflora-bracteata group’;sap 
rom vine is used in hunting rituals 


PLANTS OF PARTICULAR VALUE TO WILDLIFE 


A i fst Aaa ery P| eS . 


by bandicoots 

Aglaia lepidopetala Harms; mature fruits eaten 

ssums 

Archidendron aruense (Warburg) de Wit; flower 
nectar crise by bandicoots; seeds eaten 
by ban 

Arytera sp., aff litoralis al ‘litoralis complex’; 
fruit eaten by poss 

goodies mmacocapo ne fruits eaten by 

SSO and other birds 

Gio aera Becc,; mature fruits 
eaten wa 

ae ue Gilg or aff; fruits eaten by 


eee floribunde K. Schum.,; fruits eaten by 
casso 

cyptocnya ee Blume; fruits eaten by 
cassowaries 

Cece peseneck — or aff; ripe fruits 
eaten by cassowa 

Diospyros papuana leion ex Bakh.; fruits swal- 

by San 
err. & Perry; ripe fruits eaten 


by possums 

Dysoxylum pettigrewianum F.M. Bailey; ripe fruits 
eaten SSuUMS 

Ficus botryocarpa Miq. var. subalbidoramea 


Saal Corner; mature fruits eaten by bandi- 
ots and bats 
Ficus sae Roxb.; ripe fruits eaten by 


andic 
Ficus ae Ridl.; fruits eaten by 
bandicoots 
Ficus pungens sass ex Blume; eaten by birds 
and bandico 


Lali-i. ary. | f, H + h 


[ i a | lh H frist + kK 
t 


assowari 
ae ia macrpetal Laut.& K.Schum. ripe fruits 
nb by cassowa ries 


Pi ; | 


Pipturus aa (Forst. f.) Wedd; fruits eaten 


y 
oy anae sp.; fruits are swallowed by cas- 
ri 
sychotria micralabastra (Laut. & K. Schum.) 
eton; fruits eaten by birds 
Pyrrosia pri oe (Mett.) Morton; used as shelter 


Syzygium ccranthum (Diels) Merr.& Perry; many 
animals eat the fruit and seeds 

Syzygium | ce Merr. & Perry; fruits eaten by 
cas 

Syzygium pteropodum (Laut. & K. Schum.) Merr. 
& Perry; fruits eaten by cassowaries 


PLANTS USED ON DOGS 


Aglaia sapindina (Fv. M.) Harms; young leaves are 


Alocasia aequiloba N.E. Br, petiole base and roots 
fed to hunting dogs to stimulate aggression 


Pre pa + 


to stimulate aggression in the hu 
ese laevigata Blume; fed to ae to in- 
heir ability to hunt bandicoots 
eee cuspidispathum Alderw.; peduncle 
and spadix cooked in bamboo and fed to 
hunting dogs to promote aggression 


PLANTS WITH OTHER CULTURAL APPLICATIONS 


Arytera sp., aff. litoralis Blume, ‘litoralis complex: 
ell 


Ca alsa yx holl fase Becc,; leaves used for 


resin iS burned at night as GLGIIUNE substitu te 


fresh meat caught in the 


WI dpE 


bush 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 


Cleistanthus sp., aff. 2>apuanus (Laut) Jabl.; said to 
be a particularly good firewood for cooking 

Coix lachryma-obi L, fruits used to make necklaces 

Cominsea cf. minor Valeton; leaves used as wrap- 
ping for sago 

Curcuma cf. australasica Hooker f; a source of 

yellow dye 

Harpulli ia amie Radlk; bark is stripped and 
used as a fish poison 

Helicia affinis Sleumer; ee fruits provide a dark 
purple dye (e.g. for bilums) 

Heliconia papuana W.J. s; leaves used for 
wrapping food, eg. ‘es ert in the 


— 


Hibiscus ellipticifolius Borss.; bark is peeled in 
strips as a sleeping ma 
Leucosyke cf.capitellata (Poir.) Chew; leaves used 


59 


Microcos sp. ?nov.; used as a fish poison, cf. NGF 


102 

Morinda ee a root bark is used for 
dye, cf. N 

Ocimum ee imum cin used for perfume 

Phrynium cf. macrocephalum K. Schum.; leaves 
used as a wrap for garden vegetables and 


bush meat 

Phrynium pedunculatum Warburg, or aff; leaves 
used for wrapping sago 

Pittosporum sinuatum Blume var. sinuatum; 
young leaves are mashed and rubbed on 
diving goggles to prevent foggi 

Planchonia Be Knuth; bark is used as fish 
poison, cf. NGF 10250 

Psychotria amplithyrsa Valeton; mature fruits 
mixed with pig food to promote fattening 


as toothbrush Psychotria membranifolia Bartl.ex DC; leaves fed 
Lygodium circinnatum (Burm. f.) Swartz; stems to pigs to promote fattening 
sed to make arm bands 


ACKNOWLEDGMENTS 


The Josephstaal expeditions were funded by the Nature Conservancy (TNC). Survey data 
cited in the present account are used by permission of that organization. My studies in 
PNG are otherwise maintained by a grant from the John D.and Catherine T. MacArthur 
Foundation, to BRIT Head of Floras John Pipoly Ill. The PNGFRI and Lae National Her- 
barium provided facilities support. 

For the findings reported here | am indebted to the field assistance by Maya Gorrez 
(TNC), Ali Towati (BRIT parabotanist), and Joseph Wiakabu (LAE). Dr. Earl Saxon (TNC re- 
gional ecologist) was the senior investigator. Other participants whose efforts contrib- 
uted to the expeditions’ successes included Francis Bebe, Susan Brown, Edward Mayer, 
and Rene Weterings. 

The Latin diagnoses were corrected or written by John Pipoly, and the illustrations 
of new taxa were prepared by N.H.S. Howcroft. John Pipoly also responded to my re- 
quests for information from the BRIT botanical library. Several identifications of difficult 
collections were successfully made by senior dendrologist Kipiro Damas (LAE). My ecol- 
ogy associate Hitofumi Abe of the Japan International Cooperation Agency, kindly pro- 
vided the translation to Japanese script. Taxonomic specialists assisting with ongoing 
work on the specimens include P.Forster (Asclepiadaceae), N.H.S. Howcroft (Orchidaceae), 
K.L. Huynh (Freycinetia), and PF. Stevens (Clusiaceae and passim). Referees PF. Stevens 
and J.Pipoly made numerous improvements to the draft with their effective criticism. 


REFERENCES 
Airy SHaw, H.K. 1975. The Euphorbiaceae of Borneo. Kew Bull. Addit. Ser.4:1-245. 
. 1978. Notes on Malesian and other Asiatic Euphorbiaceae. Kew Bull. 32: 
361-418. 


60 SIDA 19(1) 


.1980.The Euphorbiaceae of New Guinea. Kew Bull. Addit. Ser. 8:1-243. 

AUSTRALIAN SurveY Corps. 1973. Papua New Guinea 1:100,000 Topographic Survey Map. Nubia 
sheet 7889 (reprinted by the National Mapping Bureau 1985). 

__.: 1974a. Papua New Guinea 1:100,000 Topographic Survey Map. Annanberg 
sheet 7888 (reprinted by the National Mapping Bureau 1981). 

.1974b. Papua New Guinea 1:100,000 Topographic Survey Map.Adelbert sheet 
7988 (reprinted by the National Mapping Bureau 1986). 

.1974c. Papua New Guinea 1:100,000 Topographic Survey Map. Manam sheet 
7989 (reprinted by the National Mapping Bureau 1987). 

Batcooy, M.MJ. van. 1971, Plant geography of the Pacific. Blumea suppl. 4:1-222. 

/P.Hovenkame, and P. van Wetzen. 1996, Phytogeography of the Pacific—floristic 
and historical distribution patterns in plants. In: A. Keast and S. Miller, eds. The origin 
and evolution of Pacific island biotas: New Guinea to Eastern Polynesia: patterns and 
processes. SPB Academic Press, Amsterdam, the Netherlands. Pp 191-214. 

Barer, R.M. 1986. A taxonomic revision of Australian Acanthaceae. J. Adelaide Bot. Gard. 
9:1-286. 

BLeEKER, P.1983. Soils of Papua New Guinea.CSIRO and Australian National University Press, 
Canberra. 

Conn, B.J. 1994. Documentation of the flora of New Guinea. In: C-1 Peng and C.H. Chou, 
eds. Biodiversity and terrestrial ecosystems. Institute of Botany, Academia Sinica Mono- 
graph 14:123-156. 

Copetanp, E.B. 1949. Aspleniaceae and Blechnaceae of New Guinea. Philip. Journ. Sci. 
78(2):207-229. 

Crort, J.R. 1981. Hernandiaceae. In: E.E. Henty, ed. Handbooks of the flora of Papua New 
Guinea 2:190-201. 

Darwin, S.P. 1979. A synopsis of the indigenous genera of Pacific Rubiaceae. Allertonia 
11-44. 

DeparTMENT Or District ADMINISTRATION. 1968. Village Directory. Territory of Papua and New 
Guinea. G.W. Reid, Acting Govt. Printer, Port Moresby, Papua New Guinea. 

Duyries, B.E.E. 1996. Hernandiaceae. Flora Malesiana ser. 1, 12(2):737-761. 

Essic, F.B. 1977. The palm flora of New Guinea. A preliminary analysis. Papua New Guinea 
Office of Forests, Bot. Bull. 9:1-39 

Foreman, D.B. 1976. A taxonomic study of the genus Helicia Lour. (Proteaceae) in New 
Guinea and Australia with notes on origin, distribution and ecology. MS. thesis. Uni- 
versity of New England, Armidale, New South Wales. 

.1995.Proteaceae.In:B.J.Conn, ed. Handbooks of the flora of Papua New Guinea 
3:221-270. 

Forman, L.L. 1986. Menispermaceae. Flora Malesiana ser. 1, 10(2):157-253. 

Forster, Pl. 1992.Circumscription of Tat | lacaqui (Apocynaceae) in Aus- 
tralia. Austr. Syst. Bot. 5:521-531. 

GIDEON, O.G. 1998. Systematics and evolution of the genus Tapeinochilos Miq. (Costaceae- 
Zingiberales). Ph.D. thesis. James Cook University of North Queensland, Australia. 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 61 


HAMMERMASTER, E.T. and J.C. Saunvers. 1995a. Forest resources and vegetation mapping of 
Papua New Guinea. PNGRIS Publ. 4, Canberra, CSIRO and AIDAB. 

—____ ,and _____. 1995b. Forest resources and vegetation mapping of Papua 
New Guinea. 1:250,000 vegetation map overlays separately issued as working copies 
to PNGRIS Publ. 4, Canberra, CSIRO and AIDAB. 

Harms, H. 1925.Die Cucurbitaceen Papuasiens. Bot. Jahrb. Syst.60:150-161. 

HarrLey, 1.G. 1966. A revision of the Malesian species of Zanthoxylum (Rutaceae). J. Arnold 
Arbor.47:171-221. 

Hay, A. 1999. Revision of Homalomena (Araceae-Homalomeneae) in New Guinea, the 
Bismarck Archipelago and Solomon Islands. Blumea 44:41-71 

and R. Wise. 1991. The genus Alocasia (Araceae) in Australasia. Blumea 35: 
499-545. 

HENoRIAN and D.J. MindLeTON. 1999. Revision of Rauvolfia (Apocynaceae) in Malesia. Blumea 
44:449-470. 

Heusoen, E.C.H. van. 1992. Flowers of Annonaceae: morphology, classification, and evolu- 
tion. Blumea suppl. 7:1-218. 

Howttum, R.E. 1954. A revised flora of Malaya. Volume Il. Ferns of Malaya. Govt. Printing 
Office, Singapore 

Jaques, A.L.and G.P, Rosinson. 1975. Explanatory notes on the Bogia geological map.Report 
75/12.Geological Survey of Papua New Guinea. Dept. of Lands, Surveys, and Mines. Pp 
1-20 and 1:250,000 map. 

JoHNs, R.J.1977.The vegetation of Papua New Guinea. Part 1:An introduction to the veg- 
etation. PNG Office of Forests, reprinted 1984 

________.. 1986. The instability of the tropical ecosystem in New Guinea. Blumea 31: 
341-371. 

KARTAWINATA, K. 1990. A review of natural vegetation studies in Malesia, with special refer- 
ence to Indonesia. In: P. Baas, K. Kalkman, and R. Geesink, eds. The plant diversity of 
Malesia, proceedings of the Flora Malesiana symposium commemorating Prof. Dr. 
C.G.GJ.van Steenis. Kluwer Academic Publishers. Pp 121-132. 

Mcavpine, J.R., G. Keic, and R. Fats. 1983. Climate of Paoua New Guinea. CSIRO and Austra- 
lian National University Press, Canberra. 

Mack, A.,ed.1998.A biological assessment of the Lakekamu Basin, Papua New Guinea. Rapid 
Assessment Program Working Papers 9, Conservation International, Washington, D.C 

Mansretp, R. 1925. Die Melastomataceen von Papuasien. Bot. Jahrb. Syst.60:105-148. 

Merritt, E.D.and L.M.Perry. 1939.On the Brass collections of Pandanaceae from New Guinea. 
J.Arnold Arbor. 20:139-186. 

. 1943. Plantae Papuanae Archboldianae, XIll. J. Arnold Arbor. 


and 
24:422-439. 

i and__—s«d1:946, Plantae Papuanae Archboldianae, XVII. J. Arnold Arbor. 
PA eel oa a oy 

PaumaNs, K. 1975. Explanatory notes to the vegetation map of Papua New Guinea. Land 


62 SIDA 19(1) 


Research Series 35,CSIRO, Melbourne, Australia.Pp 1-25 with 20 plates and 1:1,000,000 
vegetation map. 

ed. 1976. New Guinea vegetation. CSIRO and Australian National University 
Press, Canberra. 

PANNett, C.M. 1992. A taxonomic monograph of the genus Agiaia Lour. (Meliaceae). Kew 
Bull. Addit. Ser. 16:1-379. 

Payens, J.P.D.W. 1967. A monograph of the genus Barringtonia (Lecythidaceae). Blumea 
15:157-263. 

Petir, A., D. Marerem, P. Yaron, S. Mukarek, M. Oxira, and T. PLatt-Mitts. 1998. Useful plants of 
Salemben Village, Madang Province, Papua New Guinea. Christensen Research Insti- 
tute Publication 13. 

Picram, C.J.and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea 
orogen. J. Austr. Geol. Geoph. 10:193-211. 

Purr,C.and K.M.Wonc. 1993. A synopsis of the genera of Rubiaceae in Borneo. Sandakania 
2:13-34. 

ReicH, A. 1998. Vegetation part 1: A comparison of two one-hectare tree plots in the 
Lakekamu Basin.In:A.Mack,ed.A biological assessment of the Lakekamu Basin, Papua 
New Guinea. Rapid Assessment Program Working Papers 9, Conservation International, 
Washington, D.C. Pp 25-35, 97-104. 

Riospate, C.E. 1974. A revision of the family Leeaceae. Blumea 22:57-100. 

R.C. BAKHUIZEN VAN DEN Brink, and J. Koek-Noorman. 1972. Notes on New Guinea 
Rubiaceae. Versteegia and Maschalodesme. Blumea 20:339-348. 

Royen, P. van. 1964. Manual of the forest trees of New Guinea. Part 3. Sterculiaceae. Depart- 
ment of Forests, Division of Botany. 

SAuNpers, J.C. 1993. Forest resources of Papua New Guinea. Explanatory notes to map. 
PNGRIS Publication 2, CSIRO and AIDAB, Canberra. Pp 1-18 and color maps (4) at 
1:1,000,000 scale. 

SCHIRAREND, C. 1995. Rhamnaceae. In: E. Soepadmo, and K.M.Wong, eds. Tree flora of Sabah 
and Sarawak 1:305-319. 

Steoce, W.A. 1962. Asplenium affine Sw. and A. spathulinum J. Sm. ex Hook. Kew Bull. 15: 
401-410. 

Steumer, H. 1954. Flacourtiaceae. Flora Malesiana ser. 1, 5(1):1-106. 

. 1955. Proteaceae. Flora Malesiana ser. 1,5(2):147-206. 
. 1971. Icacinaceae. Flora Malesiana ser. 1, 7(1):1-87. 

Smith, R.M. 1986. New combinations in Etlingera Giseke (Zingiberaceae). Notes Royal Bot. 
Gard. Edinb. 43:243-254. 

Somer, S.H. 1988. The nonclimbing species of the genus Psychotria (Rubiaceae) in New 
Guinea and the Bismarck Archipelago. Bishop Museum Bull. Bot. 1:1-339. 

Stevens, PF. 1989, New Guinea. In: D.G. Campbell and H.D. Hammond, eds. Floristic inven- 
tory of tropical countries: the status of plant systematics, collections, and vegetation, 
plus recommendations for the future. New York Bot. Gard., New York. Pp. 120-132. 


TAKEUCHI, BOTANICAL RESULTS FROM JOSEPHSTAAL 63 


Stone, B.C. 1985. New and noteworthy paleotropical species of Rutaceae. Proc. Aca. Natl. 
Sci. Phil. 137:213-228. 

Swinc_e, W.T. 1967. The botany of Citrus and its wild relatives. In: W. Reuther, H.J. Webber, 
and L.D. Batchelor, eds. The Citrus industry, vol. 1. History, world distribution, botany, 
and varieties. University of California, Berkeley. Pp 190-430. 

TAKEUCHI, W. 1999a. Botanical results from the 1995 Bismarck-Ramu expedition in Papua 
New Guinea. Sida 18:751-782. 

. 1999b. New plants from Crater Mt., Papua New Guinea, and an annotated 
checklist of the species. Sida 18:961—1006. 

,and J. KuLanc. 1998. Vegetation part 2: botanical survey. In: A. Mack, ed. A bio- 
logical assessment of the Lakekamu Basin, Papua New Guinea. Rapid Assessment 
Program Working Papers 9, Conservation International, Washington, D.C. Pp 36-39, 
105-130. 

Turner, H. 1994. Arytera. In: Sapindaceae. Flora Malesiana ser. 1, 11(3):467-479. 

Vateton, Th. 1911. Rubiaceae. Nova Guinea 8:437-519. 

. 1913. Zingiberaceae. Nova Guinea 8:923-988. 

. 1914. Die Zingiberaceen Deutsch-Neu-Guineas. Bot. Jahrb. Syst. 52:40-100. 

. 1918. New notes on the Zingiberaceae of Java and Malaya. Bull. Jard. Bot. 
Buitenzorg, ser. 2,27:1-166. 


_______-. 1927. Die Rubiaceae von Papuasien. Il. Zweiter Teil: Coffeoideae. Bot. Jahrb. 
Syst.61:32-163. 

Verbcourt, B. 1979.A manual of New Guinea legumes. Papua New Guinea Office of Forests, 
Bot. Bull. 11:1-645. 

Wetzen, P.C. VAN. 1994. Guioa. In: Sapindaceae. Flora Malesiana ser. 1, 11(3):548-598. 

Wide, W.JJ.O. be. 1985. A new account of the genus Horsfieldia (Myristicaceae). Part 2. 
Gard. Bull. Straits Settlem. 38:55-144. 


64 SIDA 19(1) 


BOOK REVIEW 


Peter D. StiLING. 1999. Ecology: Theories and applications, 3rd ed. (ISBN 0-13-915653-4, 
hbk.) Prentice-Hall, Upper Saddle River, New Jersey 07458, U.S.A. 638 pp., b/w and 
color, and line drawings. 


Ecology: theories and applications is a well-written, comprehensive overview of basic key concepts 
and theories in the fields of ecology and conservation biology. Headings for each of the subsec- 
tions are statements that summarize the key ideas and provide a helpful outline of the major con- 
cepts of the chapter. Tables, photographs, and diagrams throughout the book are well presented 
and effectively summarize or illustrate important concepts. The book is organized in a fairly tradi- 
tion format, covering a wide range of topics. It begins with a discussion of the field of ecology in 
general, then moves mlouan evolutionary cele Ee iauore aoe Laat ecology (in- 
cluding a nic f abiotic factors), c gy, and ecosystem ecology. Stiling 
also includes a Gietsh of the relatively young ‘field tan eon ate Within each section 
are presentations of modern-day applications relating to the theories that support them. These 
sections help the student recognize the significant role that the science of ecology has to soe in 
decision-making and policy at local to global scales. The book is a great resource for university 
students and others interested in the theories forming the foundation for the ein sciences. 
—Charlotte Bryant. 


ERAGROSTIS ANCASHENSIS (POACEAE: CHLORIDOIDEAE), 
A NEW SPECIES FROM ANCASH, PERU 


Paul M. Peterson 


Department of Botan 
National Museum of Natural History 


Smithsonian Institution 


Washington, DC 20560-0166, U.S.A. 


Nancy Refulio Rodriguez and Oscar Tovar 


Historiq Na / 


NA di 
Universidad Nacional Mayor de San Marcos 
Lima, 14-0434, PERU 


ABSTRACT 


Eragrostis ancashensis PM. Peterson, Refulio & Tovar, sp. nov., is described and illustrated. The new 
species occurs on steep rocky slopes in three distinct locations in Departamento Ancash: near the 
southern end of the Cordillera Blanca, approximately 20 km E oF Raquia, near eave: ani near 
Bambas, just north of the Cordillera. The new spec ely allied 

Hitchc. but differs e its shorter culms (26-84 cm long); shorter a narrower leaf Blades [(6-)10- 
22(-26) cm long x 1-2.5(-3.0) mm wide]; shorter panicles [10-20(-38) cm long]; shorter branches 
(1.5-11 cm long) that are widely spreading; ovate spikelets (3-6.1 mm long x 2-4.5 mm wide) with 
a long ciliate (the hairs up to 1.5 mm long) and flattened rachilla; veins of the glumes, lemmas, and 
paleas usually with minute, whitish, raised glands; broadly ovate lemmas 2—3.2 mm long; and an- 
thers 1.2-2.0 mm lon 


RESUMEN 


Eragrostis ancashensis PM.Peterson, Refulio & Tovar, sp.nov., es pumualbie e ea ege eal nueva especie 
habita en pendientes de suelos rocosos, en | de Ancash: 
cerca del extremo sur de Cordillera Blanca, aproximadamente a 20 km E de: Raquia; cerca a Huaylas 
y cerca a Bambas,al norte de la Cordillera Blanca. Esta nueva especie esta fuertemente relacionada 
con Eragrostis magna Hitchc., pero difiere de ella por sus culmos mas cortos (26-84 cm de largo); 
sus laminas foliares mas cortas y menos anchas [(6—-)10-22(-26) cm de largo 3 1-2.5(-3.0) mm de 
acne); panieules) mas cortas i 20(-—38) cm bie largo];ramas de la panicula mas cortas (1.5-11 cm 

3 mm de largo 3 2-4.5 mm de ancho), raquila peas 
y yaleda ene hasta 1 Sa mm de largo); nervios del las glurmas, lemas y Palege usualamente ¢ 
glandulas Coe y blanquecinas;lemas anchamente ovadas 2-3.2 mm de largo;y anteras 1 Be 
2.0 mm de Oo. 


While making determinations of Peruvian material using the treatments of Tovar (1993), 
Renvoize (1998), and Laegaard and Peterson (2000), the first and second authors recog- 
nized the unique features of three grass collections. These specimens distinguished 
by their dark green and plumbeous-spotted spikelets with a ciliate, flattened rachilla. 
The new species is clearly a member of subfamily Chloridoideae, tribe Eragrostideae, 


SIDA 19(1): 65-70. 2000 


66 SIDA 19(1) 


subtribe Eragrostidinae (Peterson et al. 1995, 1997). We describe these specimens as a 
new species of Eragrostis,and ascribe the specific epithet to the Departamento Ancash. 


= ancashensis P.M.Peterson, Refulio & Tovar, sp.nov. (Fig. 1). Tyee: PERU. Departamento 
NCASH, Provincia Recuay: Cordillera Blanca, approximately 20 km E of Raquia on Route 02- 
ee road towards Huaraz (10° 8' 55.8" S—77° 19' 48.8" W ), 3000 m, 20 Mar 1997, PM. Peterson 

& N. Refulio Rodriguez 13793 (HoLotype: USMI: isotype: KE MO! NY! RSA! TAES! UC! US! WIS!). 


Ab Eragrostis magna Hitchce. rhizomatibus nullis, culmis 26-84 cm altis, laminis (6-)10—22(—26) cm 
longis 1—2.5(-3.0) mm latis, paniculis 10-20(-38) cm longis ramis 1.5-11 cm longis Sie spiculis 
3-6.1 mm longis 2-4.5 mm latis ovatis, rhachilla Cones elles adios usque 


longis, venis glumarum lemmatum palearum p yue g ti te 12-20 


7 


mm longis recedit. 


Caespitose perennials. Culms 26-84 cm tall, erect, terete near base, glabrous below the 
nodes; nodes mostly basal or 1 rarely 2 above; internodes glabrous. Sheaths 6-16 cm 
long, longer that the lower internode if present, mostly glabrous or with scattered hairs 
near the summit, the hairs up to 1.3 mm long; margins mostly smooth usually with a 
large tuft of hairs near the summit, these hairs up to 3 mm long; collar visible, yellowish. 
Ligules 0.4-0.7 mm long, a line of hairs, sometimes these hairs extending up to 2.5 mm 
long, these breaking off at maturity. Blades (6—-)10-22(—26) cm long, 1-2.5(-3.0) mm wide, 
flat above the ligule to tightly involute above, apically acuminate, usually densely pilose- 
villous near base above and below to sparsely pilose-villous near base and glabrous 
above, the hairs up to 2.5 mm long. Panicles 10-20(-38) cm long, 5-15 cm wide, open, 
the loosely flowered branches spreading 20-80 o from the culm axis; inflorescence 
branches mostly 1.5-11 cm long, naked near base, with spreading secondary branches, 
one to three per node; pulvini in the axils of primary and secondary branches villous, the 
hairs up to 5 mm long; pedicels 1.2-6 mm long, delicately spreading, sinuous to flexuous, 
scaberulous. Spikelets 3-6.1 mm long, 2-4.5 mm wide, florets 3-8, ovate, compressed, 
dark green with small plumbeous spots; rachilla flattened, usually densely ciliate along 
the margins, the hairs up to 1.5 mm long; disarticulation with the glumes first then the 
lemmas falling individually leaving the paleas on the rachilla. Glumes 2-2.8 mm long, 
lanceolate to ovate, membranous, shorter than the lower lemma, about equal in length, 
1-veined, keeled, scaberulous along the keel and usually with minute, whitish, raised 
glands; apex acute to acuminate, often mucronate, the mucro up to 0.5 mm long. Lem- 
mas 2-3.2 mm long, broadly ovate, membranous, 3-veined, lateral veins somewhat ob- 
scure, keeled, scaberulous along the keel and near the apex, the veins usually with minute, 
whitish, raised glands; apex acute, often darker than below. Paleas 1.8-3.1 mm long, ellip- 
tic, bowed-out, membranous, the keels usually with minute, whitish, raised glands; apex 
truncate to obtuse, sometimes minutely erose. Lodicules 2, 0.2-0.3 mm long, cuneate, 
fleshy, non-vascularized. Stamens 3, anthers 1.2-2.0 mm long, yellow to purplish at matu- 
rity. Ovaries 0.3-0.5 mm long; styles 2, separate, glabrous, stigmas 2, feathery, white to 
purplish.Caryopses 0.7—0.9 mm long,rectangular-prismatic, deeply grooved on the adaxial 
(ventral) surface, striate, dark reddish brown. 


PETERSON ET AL., A NEW ERAGROSTIS FROM PERU 67 


u 
I Ry 
i i 


IAN 


TR Anica 2090 


Fic. 1.£ragrosti hensis (Pet & Refulio Rodrig 13793).A.Habit. B. Sheath and ligule. C. Inflorescence. D. Spike- 
let. E. Lower glume, dorsal view. F. Upper glume, dorsal view. G. Floret, as viewed from the dorsal side of lemma. H. 
Lemma, ventral view. |. Palea dorsal view. J. Palea, ventral view. K, Stamens, pistil, and lodicules. L. Gynoecium, mature. 
M. Caryopsis, lateral view. N. Caryopsis, ventral view. 


68 SIDA 19(1) 


Phenology.—Flowering in mid to late March through May, with caryopses in June 
(R. Ferreyra 145/77). 

Distribution.—Eragrostis ancashensis is known only from Departamento Ancash in 
three distinct locations: near the southern end of the Cordillera Blanca (type locality in 
Provincia Recuay), near Huaylas (Provincia Huaylas), and near Bambas (Provincia Corongo), 
just north of the Cordillera. Individuals of E.ancashensis can be found growing on steep 
rocky slopes at mid elevations (2200-3220 m) associated with xerophytic plants such as: 
Agave, Commelina, Lupinus, Puya, Vicia, Viguiera, and other shrubby Asteraceae. 


Additional specimens examined: PERU. Departamento Ancash: Provincia Corongo, 7 km NW of 
Yupan on road towards Bambas, 3220 m, 26 Mar 1997, Peterson & Refulio Rodriguez 13915 (US,USM); 
7 km NW of Bambas, 2710 m, 26 Mar 1997, Peterson & Refulio Rodriquez 13919 (US, USM); Provincia 
Huaylas, between Cardz and Huallanca, 2200-2300 m, 2 Jun 1962, R. Ferreyra 14577 (USM); between 
Huaylas and el Callején, 2400-2500 m, 3 Jun 1962, R. Ferreyra 14594 (USM). 


LEAF ANATOMY 


Cross-sectional leaf blade anatomy was determined from hand sections of dry material 
(Peterson & Refulio Rodriguez 13915) on temporary slides. Therefore, an illustration is not 
presented, since the chlorenchyma tissue and parenchyma bundle sheath cells were 
mostly collapsed. 

The blades are typically kranz-C4, PCK-like [phosphoenolpyruvate carboxykinase or 
classical PCK type, defined as centrifugal/evenly distributed photosynthetic carbon re- 
duction (PCR) cell chloroplasts (with grana), XyMS+ (presence of cells between the met- 
axylem vessel elements and laterally adjacent chlorenchymatous tissue, see also 
Hattersley and Watson 1976),and presence of PCR cell wall suberized lamella,in Hattersley 
and Watson's (1992) sense] since the chlorenchyma appears loosely arranged and quite 
regularly is contiguous (not interrupted by a column of colorless cells) between adja- 
cent vascular bundles. The lamina are involute with primary, secondary, and tertiary vas- 
cular bundles decreasing in size. The primary vascular bundles are well differentiated 
into xylem with metaxylem, phloem, and a double bundle sheath (mestome and paren- 
chyma bundle sheath). However, the parenchyma bundle sheath of the primary vascular 
bundles is interrupted on the abaxial and sometimes the adaxial surface by a girder of 
fibers. In secondary vascular bundles only the abaxial parenchyma bundle sheath is in- 
terrupted by a girder of fibers whereas in tertiary bundles the parenchyma bundle sheath 
is contiguous. The ribs are flattened with angled sides (rectangular) and the furrows are 
1/5 to 1/2 as deep as the thickness of the blade adaxially and usually less than 1/5 as 
deep abaxially. The medium vascular bundle structure consists of a simple keel with only 
a single primary vascular bundle. Per blade there are 9-13 primary vascular bundles and 
20-28 secondary and tertiary vascular bundles. There are two or three secondary or ter- 
tiary vascular bundles placed between each primary vascular bundle. The xylem of the 
primary vascular bundles contains two wide metaxylem vessels that are about the same 
size as the parenchyma bundle sheath cells. The mestome, or inner sheath, is always 


PETERSON ET AL., A NEW ERAGROSTIS FROM PERU 69 


Taste 1. Salient features comparing Eragrostis ancashensis with E. magna. 


Characters E. ancashensis E. magna 

Rhiz absent present 

Giln (pane 26-84 cm to 150 cm 

Blades, lengths (6-)10-22(-26) cm 40-60 cm 

Blades, widths 1-2.5(-3.0) mm 3-7 mm 

Inflorescence lengt 10-20(-38) cm 30-50 cm 

Inflorescence a lengths —_1.5-11 cm, spreading 12-20 cm, ascending 

Pedicels aspect sinuous to flexuous, reflexed flexuous, ascending 

Spikelet shape ovate linear-lanceolate 

Spikelet lengths 3-6.1 mm 7-10 mm 

Spikelet widths 2-4.5 mm 2-2.3 mm 

Rachilla vestiture ciliate, hairs up to 1.5 mm ciliate, hairs less than 0.2 mm 
along entire length only at base of lemma 

Rachilla shape strongly flattened mostly terete 

Veins of the glumes, usually present absent 


lemmas, and paleas 
with minute, whitish, 
raised glands 
Lemma lengths 2-3.2 mm 5m 
Lemma shape broadly ovate elliptic 
Anther lengths 1.2-2.0 mm 1.2-1.5mm 


present in the vascular bundles surrounding the xylem and phloem. Chlorenchyma cells 
radiate just outside the parenchyma bundle sheath cells and are often contiguous be- 
tween adjacent bundles forming a loosely radiate arrangement (PCK-like). One to four 
rows of sclerenchyma fibers form the abaxial and adaxial girders which are wide near 
the epidermis and narrow toward the vascular bundle. Sclerenchyma fibers form a nar- 
row and very pointed projection along the margin of blade. 


DISCUSSION 


The new species seems allied to species of Eragrostis subgenus Caesiae Van den Borre 
since the majority of these species are PCK-like and perennial (Van den Borre & Watson 
1994).We suspect that the closest sister to the new species is E. magna (Hitchcock 1927) 
since it shares many features, e.g., dark green and plumbeous-spotted spikelets with a 
ciliate rachilla (reduced to a tuft of hairs below each floret in E magna), very few culm 
nodes (one or two), and rectangular-prismatic caryopses. Eragrostis ancashensis differs 
from E.magna by 15 characteristics (see Table 1), most notably: shorter culms (26-84 cm 
tall); shorter and narrower leaf blades [(6—-)10-22(-26) cm long x 1-2.5(-3.0) mm wide]; 
shorter panicles [10-20(-38) cm long];shorter branches (1.5—11 cm long) that are widely 
spreading; ovate spikelets (3-6.1 mm long x 2-4.5 mm wide) with along ciliate (the hairs 


70 SIDA 19(1) 


up to 1.5 mm long), flattened rachilla; veins of the glumes, lemmas, and paleas usually 
with minute, whitish, raised glands; broadly ovate lemmas 2-3.2 mm long; and anthers 
1.2-2.0 mm long. 

Another possible sister to the new species might be £. macrothyrsa Hack., men- 
tioned by Hitchcock (1927) as having the same aspect. However, F. macrothyrsa differs 
from E.ancashensis by having taller culms (1-1.6 m tall), larger blades (20-45 cm long x 
4-15 mm wide), longer panicles (40-60 cm long), shorter glumes (1.2-2 mm long), shorter 
lemmas (1.5-2 mm long), and shorter anthers (0.6—1 mm long). The distribution of E. 
macrothyrsa is also more sOHEN CHA occurring in Paraguay (type), Brazil, and Bolivia. 

There exists considerable tion among the accessions of F. ancashensis that may 
warrant taxonomic consideration in the future when more collections are available. The 
specimens from 7 km NW of Bambas (Peterson & Refulio Rodriguez 13919) and between 
Caraz and Huallanca (Ferreya 14577) are taller, more robust individuals with less pubes- 
cence (hairs shorter and less dense on the blades, sheath and inflorescence pulvini), 
slightly smaller florets, and shorter hairs present on the rachilla. Both of the collections 
from Provincia Huaylas (Ferreya 14577 & 14594) have less hairs that are shorter on the 
spikelets. 


ACKNOWLEDGMENTS 


Appreciation is extended to Alice R. Tangerini for providing the illustration, Dan H.Nicolson 
for correcting the Latin diagnosis, and Asuncion Cano Echevarria and Maria Isabel La 
Torre Acuy for local support. Jacques Cayouette, Stephan L. Hatch, Barney L. Lipscomb, 
and Harold Robinson are thanked for reviewing the manuscript on short notice. 


REFERENCES 


Hartterstey, PW.and L. Watson. 1976. C4 grasses: an anatomical criterion for distinguishing 
between NADP-malic enzyme species and PCK or NAD-malic enzyme species. Aus- 
tralian J. Bot. 24:297-308. 

and _____.. 1992. Diversification of photosynthesis. Pp. 38-116 in Grass 
evolution and domestication,ed.C. P Chapman.Cambridge University Press: Cambridge. 

HitcHcock, A.S. 1927. The grasses of Ecuador, Peru, and Bolivia. Contr. U.S. Natl. Herb. 24 
(8):291-556. 

LAEGAARD, S.and P.M. Peterson. 2000. 214 (2). Gramineae (Part 2-Chloridoideae) in G. Harling 
& L. Andersson (eds.), Fl. Ecuador # (in press). 

Peterson, PM., R.W. Weaster, and J. Vacbes-Reyna. 1995. Subtribal classification of the New 
World Eragrostideae (Poaceae: Chloridoideae). Sida 16:529-544. 

1997. Genera of New World Eragrostideae (Poaceae: 
Chloridoideae). Sheonian ae Bot. 87:1-50. 

Renvoize, S.A. 1998. Gramineas de Bolivia. Royal Botanic Gardens, Kew. 

Tovar, O. 1993.Las Graminees (Poaceae) del Peru. Ruizia 13:1—-480. 

VAN DEN Borre, A.and L.Watson. 1994. The infrageneric classification of Eragrostis (Poaceae). 
Taxon 43:383-422. 


WOOD AND BARK ANATOMY OF ACHATOCARPACEAE 


Sherwin Carlquist 


Santa Barbara Botanic Garden 
1212 Mission Canyon Road 
Santa Barbara, CA 93105, U.S.A. 


ABSTRACT 


Qualitative and quantitative data are given for Achatocarpus nigricans Triana, A. praecox Griseb., and 

ee spinescens A. Gray. The minute vessel pits, lack of successive cambia, and lack of 

talains or anthocyanins are distinctive features of the family that justify its removal from 

sar eer The ak nature of these features is unclear; libriform fibers are commonly re- 

ed as specialize orders on perforation plates accords with the placement of 

this family in oe Phytolaccineae, however. Quantitative features of vessels accord with the 

concept that Phaulothamnus has the wood of a desert shrub, whereas wood of Achatocarpus is like 
that of a shrub from a chaparral-like habitat 


Key Worbs: Achatocarpaceae, cambial phylogeny, Caryophyllales, ecological wood anatomy, 
Phytolaccaceae, systematic wood anatomy. 


RESUMEN 


Se ofrecen datos cualitativos y cuantitativos sobre los lehos de Achatocarpus nigricans Triana, A. 
praecox Griseb., y Phaulothamnus spinescens A. Gray. Punteaduras diminutas de vasos, seta 
de pigmentos, y deficiencia de cambios sucesivos caracterizan a la familia, y separ 

Achatocar saan de a eshiowsie: El estado es de estas peculiaridades es incierto; las ae 
libriformes se interpr avanzado. Los datos cuantitativos de los vasos indican una 
ecologia ieee a para 5 Phaulotiarnits yuna ee de tipo chaparral seco para Achatocarpus. 


INTRODUCTION 


Earlier systems include Achatocarpaceae (e.g.,Walter 1909) within Phytolaccaceae sensu 
lato. The family was segregated by Heimerl (1934) so as to include one species of 
Phaulothamnus and nine of Achatocarpus, and has been thus recognized by most sub- 
sequent authors (Cronquist & Thorne 1994).The ovary of Achatocarpaceae has two stig- 
mas but is unilocular and matures into a berry. This combination of features does not 
occur in Phytolaccaceae s.l. if Achatocarpaceae are segregated. More significantly, 
Achatocarpaceae are not known to produce either betalains or anthocyanins (Clement 
et al. 1994), whereas all other Phytolaccaceae s.|. contain betalains. All authors place 
Achatocarpaceae within Caryophyllales, but the position within the order is less clear. 
Placement of Achatocarpaceae just outside of suborder Phytolaccineae is currently com- 
monly accepted (Manhart & Rettig 1994; Thorne in Cronquist & Thorne 1994; Behnke 
1997), whereas Brown and Varadarajan (1985) place Achatocarpaceae outside 
Phytolaccaceae sensu stricto but inside Phytolaccaceae s1. 

of uncertainty in placement of Achatocarpaceae renders any 


q 
UCU! 


SIDA 19(1): 71-78. 2000 


72 SIDA 19(1) 


kind of data, including those from wood and bark anatomy, valuable for resolution of the 
phylogenetic relationships of the family. Data on wood and bark of Achatocarpaceae 
have been contributed by Metcalfe and Chalk (1950) and Gibson (1994). Molecular data 
are likely to provide strong evidence also; Caryophyllales are, as yet, relatively sparsely 
sampled with respect to DNA features. 

The stem of species of Achatocarpaceae has a single cambium whereas several 
genera of Phytolaccaceae have successive cambia. Only a single cambium is known in 
the phytolaccaceous genera Lophiocarpus, Microtea, Monococcus, and Trichostigma (these 


genera would fall into Rivinaceae if Phytolaccaceae s.s.is reduced to Anisomeria, Ercilla, 
and Phytolacca). The question of whether successive cambia or a single cambium are 
plesiomorphic or apomorphic in Phytolaccineae and in Caryophyllales as a whole re- 
mains to be resolved. 

The concept of Caryophyllales offered by Cronquist and Thorne (1994) or Behnke 
and Mabry (1994) is used here. Plumbaginaceae and Polygonaceae are considered 
outgroups when the order is so designated (Rodman 1994). However, the data of Will- 
iams et al. (1994) have led a working group (APG 1998) to recognize an expanded 
Caryophyllales in which Plumbaginaceae, Polygonaceae, Tamaricaceae, Droseraceae, 
Nepenthaceae and allied families are included. The Cronquist and Thorne (1994) con- 
cept of Caryophyllales is then termed ‘core Caryophyllales.”Wood and stem anatomy of 
Caryophyllales, when all families have been surveyed, may reflect the new classification 
or may tend to show subgroup patterns. 

The present paper is part of a survey of wood anatomy of Caryophyllales that has 
included Caryophyllaceae (Carlquist 1995), Portulacaceae and Hectorellaceae (Carlquist 
1998a), and Basellaceae (Carlquist 1999a). Among the genera or familial segregates of 
Phytolaccaceae s.l. studied to date are Petiveria and Rivina (Carlquist 1998b), Agdestis 
(Carlquist 1999b), Stegnosperma (Carlquist 1999c), Barbeuia (Carlquist 1999d),and rivinoid 
and phytolaccoid Phytolaccaceae (Carlquist in press). All of the families of Caryophyllales 
in the broad sense (APG 1998) will ultimately be included. 

The two genera of Achatocarpaceae considered here are shrubs to small trees 
(Heimer! 1934). They are native to areas with dry seasons, most markedly so in the habi- 
tats of Phaulothamnus in southern Texas and northern Mexico, less extreme in the habi- 
tats of Achatocarpus, which range from Mexico to Argentina (Walter 1909; Heimer! 1934). 
The relationship of wood anatomy to the ecology of this family is a focus of the present 
paper. 


eae 


MATERIALS AND METHODS 


The collections studied are as follows: Achatocarpus nigricans, Portoviejo, Manabi, Ecua- 
dor (M.Acosta-Solis 11918), USw-0020137 Cane 22 mm in diameter);A. praecox, Tucuman 
Argentina (sample 16 mm in diameter); PI 
22596,POM (sample 5 mm in cee After softening with 4% aqueous ethylene di- 
amine, sections were prepared with a sliding microtome and stained with a safranin-fast 


spinescens Sonora Mexico, Jones 


CARLOQUIST, ACHATOCARPACEAE 73 


green combination. Attempts to locate crystals were made with the use of polarizing 
equipment. Vessel diameter is measured as mean lumen diameter. Means for quantita- 
tive features reported are derived from 25 measurements per feature. Terminology for 
wood features accords with the IAWA Committee on Nomenclature (1964). Vessels per 
group is a mean based on a solitary vessel = 1, a pair of vessels in contact = 2, etc. 


RESULTS 


Wood Anatomy 

Both qualitative and quantitative features are given for A. praecox. Quantitative features 
are given for the two remaining species, but qualitative features are given for them only 
when these differ from the conditions in A. praecox. 

Achatocarpus praecox (Figs. 1-4). Growth rings present but indistinct (Fig. 1). Vessels 
grouped in radial multiples or solitary; mean number of vessels per group, 1.78. Mean 
vessel lumen diameter, 30 um. Mean number of vessels per mm?, 88. Mean vessel ele- 
ment length, 292 um.Mean vessel wall thickness, 4.0 um. Perforation plates nonbordered. 
Perforation plates simple. Lateral wall pitting opposite on ray-vessel interfaces (Fig. 3), 
alternate on other vessel faces (Fig. 4). Lateral wall pits of vessels minute, about 1.5 um in 
diameter, circular in outline or nearly so, with small elliptical pit apertures. Imperforate 
tracheary elements are all libriform fibers with very small simple pits (Fig.4,extreme left). 
Length of libriform fibers,677 um. Mean wall thickness of libriform fibers, 2.5 um. Axial 
parenchyma vasicentric scanty, in strands of four cells (Fig. 3, to left and right of vessel). 
Rays both multiseriate and uniseriate (Fig. 2), the former more abundant. Mean height of 
multiseriate rays, 365 um. Mean width of multiseriate rays, 2.63 cells. Mean height of 
uniseriate rays, 119 um.Multiseriate rays composed of procumbent cells (Fig. 3, left; Fig. 4) 
except for tip cells (Fig. 4, lower left), which are square or upright. Uniseriate rays com- 
posed of procumbent or upright cells. Ray cell walls lignified, with mostly simple pits. Ray 
cell walls about 2.2 um thick. Wood nonstoried. Crystals absent. Starch not observed. 

Achatocarpus nigricans. Mean number of vessels per group, 1.85.Mean vessel diam- 
eter, 37 um. Mean number of vessels per mm’, 17. Mean vessel element length, 273 ym. 
Mean vessel wall thickness, 2.2 um. Mean vessel pit diameter, 1.7 um.Mean libriform fiber 
length,670 um. Mean libriform fiber wall thickness, 1.5 um.Mean multiseriate ray height, 
330 um. Mean width of multiseriate rays, 3.2 cells. Ray cell wall thickness about 1.1 um. 
Many vessels filled with amorphous yellow deposits. Libriform fibers commonly filled 
with amorphous yellow or dark deposits. 

Phaulothamnus spinescens (Figs.5—7).Vessels predominantly in radial groups or soli- 
tary (Fig. 5). Mean number of vessels per group, 2.78. Mean vessel diameter, 21 um.Mean 
number of vessels per mm?, 347. Mean vessel element length, 354 um. Mean vessel wall 
thickness, 2.1 um.Mean vessel pit diameter, 1.8 um. Mean libriform fiber length, 549 um. 
Mean libriform fiber wall thickness, 1.1 um. Axial parenchyma is in strands of two cells. 
Uniseriate rays more common than multiseriate rays (Fig.6). Mean height of multiseriate 
rays, 302 um. Mean width of multiseriate rays, 2.0 um.Mean height of uniseriate rays, 229 


SIDA 19(1) 


ews Pantie rey 
a ese 


aay 


t 


7% 


qs 


J 


indicating in- 


’ 


Fics. 1-4. Achatocarpus praecox, wood sections. Fig. 1.T 


Fig.3.R 


Fig. 2. Ti 


J 


J 


tip of biseriate ray, 


.Tangential section 


|.Fig.4 


’ 
t 


I 


J 


scale below Fig. 3 


); Figs. 3-4, 


= 10 pm 


left; alternate minute pits on vessel. Figs. 1-2, scale below Fig. 1 (divisions 


(divisions 


) 


=10 pm 


CARLQUIST, ANATOMY OF ACHATOCARPACEAE 75 


im. Upright ray cells common, but procumbent ray cells also present (Fig. 7). Bordered 
pits common on tangentially oriented ray cell walls (Fig. 7). Ray cell wall thickness about 
1.1 um. Amorphous deposits present in ray cells and in libriform fibers (Fig. 7). 


Bark 
The sections of Phaulothamnus spinescens (Fig. 8) were unusually good and showed all 
regions of the bark clearly. The phellem cells (Fig. 8, top) contain dark-staining amor- 
phous deposits. Several layers of phelloderm are present; cells of these layers are all thick 
walled sclereids (Fig. 8). Outer cortex composed of tangentially widened parenchyma 
cells with nonlignified cell walls. Inner cortex composed of a continuous cylinder of thick- 
walled sclereids. Scattered fibers present in older secondary phloem (Fig. 8). 

The bark of A. praecox is similar to that of P spinescens.|In A.praecox, however, both an 
outer and an inner cylinder of thick walled cortical sclereids are present. The more com- 
plex bark of A. praecox may be related to large diameter of the sample studied here. 


DISCUSSION AND CONCLUSIONS 


Of all families once included in Phytolaccaceae, the family most universally segregated 
is Achatocarpaceae. In fact, the molecular results of Manhart and Rettig (1994) and the 
cladistic and phenetic studies of Rodman (1994) showed that Achatocarpaceae are not, 
in most analyses, a sister group of Phytolaccaceae, and might even be in a near-basal 
position in Caryophyllales (as defined by Behnke & Mabry 1994). Achatocarpaceae are 
not known to contain either betalains or anthocyanins (Clement et al. 1994); this is also 
true of Barbeuia. In two caryophyllalean families, Caryophyllaceae and Molluginaceae, 
anthocyanins are present but betalains are absent. 

The data from wood anatomy suggest that Achatocarpaceae belong to 
Caryophyllales: absence of borders on perforation plates characterizes Achatocarpaceae 
and also most families of Caryophyllales investigated thus far (Carlquist 1998b, 1999b, 
1999c, in press). Bark data are not available yet for a large enough number of 
caryophyllalean genera so that they can be presented as evidence for the familial com- 
position of Caryophyllales or the position of Achatocarpaceae within the Caryophyllales. 

The libriform fibers of Achatocarpaceae are a specialized feature in dicotyledons 
according to traditional criteria (Metcalfe & Chalk 1950, p.xlv), whereas tracheids, shown 
by Metcalfe and Chalk (I.c.) to be more primitive, occur in families often claimed to oc- 
cupy near-basal positions in the order: Caryophyllaceae (Carlquist 1995), Stegno- 
spermataceae (Carlquist 1999c), and Barbeuiaceae (Carlquist 1999d). All of these fami- 
lies, however, have successive cambia (Caryophyllaceae only in some genera), whereas 
Achatocarpaceae does not. Whether absence of betalains and absence of successive 
cambia are plesiomorphic or apomorphic is uncertain. 

The presence of very small pits on vessels of Achatocarpaceae is a feature not re- 
ported elsewhere in Caryophyllales (Metcalfe & Chalk 1950; Gibson 1994). The minute 
pits are indicative of the distinctiveness of Achatocarpaceae as a family. 

The rays of Achatocarpus can be characterized as Heterogeneous Type IIB, transi- 


SIDA 19(1) 


a oes = 


J 


1(5—7) and hark (8). Fig.5.T 


& Ph 


Fics, 5— 


/ 


Fig. 8. Transection, phellem at top, 


, 


.Fig. 7. Radial section, procumbent ray 


cells at top 


Fias.5.6.8, scale below Fig.5 (divisions 


7 


2 


be J 


7 


= 10pm); 


J 


be J 


£ 


n 
7 


¢ 


Fig. 7, scale below Fig. 3. 


CARLQUIST, F ACHATOCARPACEAE 7 


tional to Homogeneous Type | (Kribs 1935; Carlquist 1988). The predominance of upright 
ray cells in the Phaulothamnus specimen studied is related to the small diameter of that 
specimen and is indicative of a juvenile condition (see Carlquist 1988) and not phyloge- 
netically different from the conditions shown by the comparatively larger Achatocarpus 
specimens, which exhibit a rather more mature pattern. 

The moderate to high degree of vessel grouping in the three species is indicative 
of moderate to marked xeromorphy (Carlquist 1984). This is independently evident in 
the Mesomorphy Ratio (vessel diameter times vessel element length divided by num- 
ber of vessels per mm?),a convenient expression of both conductive safety and conduc- 
tive efficiency. The values for this ratio are: A. nigricans, 59; A. praecox, 99; P spinescens, 21 
The desert or near-desert habitats of Phaulothamnus correlate with the low values for 
that species. Southern California desert shrubs as a group have a Mesomorphy Ratio of 
20.9 (Carlquist & Hoekman 1985). The higher values for Achatocarpus are close to the 
values for southern Californian chaparral shrubs as a group, 66.7, or southern Californian 
coastal sage shrubs, 80.7 (Carlquist & Hoekman 1985). 


REFERENCES 


APG (AnalosperM PHYLOGENY Group). 1998. An ordinal classification for the families of flower- 
ing plants. Ann. Missouri Bot. Gard. 85:53 1-553. 
BeHnke, H.-D. 1997. Sarcobataceae—a new family of Caryophyllales. Taxon 46:495-507. 
and T.J. Masry (eds). 1994. Caryophyllales. Evolution and sytematics. Springer 
Verlag, Berlin & Heidelberg. 
Brown, G.K., and FS. VaraparaJan. 1985. Studies in Caryophyllales 1: Re-evaluation of classi- 
fication of Phytolaccaceae s.|. Syst. Bot. 10:49-63. 
Car.auist, S.1984.Vessel grouping in dicotyledon woods: Significance and relationship to 
imperforate tracheary elements. Aliso 10:505-525. 
. 1988. Comparative wood anatomy. Springer Verlag, Berlin & Heidelberg. 
. 1995.Wood anatomy of Caryophyllaceae: Ecological, habital, systematic, and 
phylogenetic implications. Aliso 14:1-17. 
98a.Wood anatomy of Portulacaceae and Hectorellaceae: Ecological, habital, 
and systematic implications. Aliso 16:137-153. 
. 1998b.Wood and stem anatomy of Petiveria and Rivina (Caryophyllales): Sys- 
tematic implications. IAWA J. 19:383-391. 
. 1999a.Wood, stem, and root anatomy of Basellaceae, with relation to system- 
atics and cambial variants. Flora 194:1-12. 
. 1999b.Wood anatomy of Agdestis (Caryophyllales): Systematic position and 
nature of the successive cambia. Aliso 18:35-43. 
. 1999c.Wood and stem anatomy of Stegnosperma (Caryophyllales): Phyloge- 
netic relationships; nature of lateral meristems and successive cambial activity.IAWA J. 
20:149-163 


78 SIDA 19(1) 


__,«1999d. Wood anatomy, stem anatomy, and cambial activity of Barbeuia 
(Caryophyllales). [AWA J. 20:431-440. 

eee I ress): woe and stem anatomy of phytolaccoid and rivinoid 
Phytolaccaceae (Caryoy les):ecology, systematics, nature of successive cambia. Aliso 

_and D.A. Hoekman. 1985. Ecological wood anatomy of the woody southern 
California flora. IAWA Bull. n.s., 6:319-347. 

Crement, J.S., TJ. Masry, H. Wycer, and A.S. Dreioine. 1994. In: H.-D. Behnke and TJ. Mabry, eds. 
Caryophyllales. Evolution and systematics. Springer Verlag, Berlin & Heidelberg. Pp. 
247-261. 

Cronauist, A.and R.F. THorne. 1994. Nomenclatural and taxonomic history. In: H-D. Behnke 
and TJ. Mabry, eds. Caryophyllales. Evolution and systematics. Springer Verlag, Berlin & 
Heidelberg. Pp. 5-25. 

Gigson, A.C. 1994. Vascular tissues. In: H.-D. Behnke and TJ. Mabry, eds.Caryophyllales. Evo- 
lution and systematics. Springer Verlag, Berlin & Heidelberg. Pp. 45-74. 

Heimert, A. 1934. Achatocarpaceae.In:A.Engler & H.Harms, Die naturlichen Pflanzenfamilien, 
ed. 2, 16c:174-178.Verlag von Wilhelm Engelmann, Leipzig. 

law COMMITTEE ON NomenctaTure. 1964. Multilingual glossary of terms used in wood anatomy. 
Verlagsbuchanstalt Konkordia, Winterthur, Switzerland 

Kriss, D.A. 1935. Salient lines of structural specialization in the wood rays of dicotyledons. 
Bot. Gaz. 96:547-55/7, 

Manuaet, J.R., and J. H. Rettic. 1994. Gene sequence data. In: H.-D. Behnke and T.J. Mabry, 
eds. Caryophyllales. Evolution and systematics. Springer Verlag, Berlin & Heildelberg. 
Pp, 235-246. 

Mercatre, C.R. and L. CHatk. 1950. Anatomy of the dicotyledons. Clarendon Press, Oxford. 

Ropman, J. E. 1994. Cladistic and phenetic studies. In: H.-D. Behnke and T.J. Mabry, eds. 
Caryophyllales. Evolution and systematics. Springer Verlag, Berlin & Heidelberg. Pp. 


Watter, H. 1909. Phytolaccaceae. Das Pflanzenreich IV(83):1-154. 
Wictiams, S.E., V.A. Acgert, and M.W. Cuase. 1994. Relationships of Droseraceae: A cladistic 
analysis of rocL sequence and morphological data. Amer. J. Bot.81:1027-1037. 


BATOPILASIA (ASTERACEAE: ASTEREAE), 
A NEW GENUS FROM CHIHUAHUA, MEXICO 


Guy L.Nesom 


North Carolina Botanical Garden 
Coker Hall CB 3280 


University of North Caro 


olin 
Chapel Hill, NC 27599, U. a 
Richard D. Noyes 


Plant Genome Mapping Laboratory 
Riverbend Research Center, Room 16? 
University of Georgia 


Athens, GA 30602, U.S.A. 


ABSTRACT 
The Mexican species byei ee nese is une: a aa and subtribe 
Conyzinae and treated as the monotypic g p gen nd species 
Batopilasia byei aneees & Nesom ) Nesom & Noyes, comb. nov. B ilasi lea ees 


cially resembles the phyletically isolated North American genera Boltonia and Chloracantha in 
morphology. DNA studies indicate that Batopilasia and Boltonia are closely related and suggest 
that Boltonia is closely related to Chloracantha. The new genus apparently is restricted to the area 
of the Barranca del Cobre in southwestern Chihuahua, Mexico. A technical description, illustration 
and distribution map are provided. 


RESUMEN 


La especie mexicana Erigeron byei Sundberg & Nesom se excluye de EFrigeron y de la subtribu 
Conyzinae y se trata como género monotipico Batopilasia Nesom & Noyes, gen. nov., y especie 
Batopilasia byei (Sundberg & Nesom ) Nesom & Noyes, comb. nov. Batopilasia byei se parece al 
menos superficialmente en su morfologia a los géneros norteamericanos filéticamente aislados 
Boltonia y Chloracantha. Los estudios de DNA indican que Batopilasia y Boltonia estan muy 
relacionados y sigieien cue Bonenla este muy reecionaas con Chloracantha. El nuevo género 
Cobre en el suroeste de Chihuahua, México 
Ge ofrecen una descripcion técnica, una ilustracion y un mapa de distribucidn. 


In the original description of Erigeron byei (Sundberg & Nesom 1990), it was regarded as 
most similar to E. ortegae Blake (= Aster spinosus Benth.) and E. oxyphyllus Greene, these 
three species constituting Erigeron sect. Spinosi (E.ortegae, the type). A number of essen- 
tial differences have since been recognized between E. oxyphyllus and E.ortegae, and the 
latter has been segregated as the monotypic genus Chloracantha (Nesom et al. 1991; 
Sundberg 1991). Frigeron — has been treated as a member of the primarily Cali- 
(Erigeron sect. Linearifolii, Nesom 1992 = sect. Pycnophyllum; 
see Nesom & Noyes 1999) NeconE: et al. (1991) noted that although “the evolutionary 
affinities of E. byei may yet prove to lie with [Chloracanthal, outside of Erigeron, there is 


fornian E. foliosus Nutt.group 


SIDA 19(1): 79-84, 2000 


80 SIDA 19(1) 


insufficient evidence at present..."to justify its exclusion, and E. byei has been maintained 
in an isolated position within Erigeron. Recent study of molecular variation in the tribe 
Astereae (Noyes & Rieseberg 1999), however, provides evidence that the origin of E. bye 
lies outside the phylogenetic nexus of Frigeron and Conyza and their closest relatives. 


Relationship to Erigeron and Conyzinae 
Erigeron byei was originally placed in Frigeron because of its perceived resemblance to E. 
ortegae and E. oxyphyllus, but after the recognition of the infra-generic relationship of 
the latter (within Erigeron sect. Pycnophyllum) and the transfer of the former to 
Chloracantha, E. byei has no close similarity to any other species of Erigeron and does not 
fit into any previously circumscribed section (Nesom 1989). The small, few-flowered heads 
and the subterete, multinerved cypselas, in particular, are unusual in Erigeron, although 
no single morphological character is sufficient to exclude the species from the genus. 
Molecular studies by Noyes and Rieseberg (1999) and Noyes (2000) include repre- 
sentative species of Erigeron and close relatives (63 Erigeron, representing 20 sections, 6 
Conyza; 2 Aphanostephus; 1 each of Apopyros, Neja,and Hysterionica) and delimit a mono- 
phyletic group that corresponds to the Conyzinae of Nesom (1994), with three excep- 
tions: (1) the North American genus Aphanostephus arose from within the group, appar- 
ently ina sister relationship to Erigeron bellioides DC. (and presumably its close relatives), 
a species native to the Carribean Islands—these two groups in a sister relationship with 
a group of species including £. quercifolius Lam.; (2) African species of Conyza are placed 
within subtribe Grangeinae rather than Conyzinae; and (3) E. byei is related to species 
outside of the Conyzinae. As so delimited, the species and genera of Conyzinae encom- 
pass considerable morphological diversity, but £. byei is the only New World species to 
be excluded. 


Relationship to Chloracantha and Boltonia 

The hypothesis that Erigeron byei is closely related to Chloracantha was based on their 
shared rhizomatous habit, cauline leaves much reduced in size and distribution, persis- 
tently green-glabrate stems and leaves, erect buds, distinctly similar phyllaries (thin-her- 
baceous, apically rounded to obtuse, with three, prominent, orange-resinous nerves),and 
deltate collecting appendages of the disc corolla style branches. Boltonia also shares 
this combination of habital and capitular features. 

In the Noyes and Rieseberg (1999) analysis, Erigeron byei is positioned as the sister 
genus to Boltonia and this pair of taxa is then most closely related to Symphyotrichum, 
Oreostemma,and genera of subtribe Machaerantherinae. Chloracantha was not included 
in their analysis, but phylogenetic proximity between Boltonia and Chloracantha had 
earlier been suggested by cpDNA analyses of Astereae—in the context of broader sam- 
pling, Morgan (1990) observed that Chloracantha |s most closely related to the 
goldenaster group (subtribe Chrysopsidinae), while Suh (1989) observed the same rela- 
tionship between Boltonia and the goldenasters. A close relationship between Boltonia 
and goldenasters, based on cpDNA evidence, is not corroborated by ITS sequence data 


NESOM AND NOYES, BATOPILASIA, 81 


from nuclear ribosomal DNA (Noyes & Rieseberg 1999), but the suggestion remains that 
Boltonia and Chloracantha are closely related. A long-standing hypothesis of close rela- 
tionship between Boltonia and the Asian genus Kalimeris was rejected by Gu and Hoch 
(1997) on morphological grounds and by Noyes and Rieseberg (1999), based on mo- 
lecular evidence. 

Boltonia remains a genus strongly isolated in morphology (see Cronquist 1980 and 
Anderson 1987 for taxonomic summaries), differing from Frigeron byei in having mostly 
cauline leaves, conical or convex receptacles, phyllari 
short-tubed disc corollas, cypselas broadly elliptic to elliptic-obovate, strongly flattened 
and broadly winged or unwinged, with a nerve at each margin, these often with con- 
spicuous, orangish oil ducts,and pappus absent or short-coronate, also with several small 
bristles and 2(-4) much longer, thickened, barbellate awns. 

Chloracantha differs from Erigeron byei primarily in vegetative features—taller stat- 
ure, thick rhizomes, perennial and lignescent stems with thorns and axillary buds, lack of 
persistent basal leaves—but also in larger heads and greater number of flowers. The 
similarities of Erigeron byei and Chloracantha in their subterete, 4—5-nerved cypselas and 
pappus of barbellate bristles are probably primitive (plesiomorphic), compared to the 
highly derived (apomorphic) fruiting features of Boltonia. 

In summary, although the divergent taxonomic position of Erigeron byei was sus- 
pected at its original description, based on morphology, its relatively generalized fea- 
tures make it difficult to place the species. In overall morphology, however, E. byei is nota- 
bly similar to the genus Chloracantha, and molecular evidence indicates that it is closely 
related to Boltonia and perhaps Chloracantha. On the basis of its morphological isola- 
tion within Erigeron and its unambigous molecular-phylogenetic placement outside of 
the Conyzinae, we treat £. byei as a monotypic genus. 


s with a thick, raised, orangish midrib, 


Batopilasia Nesom & Noyes, gen.nov. Tver:8 ilasia byei (N & Sundberg) N & Noy 


A Chloracanthae similis vestimento fere glabro, foliis caulinis amplitudine valde redactis, gemmis 
erectis, phyllariis tenui-herbaceis nervis tribus aurantiacis, flosculis radii laminis albis brevibus 
circinatis, flosculis hea rami eye aie eneicbus deltatis, et acheniis 4-5-nervatis sed differt statura 

multo minore, h tenuibus vel ramis caudicis rhizomiformibus, foliis 
ear ce came in rosulis coarctatis, sas minoribus, et flosculis radii paucioribus. Ab 
Erigeronte Conyza et affinibus late separatus datis molecularibus 


Batopilasia byei Nesom & Noyes, comb. nov. (Fig. 1) Basionyu:Erigeron byei Sundberg & Nesom, 
Phytologia 69:278. 199 


Perennial, caespitose, glabrous herbs from a system of thin, lignescent, rhizome-like cau- 
dex branches. Stems 7-20 cm tall, usually with 1-2 ascending branches near midstem. 
Basal leaves persistent in rosettes, sessile, narrowly elliptic-oblanceolate, 8-26 mm long, 

—2 mm wide, entire, 1-nerved or faintly 3-nerved, with a mucronulate apex, the cauline 
leaves few and sharply reduced in size to linear bracts 1-4 mm long. Heads 5-8 mm 
wide, erect in bud, solitary on nearly naked peduncles 5-10 cm long; phyllaries in 3-4 
strongly graduated series, elliptic-oblanceolate with obtuse to acute apices, thin-herba- 


82 SIDA 19(1) 


ceous with scarious margins and 1-3, filiform, orange-resinous nerves, sometimes purple 
tinged at least on the upper half, the inner series 4-5 mm long, 0.5-0.8 mm wide; recep- 
tacles low-convex. Ray flowers 9-18, fertile, the corollas white, drying white or purplish, 
5-7 mm long, the lamina 1.4-2 mm wide, coiling at the tips. Disc flowers 24-29, fertile, 
the corollas 3.5-4.2 mm long, not inflated or indurated, lobes triangular-deltate, erect; 
style branches with deltate collecting appendages 0.1—0.2 mm long. Cypselas sparsely 
strigose, cylindrical and terete to slightly flattened, 1.8-2.5 mm long, 0.4-0.5 mm wide, 
with (2-)4(-5) thin, orange nerves; pappus of 15-27 barbellate bristles 2.9-3.8 mm long 
and a few outer setae 0.1-0.5 mm long. Chromosome number, 2n = 18 (Sundberg & 
Nesom 1990). 

The genus is named for the region of southwestern Chihuahua to which it appar- 
ently is restricted. The known populations of Batopilasia (as cited in Sundberg & Nesom 
1990) occur in Municipio Batopilas, within a radius of about 25-30 kilometers, south of 
the town of Creel and in the general area of the Barranca del Cobre (Fig. 2). The plants 
grow on steep rocky slopes, commonly in rock crevices and ledges in arroyos and can- 
yons, in pine-oak woodlands at 2000-2400 meters elevation, flowering May through July. 
The epithet commemorates Robert Bye, botanist and ethnobotanist who has had a long- 
standing and active interest in the flora of Chihuahua. 


NESOM AND NOYES, BATOPILASIA 83 


. Texas 

Chihuahua 

A 
sf Batopilasia bye 
100 km 
Seo. A 
Fic. 2. Geographic distribution of Batopilasia byei. 
ACKNOWLEDGMENTS 


We appreciate review comments from Don Pinkava and permission from Mike Warnock 
(publisher of Phytologia) to republish Linda Vorobik’s illustration of Erigeron byei.| am 
grateful for help with publications costs provided by the Mouzon and Mouzon Fund of 
the University of North Carolina Herbarium (NCU). 


84 SIDA 19(1) 


REFERENCES 


ANbeRSON, L.C. 1987. Boltonia apalachicolensis (Asteraceae): a new species from Florida. 
Syst. Bot. 12:133-138. 

Cronauist, A. 1980. Boltonia. In: Vascular flora of the southeastern United States, Vol. |. 
Asteraceae. Univ. of North Carolina Press, Chapel Hill. Pp. 168-169. 

Gu, H. and PC. Hoc. 1997. Systematics of Kalimeris (Asteraceae: Astereae). Ann. Missouri 
Bot. Gard. 84:762-814. 

Morcan, D.R. 1990.A systematic study of Machaeranthera (Asteraceae) and related groups 
using restriction site analysis of chloroplast DNA and a taxonomic revision of 
Machaeranthera section Psilactis. Ph.D. dissertation, Univ. of Texas, Austin. 

Nesom, G.L. 1989. Infrageneric taxonomy of New World Erigeron (Compositae: Astereae). 
Phytologia 67:67—93. 

Nesom, G.L. 1992. Revision of Erigeron sect. Linearifolii (Asteraceae: Astereae). Phytologia 
72:157-208. 

Nesom, G.L. 1994. Subtribal classification of the Astereae (Asteraceae). Phytologia 76:193- 


Nesom, G.L., ¥. SuH, D.R. Morcan, S.D. Sunpeerc, and B.B. Simpson. 1991. Chloracantha, a new 
genus of North American Astereae (Asteraceae). Phytologia 70:371-380. 

Nesom, G.L.and R.D. Noyes. 1999. Notes on sectional delimitations in Erigeron (Asteraceae: 
Astereae). Sida 18:1161-1165. 

Noyes, R.D. 2000. Biogeographical and evolutionary insights on Erigeron and allies 
(Asteraceae) from ITS sequence data. Pl. Syst. Evol. 220:93-114. 

Noyes, R.D. and L.H. Rieseserc. 1999. ITS sequence data support a single origin of North 
American Astereae (Asteraceae) and reflect deep geographic divisions in Aster s.|. Amer. 
J.Bot.86:398-41 2. 

Sux, Y. 1989. Phylogenetic studies of North American Astereae (Asteraceae) based on 
chloroplast DNA. Ph.D. dissertation, Univ. of Texas, Austin. 

Sunpeere, S.D. 1991. Infraspecific classification of Chloracantha spinosa (Benth.) Nesom 
(Asteraceae) Astereae. Phytologia 70:382-391. 

Sunpeere, $.D.and G.L. Nesom. 1990. A new species of Erigeron (Asteraceae: Astereae) from 
Chihuahua, Mexico. Phytologia 69:278-281. 


A NEW COMBINATION IN 
ARCHIBACCHARIS (ASTERACEAE: ASTEREAE) 


Guy L.Nesom 


North Carolina Botanical Garden 
Coker Hall CB 3280 


= 


Univers it of North Cd 


rolin 
Chapel Hill, NC 27599-3280 U. ‘ A. 


ABSTRACT 


A new combination in Archibaccharis (Asteraceae: Astereae) is made based on an earlier name by 
FW. Klatt published in 1884. 


RESUMEN 


Se hace una nueva combinacion en Archibaccharis (Asteraceae: Astereae) basada en un nombre 
anterior de F.W.Klatt publicado en 1884 


Study of a specimen collected in Mexico by F.M. Liebmann shows that a name for an 
accepted species of Archibaccharis was published (as a species of Baccharis) by F.W. Klatt 
nearly 60 years earlier than the name now in use, A. caloneura S.F. Blake. The earlier name 
is brought into current nomenclature by the following combination. 


ac gale trichotoma (Klatt) Nesom, comb.nov.Baccharis trichotoma Klatt, Leopoldina 
0:91. 1884. Tyee: [Locality not specified on specimen], Liebmann 55,"Liebm.PI. Mus. Nr. 10.994’ 


es C, scanned image!). 
Archibaccharis caloneura S.F.Blake, Proc. Biol. Soc.Washington 55:117. 1942. Type: MEXICO. 
Oaxaca: Mt. Zempoaltépetl, lower slopes, 19-27 Feb 1937, WH. Camp 2701 (HoLoryee: 
NY, fichel; isorvee: US-photo and fragments!). 


Klatt’s citation of the type of Baccharis trichotoma was "Mexico, leg.Liebmann Nr.55.Herb. 
Hort. Bot. Hafn.” Judging from the handwriting (by comparison with Burdet 1978), the 
type specimen apparently was first identified by Schultz-Bipontinus as "Baccharis elegans 
HBK.? var."The identification as“Baccharis trichotoma FW Klatt” presumably was later and 
by Klatt himself. The provenance of the collection is not noted on the sheet or label, but 
other details of the specimen, including the earlier tentative identification by Schultz, 
match the description and protologue published by Klatt. The nature of the three-parted 
division alluded to by the epithet is not clear. 

Archibaccharis trichotoma is known to occur from central Oaxaca to Chiapas, Mexico, 
where it grows in pine-oak, pine, evergreen cloud forests, often with oaks, 1750-2900 
meters elevation, flowering in October through April Jackson 1975; Nesom in prep). 
Liebmann travelled in Mexico and Cuba during the years 1840-1843 (McVaugh 1987; 
University of Copenhagen Botanical Museum 1999); he collected in 1842 (May through 


SIDA 19(1): 85-86. 2000 


86 SIDA 19(1) 


early October) at numerous localities in central regions of Oxaca (McVaugh 1987), pre- 
sumably where the type specimen of A. trichotoma was gathered. 

The leaves of Archibaccharis trichotoma are ovate-lanceolate with long-acuminate 
apices, petiolate, thick and glabrous with prominently raised venation, mucronulate on 
the distal 2/3-1/2 of the margins, the stems are glabrous and evidently dark-colored, 
and the capitulescence is compact and primarily terminal, raised above the level of the 
upper leaves. The type specimen bears staminate heads. Archibaccharis trichotoma is 
placed within Archibaccharis sect. Stricta Nesom (Nesom 1991) with other species of 
southern Mexico and Central America that have strictly erect stems and coriaceous leaves. 

This taxonomic problem was brought to light through study of a specimen “photo- 
graphed by the Field Museum of Natural History”and distributed as the type of Baccharis 
trichotoma Klatt. The label of the specimen (W 33218, MO-photo!) identifies it as B. 
trichotoma Klatt and indicates only that the collection was made in “Mexico" by Galeotti, 
with no other specific collection information visible. H.-G. Galeotti collected in Oaxaca, 
including regions in the central part of the state where Archibaccharis trichotoma is known 
to occur, from April 1839 to early 1840 (McVaugh 1978). The two branches on the W 
sheet were taken from pistillate plants that are characteristic of the species as known 
from the type and other collections. Examination of the photo of the Galeotti collection 
(not the type) led to a search for the type, ultimately resulting in the reduction of the 
Blake name to synonymy under the Klatt name. 


ACKNOWLEDGMENTS 


| am grateful to Olof Ryding (©), who located the type, prepared a scanned image of it, 
and sent it to me digitally, the staff of US for help during a visit there, MO for a loan of photo- 
graphs of types to TEX, John Strother for encouragement and comments, and John Pruski 
and Eric Lamont for journal reviews.| am grateful for help with publications costs pro- 
vided by the Mouzon and Mouzon Fund of the University of Carolina Herbarium (NCU). 


REFERENCES 


Burpet, H.M. 1978. Cartulae ad botanicum graphicem. XIll. Candollea 33:365—408. 

Jackson, J.D.1975.A revision of the genus Archibaccharis Heering (Compositae - Astereae). 
Phytologia 32:81-194. 

McVauch, R. 1978. Galeotti’s botanical work in Mexico: The numbering of his collections 
and a brief summary. Contr. Univ. Michigan Herb. 11:291-297. 

McVaucu, R. 1987. Itinerary and gazatteer. In: J.T. Mickel. Liebmann’s Mexican ferns: His itin- 
erary,a translation of his“Mexicos Bregner,”and a reprinting of the original work. Contr. 
New York Bot. Gard. 19:5-38, 

Nesom, G.L. 1991. Two new species of Archibaccharis (Asteraceae: Astereae) from Mexico 
with a reevaluation of sectional groupings in the genus. Phytologia 71:152-159. 

Nesom, G.L.In prep. The Comps of Mexico. A systematic account of the family Asteraceae. 
Vol. 2, Astereae. Phytologia Memoirs. 

UNiversity OF COPENHAGEN BoTANicat Museum. 1999. Website: www.nathimus.ku.dk/bot/ 
botmus.htm. 


REVISION OF LOBELIA SECT. TUPA 
(CAMPANULACEAE: LOBELIOIDEAE) 


Thomas G. Lammers 


ee of Biology and Microbiology 
University of Wisconsin oo 
Oshkosh, WI 54901, 


ABSTRACT 


The classification of Lobelia L. sect. Tupa (G. Don) Benth. was revised following extensive field stud- 

ies and examination of over 2500 specimens in 43 herbaria. Four species are recognized; all are 

ne vaplolds: wih racemose Unilap late flowers, Stet to celia! one Two are branched ae 
mall tr a re 1s, found in the dw 

and xerophytic-shrub ne sclerophyllous vegetation zones of the xeric north: L polyphylla neck & 

Arn.with small wine-purple flowers, ae L excelsa Bonpl.with larger flowers that change color from 

yellow slits Olanide in Pu ue red at aun 6. The other two are large herbaceous or Subic oe 


| aes | iat {fr 
INUTELY foveolate- retic 


found in the deciduous forest vergreen temperate rain forest regions of the mesic < south: red- 
flowered L. tupa L.and pink-flowered L bridgesii Hook. & Arn. The species of each pair are at least 
partly sympatric, and their reproductive isolation (as evidenced by a near total lack of plants with 
intermediate morphology) may be related to differences in pollination syndrome: entomophily in 
L. bridgesit and L. polyphylla, ornithophily in L. tupa and L. excelsa 


RESUMEN 


La clasificacién de Lobelia L. sect. Tupa (G.Don) Benth. se revis6 como continuacién a estudios de 
campo extensivos y el examen de mas de 2500 especimenes de 43 herbarios. Se reconocen cuatro 
especies, todas ellas hexaploides con flores unilabiadas racemosas, endémicas de Chile central. 
Dos c-—O mM P| +>]// AliA { + £ >| | \ ill HI 


rbustos 


t 1 _ 
elipsoides débilmente estriadas, que se encuentran en las zonas de vegetacion de arbustos enanos 
xerofiticos y la zona esclerofila del Ore xérico: L ppoypeyila Hook. = ae con uloes penne oe 
color purpura-vino,y L. excelsa Bonp! 

y naranja en el botdn, hasta rojo en la antesis. Las eras dos son perennes herbaceas grandes o 
sufruticosas no ramosas de tallo hueco, con semillas anchamente elipsoides diminutamente 
foveolado-reticuladas, que se encuentran en las regiones de la pluvisilva caducifolia y la templada 
perennifolia del sur mésico: L. tupa L. de flores rojas y L. bridgesii nooks & Arn. de flores rosas. Las 
especies de cada par son al menos parcialmente simpatricas, y su ductivo (como 
se evidencia por la falta casi total de plantas con morfologia intermedia) ee estar relacionado 

con diferencias en el sindrome de polinizacidn:entomofilia en L. bridgesii y L. polyphylla, y ornitofilia 

nL. tupa y L. excelsa. 


Lobelia L. is the largest genus of Lobelioideae (Campanulaceae), comprising over 400 
species of annual and perennial herbs, shrubs, trees, and giant rosette plants (Lammers 
1993a).It is cosmopolitan in distribution, indigenous to six continents and several island 
groups (e.g., Hawaii, New Zealand, the Antilles). Nearly 38% of the species are African and 
another 29% North American; Asia and Australasia each have about 10% of the species, 
South America 8%, and Polynesia 3%, while only two species occur in Europe (Lammers, 
unpublished data). 


SIDA 19(1): 87-110. 2000 


88 SIDA 19(1) 


The genus was last monographed by Wimmer (1953, 1968), who divided it into 
three subgenera: Lobelia with two sections; Mezleria (C. Presl) E.Wimm., nom. illeg. (cf. 
Lammers 1999) with two sections; and Tupa (G. Don) E.Wimm. with six sections. Further 
taxonomic structure was indicated by dividing many of the sections into subsections 
and other subordinate taxa (cf. Lammers 1993a). 

Wimmer's classification of Lobelia was recently revised by Murata (1995). Though 
based primarily on seed coat morphology (Murata 1992), this revision was also concor- 
dant with other data not available to Wimmer, including chromosome numbers (Lammers 
1993a) and chloroplast DNA (cpDNA) restriction-site analyses (Knox et al. 1993). In this 
classification, Wimmer's three subgenera were maintained (though the illegitimate name 
of the second one should be replaced by subg. /solobus (A. DC.) Y.S. Lian; Lammers 1999). 
However, Wimmer's division of the subgenera into sections was greatly altered. 

Among the sections of subg. Jupa which were remodeled by Murata was sect. Tupa 
(G. Don) Benth. Wimmer (who used the now illegitimate name sect. Eutupa E.Wimm.) 
had construed this section to include 69 diverse species from Asia, Africa, south America, 
and the West Indies. (Within the section, however, Wimmer did segregate the Chilean 
species as “§1. Species chilenses” under the invalid subsectional name “Primanae’ nom. 
nud.) In Murata’s revision, however, all extra-Chilean species were removed to sect. 
Colensoa (Hook. f.) J. Murata or sect. Homochilus A. DC., leaving only a small group of 
species endemic to central Chile in sect. Tupa. 

Thus circumscribed, sect. Tupa is characterized by its combination of robust 
iteroparous (polycarpic) habit; large racemose flowers; unilabiate red, pink, or wine-purple 
non-spurred corolla with monomorphic deflexed lobes coherent at apex; staminal col- 
umn shorter than the corolla and exserted from its dorsal slit; ventral anthers bearded 
apically with tufts of stiff white trichomes; smooth seed coat comprising a single layer of 
cells with long lumina (Type D of Murata 1992, 1995); and hexaploid (n = 21) chromo- 
some number. The latter two features appear to be unique within subg. Jupa (Murata 
1992, 1995; Knox et al. 1993; Lammers 1993a). While hexaploidy is regarded as derived 
within Lobelioideae (Lammers 1993a), the Type D seed coat is considered plesiomorphic 
(Murata 1992, 1995), 

This more restricted circumscription was supported strongly by the phylogenetic 
analyses based on cpDNA data (Knox et al. 1993). In the concensus tree, the species of 
sect. Tupa s. str. formed a monophyletic group that was supported by 10 cpDNA restric- 
tion-site mutations (bootstrap value 99%, decay value >5).The remaining species of subg. 
Tupa examined, including two (L. boninensis Koidz. and L. nicotianifolia Roth ex Schult.) 
that had been included in sect. Tupa by Wimmer, formed its sister-group. These plants 
were all tetraploid (n = 14), so far as known (Lammers 1993a). 


J 


eld 


TAXONOMIC HISTORY 
Plants referable to Lobelia sect. Tupa were first made known to Europeans through the 
activities of the French missionary Louis Feuillée, who visited Chile between 1707 and 
1712.In the published account of his travels (Feuillée 1714), he described and illustrated 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 89 


a highly toxic plant known to the indigenous Mapuche as tupa. Feuillée gave the plant 
the Latin polynomial“Rapuntium spicatum, foliis acutis, vulgo Tupa."In Species plantarum, 
Linnaeus (1753) accepted the species on the sole basis of Feuillée’s description and plate, 
and assigned it to his genus Lobelia. In taking up the vernacular name as the nomen 
triviale, he misspelled it “trapa.” This error was corrected to L. tupa in the next edition 
(Linnaeus 1762). 

In the first half of the nineteenth century, 17 species related to L. tupa were de- 
scribed from Chile. More than one-third of these descriptions (e.g., Sims 1810; Lindley 
1826, 1830;D.Don 1835) were based on plants cultivated from seed in various European 
botanic gardens, and were published in illustrated semi-popular venues such as Curtis’s 
Botanical Magazine. In the latter half of the century, eight more species were described, 
primarily by resident botanist Rodulfo Amando Philippi (1808-1904) in the course of his 
studies of the Chilean flora (e.g., Philiopi 1873, 1895). A number of varieties and forms 
were also described over the years by various workers. Altogether, 35 heterotypic taxa 
referable to Lobelia sect. Tupa have been described, all from Chile and nearly all in the 
nineteenth century. 

Differences of opinion on the circumscription and typification of genera created 
additional combinations in the group. Presl (1836) believed that Rapuntium Mill. was the 
nomenclaturally correct name for most of the species treated under Lobelia by Linnaeus, 
while Kuntze (1891) argued for Dortmanna Hill. George Don (1834) segregated robust 
species of Lobelia with unilabiate corollas as the genus Tupa G. Don, a move supported 
by Candolle (1839) but subsequently reversed by Bentham (1876). These differences of 
opinion (together with changes in rank by various workers over the years) have resulted 
in an additional 61 homotypic combinations based on the original 35 taxa, giving a total 
of 96 validly published names referable to Lobelia sect. Tupa. 

In the most recent flora of Chile, Reiche (1905, 1910) synonymized or excluded many 
of these names, recognizing just six species and six additional heterotypic varieties in 
sect. Tupa. In the most recent monograph of the section (as “$1. Species chilenses’), 
Wimmer (1953, 1968) recognized 20 taxa:seven species plus thirteen additional hetero- 
typic varieties and forms. This classification was implicitly accepted by Murata (1995) 
when he remodeled the section, and was also embodied in the most recent catalogue 
of the Chilean flora (Marticorena & Quezada 1985). However, when | began to prepare a 
treatment of the Campanulaceae for Flora de Chile (cf. Marticorena & Rodriguez 1995), it 
became apparent that this classification of the section was less than optimal, and that a 
thorough revision was required. 


ETHNOBOTANY 


Though tupa (sometimes rendered as trupa) is the Araucanian name for L. tupa, the plant 
is also widely known in Spanish as tabaco del diablo (Santa Cruz 1932;IbaAez 1955; Mariani 
1965; Hoffmann 1997). This common name (‘devil's tobacco") relates to early reports 
that the dried leaves were smoked by the Mapuche for “unholy” purposes, i.e., as a nar- 
cotic and alleged hallucinogen. Although its hallucinogenic status has not been dem- 


90 SIDA 19(1) 


onstrated clinically, the plant was considered by Schultes (1981, 1990) to be “definitely 
psychoactive.” The actual compounds responsible for this activity are not known. The 
latex does contain pyridine alkaloids, including lobeline and its diketo- and dihydroxy- 
derivatives, lobelanidine and nor-lobelanidine (Santa Cruz 1932; Hill 1970; Raffauf 1970; 
Gibbs 1974); however, these substances are not pyschoactive (Schultes & Hoffmann 1980). 
Lobeline has been used pharmaceutically as a respiratory stimulant, in the treatment of 
bronchial asthma symptoms, and in overcoming nicotine addiction (Blacow 1972;Lewis 
& Elvin-Lewis 1977).In Chile, the expressed latex of L. tupa has been used by rural folk to 
relieve the pain of dental caries, and in compresses to treat joint and hoof pain in horses 
(Murillo 1889;Santa Cruz 1932). The pharmacological bases for such uses are not known. 

Hallucinogenic or not, L. tupa is definitely toxic. Feuillée (1714) reported that the 
odor of the flowers alone was enough to induce vomiting. Despite several trials, | was 
not able to confirm that observation, though the scent is by no means pleasant. In any 
event, beekeepers consider the plant a nuisance, as its nectar gives honey an acrid, un- 
pleasant flavor (Murillo 1889). Though ostensibly a hummingbird-pollinated plant (see 
below),| have observed large orange bees stealing nectar from the flowers via the dor- 
sal slit of the corolla, without contacting the anthers or stigma. 

Feuillée also reported that small quantities of latex rubbed in the eyes would cause 
blindness, a statement | can partly confirm. On more than one occasion during field 
work, | absent-mindedly rubbed my eye after handling material of L. tupa. Though latex 
was no longer perceptible on my hands, in a short time my eye began to sting and water 
profusely and my vision blurred, while the surrounding skin of the face became swollen 
and numb. The effects persisted for nearly an hour. Murillo (1889) reported that inges- 
tion of the plant or its latex could result in intestinal distress and bloody diarrhea. Most 
recently, Matthews (1988) documented the case of a London gardener who was immo- 
bilized for 10-15 minutes after accidentally inhaling airborne dried latex of the plant. 

On a more pleasant note, the species of Lobelia sect. Tupa have considerable horti- 
cultural potential. When seed was first imported to Europe early in the nineteenth cen- 
tury, the resulting plants generated a great deal of interest among gardeners and plant 
aficionados (Sims 1825; Lindley 1826, 1830, 1833;G.Don 1834;D.Don 1834, 1835;Lemaire 
1843; Loudon 1844). Though most were soon lost from cultivation, L. tupa apparently 
remained in the trade (Voss 1894;Finnis 1966; Matthews 1988;Thomas 1990; Huxley 1992) 
and may have been the object of some selective breeding; Chittenden (1923) mentions 
a cultivar from Stokes’ nursery named ‘Brilliant.’ However, seed of the three other species 
recognized here has been reintroduced to botanic gardens (Lammers 1993b), which 


may lead to a renewal of interest in these plants as horticultural subjects. 


SYSTEMATICS 


Much of the data upon which this revision is based was gathered from over 2500 speci- 
mens deposited in 43 herbaria (see Acknowledgments for a complete list of institu- 
tions). This was supplemented by 10 weeks of intensive field work in Chile during January 
1989 and October-November 1990. In company with one to three Chilean colleagues, | 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 91 


travelled over 6500 miles by jeep, covering the entire geographic, elevational, and eco- 
logical range of sect. Tupa, from Copiapé (27°20'S) in the north to Puerto Montt (41°30'S) 
in the south. Fifty-six natural populations, representing all species recognized here, were 
visited and adigisg: - was fale eet to examine several naturalized populations of L. tupa in 
the Juan Ferna dition in January-February 1986 (cf.Lammers 1997). 

During these studies, saciculey arenes was paid to patterns of morphological 
variation within populations. Though some of the infraspecific taxa recognized by 
Wimmer (1953, 1968) are quite striking, if they do not at a minimum form discrete popu- 
lations, they do not merit taxonomic recognition. Emphasis was also placed on discern- 
ing possible geographic, elevational, or ecological correlates of morphological variation. 
In addition, material was also gathered for cytological investigations. The results of that 
work were reported separately (Lammers & Hensold 1992). 

In the end, it was concluded that only four species and no infraspecific taxa could 
be distinguished in sect. Tupa: L. bridgesii Hook.& Arn. [including L. blanda (D.Don) Endl], 
L. excelsa, L. polyphylla [including L. ovata Reiche],and L. tupa [including L. mucronata Cav]. 
The rationale for each of these synonymizations is discussed under the respective species. 

These four species can be divided readily into two pairs on the basis of habit, seed 
morphology,and habitat:(1) a pair of branched solid-stemmed shrubs (rarely small trees) 
with faintly striate oblong or ellipsoid seeds, in the dwarf- and xerophytic-shrub and 
sclerophyllous vegetation zones of the xeric northern portion of the range (L. excelsa 
and L. polyphylia); and (2) a pair of robust herbaceous or suffruticose unbranched hol- 
low-stemmed perennials with minutely foveate-reticulate broadly ellipsoid seeds, in the 
deciduous forest and evergreen temperate rain forest regions of the mesic southern 
portion (L. bridgesii and L. tupa). 

The two pairs are largely allopatric, though populations of L. excelsa and L. tupa do 
approach within a few miles of each other in Prov. Colchagua, Curicd, and Talca. The 
members of each pair are definitely sympatric. In the xeric north, the geographic ranges 
of L. excelsa and L. polyphylla overlap by roughly 90%; the former extends a little farther 
south and not so far north, while the latter generally grows nearer the sea and not so 
high into the Andean foothills. In the mesic south, L. bridgesii has a very restricted distri- 
bution that lies entirely within the broader range of L. tupa. 

Not only are the members of each pair sympatric, they often form mixed popula- 
tions.Of the 17 populations studied in the mesic south, 76% comprised L. tupa only, 18% 
L. bridgesii only, and 6% were mixed. Of the 39 populations studied in the xeric north, 
54% comprised L. excelsa only, 26% L. polyphylla only,and 21% were mixed. 

Despite this apparent opportunity for miscegenation, | could not locate a single 
morphologically intermediate individual during field work, despite intensive searches. 
Furthermore,| have examined only one herbarium specimen that clearly is intermediate 
between two accepted species. The holotype of Tupa kingii Phil., collected at Valparaiso 
in 1868, appears to represent L. excelsa x L. polyphylla (see below). 

From these facts, it is apparent that some very efficient isolating mechanism is at 
work among these species. Differences in chromosome number may be ruled out; all 


92 SIDA 19(1) 


have n = 21, interpreted as hexaploid (Lammers & Hensold 1992;Lammers 1993a;but cf. 
Stace & James 1996 for an alternative interpretation). Phenological differences likewise 
do not appear to be operative. Though L. polyphylla begins to bloom a few weeks earlier 
than L. excelsa, there is still very considerable overlap in their respective phenologies; L. 
bridgesii and L. tupa bloom concurrently. 

The best hypothesis is that isolation within each pair is a product of pollinator dif- 
ferences, a situation similar to that hypothesized for L. cardinalis L.and L. siphilitica L. 
(Thompson & Lammers 1997). Judging from floral morphology, it appears that one mem- 
ber of each pair is adapted to entomophily, the other to ornithophily. The flowers of L. 
bridgesii and L. polyphylla appear suitable for effective visits by bees and other insects. | 
have not observed pollinator visits to L. polyphylla, but large orange bees were observed 
visiting flowers of L. bridgesii, contacting the anthers and stigmas with their backs. When 
the same bees visited L. tupa, it was as a nectar thief, removing nectar via the dorsal slit 
without touching anthers or stigma. With their larger red flowers, L. excelsa and L. tupa 
appear to be adapted to pollination by hummingbirds, though | have not observed this. 
Further biosystematic research clearly is needed to test these hypotheses. 

While the four species fall into two pairs on the basis of habit, seed morphology, 
and habitat, this does not appear to be an accurate reflection of the underlying phylogeny, 
as inferred from cpDNA restriction-site mutations (Knox et al. 1993). Though the south- 
ern pair (L. bridgesii and L. tupa) is supported as a clade by three shared mutations (boot- 
strap value 95%, decay index = 3), the northern pair is not. Instead, L. polyphylla forms the 
basal branch of the tree, sister to the remaining three species; L. excelsa is then the next 
branch, sister to the southern pair. The sister-relationship of L. exce/sa to the southern 
pair is likewise supported by three mutations (bootstrap value 93%, decay index = 3). 

Though no detailed non-molecular analyses have been performed, it does appear 
that morphological data are concordant with the coDNA phylogeny. Lobelia polyphylla, 
the basal branch, is the most discordant member of the section, with its much smaller 
wine-purple flowers. The members of the (L. exce/sa + southern pair) clade all share larger 
red (or pink) flowers, while the southern pair is characterized by its unbranched hollow 
herbaceous or suffruticose stems, usually decurrent leaf margins, and minutely foveate- 
reticulate broadly ellipsoid seeds. Unique traits within the (L. excelsa + southern pair) 
clade are the pink corolla of L. bridgesii; and the tan or pale yellow latex, bibracteolate 
pedicels, and floral color change (Weiss 1995) of L. excelsa. Statements regarding the 
apomorphy or plesiomorphy of any of these traits are best deferred pending better knowl- 
edge of the outgroup. 


= 


Lobelia sect. Tupa (G. Don) Benth.in Benth.& Hook.f.,Gen.PI.2:552. 1876. Tupa G. Don, Gen. 
Hist. 3:700. 1834. Lobelia [unranked] Tupa (G.Don) Heynh., Nom. Bot. Hort. 1:473. 1840. Lobelia 
sect. Futupa E.Wimm., Ann. Naturhist. Mus. Wien 56:365. 1948, nom. illeg. Type (under Art.22.5, 
first confirmed by Pfeiffer 1874):L. tupa L. 


iteroparous shrubs, 0.6-4 m tall (rarely small trees to 6.8 m tall) or robust perennials, 0.5— 
3 m tall; stems woody, solid, and repeatedly branched, or herbaceous or suffruticose, 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 93 


hollow, and unbranched above the base, pubescent or glabrous; latex white, less often 
pale yellow or tan. Leaves simple, alternate, exstipulate, pinnately veined, sessile, pubes- 


cent or glabrous; margin entire or finely callose-toothed, sometimes forming a sagittate 
base decurrent on the stem below the point where the midrib meets the node. Flowers 
perfect, protandrous, resupinate, epigynous, zygomorphic, pedicellate, solitary in the ax- 
ils of the upper leaves or these reduced in size, creating a terminal 10—65-flowered 
bracteate anauxotelic (rarely auxotelic) racemose inflorescence; pedicel equalling or 
shorter than its flower, ebracteolate or with a pair of linear bracteoles at or below the 
middle. Hypanthium obconic, hemispheric, campanulate, or obovoid, pubescent or gla- 
brous, adnate to the ovary; calyx lobes 5, valvate, triangular or narrowly triangular, pubes- 
cent; margin entire or very rarely with a few teeth. Corolla sympetalous, unilabiate, 15-65 
mm long, red (sometimes yellow and orange in bud, or very rarely all yellow throughout 
anthesis), pink, or wine-purple, lacking a nectar spur, glabrous or minutely pubescent; 
tube suberect, curved, or arcuate, slit dorsally to base; lobes 5, valvate, monomorphic, 
shorter than the tube, deflexed, coherent at apex. Stamens 5, alternating with the corolla 
lobes, connate, forming a staminal column shorter than the corolla and free from it; fila- 
ment tube exserted through dorsal slit in corolla, bearing small flattened triangular tri- 
chomes ventrally at base, otherwise glabrous; anthers dehiscing introrsely and longitu- 
dinally, the dorsal three longer than the ventral two, occluding the orifice of the tube, the 
surface of the tube glabrous or the dorsal three with scattered long hairs, the ventral 
two with tufts of white bristles at apex. Ovary inferior, bilocular; placentae axile, large; 
ovules numerous, small, anatropous. Fruit a half-inferior capsule, broadly ellipsoid, ovoid, 


obovoid, oblate, or subspherical, dehiscent by two valves at the rounded or truncate and 
apiculate apex. Seeds small, light brown or golden brown, oblong or ellipsoid and faintly 
striate, or broadly ellipsoid and minutely foveolate-reticulate; testa comprising a single 
layer of cells with long lumina (Type D of Murata 1992, 1995). Chromosome number n = 
21.Four species, endemic to central Chile. 


KEY TO THE SPECIES 


. Shrubs (rarely small trees); stems woody, hee pianeeG. solid; leaf base not 
lon 


lobed, = stem;seeds oblong 1, the surface faintly 

striat 

2. latex wale peleels ciactCe gi or vel rarely with a pair of bracteoles 2-5 
lla 15-25 mm long, wine-purple, 


the tube (7-)10-1 7mm ‘loneeane lobes 4- 12 mm long; filament tube 9-15 mm 

long; anther tube 4-7 mm long, glabrous or very rarely sparsely pubescent; 

capsule 6-9 mm in diameter 1.L. polyphylla 
. Latex tan or pale yellow; pedicels with a pair of bracteoles 2.5-10 mm long 

hypanthium 8-15 mm in diameter; corolla (38-)45-65 mm long, yellow and 

orange in bud, becoming red at anthesis (very rarely all yellow throughout 

anthesis), the tube 25-42 mm long, the lobes 12-33 mm long; filament tube 

29-47 mm long; anther tube 8-11 mm long, pubescent; capsule 11-18 mm in 

diameter 2.L.excelsa 


N 


94 SIDA 19(1) 


1. Robust perennial herbs; stems herbaceous or suffruticose, unbranched above the 
base, hollow; leaf base sagittate, the pair of triangular basal lobes decurrent on the 
stem for 2-30 mm (rarely cuneate or obtuse and non-decurrent); seeds broadly 
ellipsoid, the surface minutely foveolate-reticulat 
3. Leaf apex acute or acuminate, ee ne or aii lamina, 
hypanthium, and calyx lobes oye ubes m long, red 
(very rarely yellow), densely pubescent, the cube 19-35 m mm een tube 
22-33 mm long 3.L.tupa 
3. Leaf apex long acuminate, aristate or cirrhose; lamina, hypanthium, and calyx 
lobes glabrous; corolla 25-36 mm long, pink, glabrous, the tube 13-19 mm long; 
filament tube 14-17 mm long 4.L. bridgesii 


oo 


1. Lobelia polyphylla Hook. & Arn., Bot. Beechey Voy. 33. Dec 1830. Tupa polyphylla (Hook. 
& Arn) G.Don, Gen. Hist. 3:700. 1834.Rapuntium polyphyllum (Hook. & Arn.) C.Presl,Prodr. Monogr. 
Lobel. 29.1836.Dortmanna polyphylla (Hook.& Arn.) Kuntze, Revis.Gen.PI.972.1891. Tyee: CHILE. 
Valparaiso, (26 Oct-2 Nov 1825 or May 1828, leg. Lay & Collie,] Beechey s.n. (HoLotyPe: K!). Infor- 
mation in brackets is taken from the official itinerary (Hooker & Amott 1830). 


Lobelia purpurea Lindl., Edwards’ Bot. Reg. 16:pl. 1325. Jun 1830; non Breiter, Hort. Breiter. 249. 
1817. Tupa purpurea G. Don, Gen. Hist. 3:700. 1834. Rapuntium purpurea (G.Don) C. Presl, Prodr. 
Monogr. Lobel. 28. 1836. Dortmanna purpurea (G. Don) Kuntze, Revis. Gen. Pl. 972. 1891. Tyee: 
CHILE. Valparaiso, Feb 1825, MacRae s.n. (iectotyee [designated by Wimmer 1953: 616]: CGE; 
ISOLECTOTYPES: E! K!). 

Rapuntium besserianum C.Presl, Prodr. Monogr. Lobel. 28. 1836. Tupa besseriana (C.Presl) A.DC. in 
DC., Prodr. 7:393. 1839. Tupa polyphylla var. besseriana (C. Presl) Vatke, Linnaea 38:/27. 1874. 
Dortmanna besseriana (C.Presl) Kuntze, Revis.Gen. PI.972. 1891. Lobelia polyphylla var. besseriana 
(C. Presl) Reiche, Anales Univ. Chile 117:459. 1905. Type: CHILE. Vatparaiso, 2.1 km S of Punta de 
Angeles, on the coastal road from Valparaiso to Laguna Verde, steep rocky slopes of quebrada 
above the road, elev. 120 m, 2 Nov 1990, Lammers, Baeza & Penailillo 7741 (weotyee here desig- 
nated: Fl; isoneorype: CONC!). As no original material could be located, a recent collection that 
agrees with the protologue is here selected to serve as the ne 

ie subdentatum C. Presl, Prodr. Monogr. Lobel. 28. 1836. si Dpeiee (C. ie A.DC. 

n DC.,Prodr. 7:393. 1839.Dortmanna subdentata (C. Presl) Kuntze, Revis.Gen. PI.972. 1891.Lobe- 
fa polyphylla f.subdentata (C.Presl) E.Wimm., Pflanzenr.|V.276b:615.1953. Tyee:CHILE. In ee 
anonymous s.n. (HOLOTYPE: PR! 

ie um bracteosum C. Pres, Prodr. Monogr. Lobel. 29. 1836. Tupa bracteosa (C. Presl) A. ae in 
rodr. 7:393. 1839. Tupa polyphylla var. bracteosa (C. Presl) Vatke, Linnaea 38:727. 
ortmanna bracteosa (C.Presl) Kuntze, Revis.Gen.PI.972.1891.Lobelia polyphylla var. eos 
‘ Presl) Reiche, Anales Univ. Chile 117:459. 1905. Lobelia polyphylla f. bracteosa (C. Presl 
_, Ann. Naturhist. Mus.Wien 56:365. 1 an Type: CHILE. Quillota, fruticetis petrosis collium, 
on [1829], Bertero 1103 (HoLoryee: PRI; Isory V2]! aes M! NY! P! TUB! W!) 

lag hyssopifolium C. Pres|, Prodr. Monogr. eealo 1836. Tupa be Coes (C. Presl) A.DC. 

DC., Prodr. 7:393. 1839. Lobelia hyssopifolia (C. see C. Gay, Fl. Chile 326. 1849. Dortmanna 
ea ifolia (C. Presl) Kuntze, Revis. Gen. Pl.972. 1891.Lobelia polyphylila f. ee (C.Presl) E. 
Wimm., Ann. Naturhist. Mus. Wien 56:365. 1948. Tyee:Peruvia? Chile?’ anonymous 5.n. (HOLOTYPE: 
PRI). 

Tupa polyphylla var. angustifolia Hook. & Arn. ex A.DC.in DC., Prodr. eee 839. Lobelia polyphylla 
var. angustifolia a (Hook. & Arn.ex A. DC.) Heynh., Nom. Bot. Hort. 1:473. 1840. Tyre: CHILE. Prope 
Coquimbo, 1831, Cuming 888 (\ectoryre here designated: K!; sovectoryee: K!). The other syntype 
was Bertero 110 (G-DC [microfiche] 


Te. 


amr 


a 


a 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 95 


Tupa polyphylla var. eine A.DC.in DC,, Prodr. 7:393. 1839. Lobelia polyphylla var. latifolia (A.DC,) 
Heynh., N B 1:473. 1840. Type: CHILE. Prope Valparafso, 1831, Cuming 599 (Lectotype 
is desig Dee oe BM! CONC! E! GH! RSA! W!). The ere syntype was Poeppig 290 
(G-DC [microfiche!]; isosyntypes: BM! W[3]}). 

Tupa atropurpurea Vis., \II. Piant. Nuov. 2:23. 1844. Type: CHILE. Prov. Vavparalso: 3 km N of se ae 
Verde, on the coastal oe from Valparaiso, steep rocky slopes overlooking Bahia Laguna Verde, 
below the road, elev. 150-220 m, 2 Nov , Lammers, Baeza & Penailillo 7760 (Neotype here 
designated: F! isoneotypes: CONC! MO!). Described from living plants cultivated at the botanic 
garden in Padua, Italy.As no original material could be located at PAD (R.Marcucci,in litt. 3 Dec 
1997) or elsewhere, a recent collection that agrees with the protologue is designated as the 
neotype. 

Tupa ovata Phil. Anales Univ. Santiago 43:506. 1873; non G. Don, Gen. Hist. 3:700. 1834. Lobelia 
ovata Reiche, Anales Univ. Chile 117:460. 1905. Type: CHILE. Carrizal bajo, Dec 1871, King s.n. (Ho- 
Lorype: SGO-043561 [three small branches on right side of sheet]!; isotypes: BM! K!). The one large 
branch on the left side of the sheet is A specimen [Bandurrias, Geisse s.n.] which bears 
the manuscript name “Tupa glabrat 

Tupa polyphylla var. coquimbana ae ae 38:727. 1874. Lobelia polyphylla var.coquimbana 
(Vatke) Reiche, Anales Univ. Chile 117:459. 1905. Tyre: CHILE. Prov. Coquimeo [Elqui]: entrada al 
camino del Mineral La Hiquera, al norte de portezuelo de la Cuesta de Buenos Aires, 13 Oct 
1963, Marticorena & Matthei 172 (Neoryee here designated: CONC!: isoneorype: OS!). Because the 
original material presumably was destroyed during World War II (cf. Lammers 1994) and no 
duplicates could be found, a more recent gathering that conforms to the protologue is here 
designated as the neotype. 

Tupa poeppigiana Phil., Anales Univ. Santiago 90:188. 1895. Tyee: CHILE. Quillota, Germain s.n. (Lec- 
torype here designated: SGO-057199!). No specimen was cited in the protologue, merely the 
locality “Quillota.” The sheet designated as lectotype was the only one in Philippi’s herbarium 
that agreed with the protologue and that was annotated with this name in his handwriting. 

Tupa axilliflora Phil., Anales Univ. Santiago 90:188. 1895. Lobelia axilliflora (Phil.) Reiche, Anales 
Univ. Chile 117:460. 1895. Type: CHILE. Coquimbo, Soza s.n. (Lectotyee here designated: SGO- 
057182!).The other syntype cited in the protologue [Catemu, Sep 1860, Philippi s.n. (SGO-057183! 
[photograph: ie isosynTyPe: W!)] was never annotated by Philippi with this name. Although 
Munoz (1960) cited both localities, only the accession number of the latter was listed. 

Tupa eee Phil. oe Univ. mantiage 90:189. 1895. Type: CHILE. Prov.Coquimeo: ad rivulos Serena, 
Oct 


_ Gay 1466 (H GO! [photograph: GH!]). 
Ce eae Phil. sendies ee Santiago 90:189. 1895. Lobelia dei a f. linearifolia (Phil.) E 
turhist. Mus.Wien 56:365. 1948. Type: CHILE. Coquimbo, 1889/90, Geisse s.n. ae 


Totype here aod: 057195!). No specimen was cited in the Se merely the 
locality “Coquimbo."Philippi’s herbarium contains three sheets that matched the protologue 
reasonably well (Munoz 1960). The specimen here designated as the lectotype was the only 
one annotated with this name in Philippi’s handwriting. 
Tupa serrata Phil., Anales Univ. Santiago 90:189. 1895. Type: CHILE. Valparaiso, King s.n. (HOLOTYPE: 
SGO-057185![photograph: GH!)). 
Shrubs, 0.6-2 m tall; stems woody, repeatedly branched, solid, pubescent or glabrous; 
latex white. Lamina 1.8-16 cm long, 0.4-4.8 cm wide, ovate, widely ovate, narrowly ob- 
long, widely elliptic, elliptic, narrowly elliptic, lanceolate, or linear, minutely pubescent or 
glabrous; margin entire, denticulate, serrulate, or serrate; apex rounded, obtuse, acute, 
acuminate, or long acuminate, sometimes mucronate, cuspidate, or aristate; base trun- 


96 SIDA 19(1) 


cate, rounded, obtuse, cuneate, or attenuate. Flowers solitary in the axils of the upper 
leaves, or aggregated into a 10-30(-45)-flowered raceme; bracts 7-50(-75) mm long, 
2.5-28 mm wide, widely elliptic, elliptic, narrowly elliptic, lanceolate, or ovate, glabrous or 
minutely pubescent; pedicels 7-17 mm long, ebracteolate (very rarely some with a pair 
of linear bracteoles 2-5 mm long at or below the middle), minutely pubescent. Hy- 
panthium 4-7 mm long, 4-8 mm in diameter, obconic, campanulate, or rarely obovoid, 
pubescent or subglabrous. Calyx lobes 3-8 mm long, 1-3 mm wide, triangular or nar- 
rowly triangular, pubescent; margin entire or very rarely with a few teeth; apex acumi- 
nate or long acuminate. Corolla 15-25 mm long, wine-purple, minutely pubescent; tube 
(7-)10-17 mm long, 1.5-3 mm in diameter at middle, suberect or curved; lobes 4-12 
mm long, 0.8-2 mm wide. Filament tube 9-15 mm long, purple; anther tube 4-7 mm 
long, 1.2-2.5 mm in diameter, grey with pale stripes, glabrous or the dorsal three rarely 
with scattered long pubescence. Capsule 9-12 mm long, 6-9 mm in diameter, broadly 
ellipsoid or ovoid. Seeds 0.7—-1 mm long, 0.3-0.4 mm in diameter, oblong or ellipsoid, 
light brown or golden brown, faintly striate (Murata 1995, figs.52-53).Chromosome num 
bern = 21 (Lammers & Hensold 1992;Lammers 1993a). 

lcones.—Lindley (1830) [as L. purpurea];D.Don (1834); Hooker (1837);Loudon (1844), 
pl. 66, fig. 3; Navas (1979), pl.42D-G. 

Distribution, Habitat,and Phenology.—Endemic to xeric north-central Chile between 
latitude 27°S and 34°S, in the regions of dwarf- and xerophytic-shrub and sclerophyllous 
vegetation (cf.Walter 1973). Frequent on dry rocky slopes and cliffs, sometimes in sandy 
soil, from near sea level (often in sight of the ocean) up to 900 m, rarely as high as 1200 m 
above sea level, often in company with L. excelsa. Flowering August through February. 

Discussion.—This species is extremely variable in foliar features, especially size and 
shape of the lamina and the degree of reduction in leaves subtending flowers (ie., 
whether the plant forms a discrete bracteate inflorescence or bears solitary flowers in 
axils of unreduced leaves). This inordinate variability is the source of the species’ exten- 
sive synonymy. Though the extremes [e.g., plants distinguished by Wimmer (1953) as L. 
polyphylla f.linearifolia and L. ovata] appear most distinct, it was noted during field study 
that many populations were quite heterogeneous in foliar morphology. Individuals that 
differed considerably in leaf length, width, and outline often were found growing side- 
by-side. Furthermore, leaves that emerged and expanded early in the season often dif- 
fered significantly from those appearing later. In some cases, two or three of Wimmer's 
“taxa” could be found on a single individual.For example, in Marticorena et al. 1367 (CONC), 
the lower leaves key to var. besseriana while the upper leaves key to var. /atifolia. This 
suggests that at least some of the observed variation is environmentally induced (re- 
lated to rainfall or day-length perhaps) rather than under genetic control, or perhaps 
related to heteroblasty. 

In detailed multivariate analyses of this species (Lammers & Glass 1998), no pat- 
terns were discernible in the data, which were gathered from 64 herbarium sheets, in- 
cluding nearly all type specimens. Variation in foliar features was absolutely continuous, 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 97 


with no gaps by which meaningful taxa (including L. ovata) could be distinguished. No 
correlations were detected between foliar morphology and any geographic, ecologic, 
or elevational parameters. Furthermore, the plants examined cytologically (Lammers & 
Hensold 1992) well represented the diversity of foliar morphology; all showed n = 21. 

In contrast to foliar structures, the flowers of L. polyphylla are remarkably homoge- 
neous. They are distinctive within the section because of their small size and the dark 
pigmentation. The hue of the corolla is a very intense red-purple which compares favor- 
ably to the color of a good Chilean red wine (‘vino tinto’). The only variation in floral 
features which appeared to be geographically correlated was anther pubescence, which 
showed a clinal pattern of variation: pubescent anthers were commonest towards the 
north, glabrous anthers towards the south. 

This species is unusual among Lobelicideae in showing some variability in the pres- 
ence or absence of bracteoles. However, the single specimen found that had bibracteolate 
pedicels (Marticorena et al. 1379, CONC) also showed typical ebracteolate pedicels in the 
same inflorescence. Similarly, inflorescences in sect. Tupa ordinarily are anauxotelic (ie., 
incapable of renewing vegetative growth; cf. Stein 1987), with growth resuming via 
branching from axillary buds below the spent inflorescence. However, one collection 
(Schlegel 2752, CONC, SGO) was seen that was clearly auxotelic, with an inflorescence in 
full anthesis surmounted by ca. 15 cm of new vegetative growth. 


— 


eee specimens. CHILE. Prov. Copiapo: 20-25 km W of Totoral, Jaylor et al. 10807 (ASU, 

MO). Prov. Huasco: Las Lozas, Bocher et al. 545 (C); Huasco, Kubitzki 290 (VALD); Mina Los Cristales, 
eae et aa 1700 (CONC); Huasco, Montero 7611 (CONC, OS); Quebrada del Pretil, Ricardi & 
Marticorena 3966 (CONC); 3-4 km SW of Huasco, Worth & Morrison 16245 (GH, UC). Prov. Elqui: Punta 
Arrayan, Dillon et al. 5437 (F); La Higuera, Jan 1886, Fonck s.n. (SGO);La Serena, Frodin 153 (UPS); entre 
La Serena y Vallenar, Garaventa 4241 (CONC); entre Tongoy y Guanaqueros, Gleisner 14 (CONC); El 
Tofo, Kubitzki 279 (CONC, VALD); Punta Herradura, Lammers 7650 (ASC, CONC, F, MU, OSH, UC, US); 
Punta Teatinos, Landrum & Landrum 5634 (ASU, GH, NY, RSA, UC); Coquimbo, Oct 1878, Philippi s.n. 
(SGO); La Serena, Sep 1898, Reiche s.n. (SGO); Cuesta de las Cordas, Ricardi & Marticorena 4564/949 
(CONC, OS); 20 km S of Incahuasi, Ricardi & Marticorena 4896/1281 (CONC, OS, Universidad de Talca); 
La Serena, Sparre 2793 (S, SGO); N of La Serena, Taylor et al. 10654 (ASU, F, MO). Prov. Limari: Talinay, 
Jiles 439 (CONC); Fray ae Kubitzki 100 (VALD); 1 km N of Mantos de Hornillo, Lammers et al. 6372 
(CONC, F); 11 km N of Mantos de Hornillo, Lammers et al.6386 (CONC, F), 6389 (B, CONC, F, MU), 7665 
(CONC, F), 7666 (CONC, F); ie Munoz B-152 (SGO); Fray Jorge, Skottsberg & Skottsberg 887 (GB, 
NY, S); Fray Jorge, Werdermann 901 (A,B, BM, E,F, GH, M, MO, NY, UC, US). Prov. Choapa: Pichidangui, 
Correa 67 (SGO); Puente Negro, Jiles 4354 (CONC); 0.2 km N of Los Vilos, Lammers et al. 6331 (B,C, 
CONC, GB, F, MEXU, MU); 7 km N of Los Vilos, Lammers et al.6341 (CONC, F, MU, NY); 8.9 km N of Los 
Vilos, Lammers et al. 7536 (CONC, F, TEX); 8.3 km N of Puerto Oscuro, Lammers et al. 7668 (B, CONC, F, 
TEX); 23.6 km S of Los Vilos, Lammers et al. 7677 (CONC, F, MU, OSH, US); Estacion Ingenerio Barriga, 
Marticorena et al. 210 (CONC); Agua Amarilla, Marticorena et al. 332 (B); N of Los Vilos, McGill 1028 
(ASU); IIlapel, Rose & Rose 19247 (NY, US); 33 km SW of Illapel, Worth & Morrison 16646 (BH). Prov. 
Petorca:4 km N of Longotoma,Lammers et al.6397 (CONC, F,MU, OS, UB); 1 km.S of Papudo,Lammers 
et al. 6402 (CONC, F, MU); 1.3 km N of northernmost road to Zapallar, Lammers et al. 7700 (CONC, F). 
Prov. San Felipe de Aconcagua: Quebrada de Las Palmas, Sep 1965, Ramirez s.n. (VALD). Prov. 
Valparaiso: Limache, Garaventa 299 (CONC); Valparaiso, queen 129 (CONC); Concon, Gunckel 
19081 (CONC); Renaca, Jaffuel 1752 (CONC); Quintero, Lammers et al.6412 (CONC, F),6413 (CONC, F), 


98 SIDA 19(1) 


6414 (F),6415 (F), 7723 (ASC, CONC, F), 7724 (CONC, F), 7725 (BM, CONC, F, UC); 2.6 km S of Punta de 
Angeles, Lammers et al. 7753 (CONC, F, NY, OS); Mirasol, Landrum 3821 (MICH, MO, NY, SGO);Valparaiso 
Dec 1851, Philippi s.n. (SGO);Placeres, Stebbins 8542 (UC); Quintero, Werdermann 30 (BM, CAS, E,F, GH, 
M, MO, S, U, UC, US). Prov. San Aaionie: El Tabo, Gunckel 25483 (CONC); Algarrobo, Lammers et al. 
7800 (B, CONC, F); Quebrada de Cordoba, Jan 1980, Navarreto s.n. (VALD).Regién Metropolitana: 
Tejas Verdes, Gunckel 24556 (CONC); Pentaflor, Montero 1748 (CONC); Cartagena, Navas 2298 (CONC); 
Dunas de Las Cruces, 19 Oct 1950, Pfister & Ricardi s.n. (CONC, OS). 

CULTIVATION.U.S.A. California: University of California Botanical len, Berkeley, Jul 1963, 
Hutchinson s.n. (MIC 


2.Lobelia excelsa Bonpl.,Descr.P]. Malmaison 112.1816.Rapuntium excelsum (Bonpl.) C.Presl, 
Prodr. Monogr. Lobel. 29. 1836. Dortmanna excelsa (Bonpl.) Kuntze, Revis. Gen. Pl. 972. 1891 
Type: FRANCE. Hort. Malmaison, Jun 1813 (Lectorype here designated: Bonpland 1816, pl. 46!). 
As no original material was located, the plate published with the protologue is here desig- 
nated as the lectotype. 

Lobelia gigantea Sims, Bot. Mag. 32:pl. 1325. 1810; non Cav., Anales Hist. Nat. 2:104. 1800. Lobelia 
salicifolia Sweet, Hort. Suburb. Lond. 37. 1818. Tupa salicifolia (Sweet) G. Don, Gen. Hist. 3:700. 
1834. Type: CHILE. Valparaiso, 1794, Menzies s.n. (Lectorype here designated: BM! isovectorype: MO!), 
The name was based on this collection and on living plants growing ina ae nursery. 


Lobelia arguta Lindl., Edwards’ Bot. Reg. 12:pl. 973. 1826. Tupa arguta (Lindl.) G. Don, Gen. Hist. 
3:/00. 1834. Dortmanna arguta (Lindl.) Rane dale Gen. PI. 972. ie Type: ne ee 
Chiswick Garden, Sep 1825 (Lecrorype here dley vee pl.973!).A 
rial was located, the plate published with the ean is here designated as ie. be aeaihann 


Lobelia neriifolia Moris, Enum.Sem Ha tite cra ree eG CaRDENAL CARO: 
1.2 km E of Pichilemu on highway to Nancagua, frequent on arid rocky bluffs above the road, 
elev. 45 m, 15 Nov 1990, Lammers, Baeza & Penailillo 7890 (Neorype here designated: OSH!; 
isoneorypes: CONC! F! MU! TEX! US!). Described on the basis of living plants grown in the botanic 
garden at Turin, Italy, from seeds sent by Bertero. As no original material or Bertero specimens 
could be located, a recent collection that conforms to the protologue is here designated as 


the neotype. 
Tupa glaucescens Phil. Anales Univ. Santiago 90:187. 1895. Type: CHILE. San Isidrio, Dec 1882, Philippi 
s.n. (HOLOTYPE: SGO)), 


Shrubs, 2-4 m tall, rarely trees to 6.8 m; stems woody, repeatedly branched, solid, gla- 
brous or minutely pubescent; latex tan or pale yellow. Lamina 2.8-15 cm long, 0.7-3.1 
cm wide, oblong, narrowly oblong, elliptic, or narrowly elliptic, rarely lanceolate, glabrous 
or minutely pubescent; margin serrulate, minutely serrulate, or subentire; apex obtuse, 
acute, or acuminate, sometimes cuspidate, mucronate, or aristate; base rounded, obtuse, 
cuneate, or attenuate. Flowers solitary in the axils of the upper leaves; pedicels 12-45 
mm long, bibracteolate on the lower third or rarely at the middle, minutely pubescent; 
bracteoles 2.5-10 mm long, linear, minutely pubescent. Hypanthium 5-10 mm long, 8- 
15 mm in diameter, hemispheric, broadly hemispheric, broadly campanulate, obconic, or 
broadly obconic, glabrous or minutely pubescent. Calyx lobes 3.5-9(-20) mm long, 1-4 
mm wide, triangular or narrowly triangular, minutely pubescent; apex acuminate or long 
acuminate. Corolla (38-)45-65 mm long, yellow and orange in bud, becoming red at 
anthesis (very rarely all yellow), glabrous or sparsely pubescent with minute hairs; tube 
25-42 mm long, 2.5-6 mm in diameter at middle, suberect or curved; lobes 12-33 mm 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 99 


long, 1-3 mm wide. Filament tube 29-47 mm long, red; anther tube 8-11 mm long, 2-4 
mm in diameter, pale straw-colored, the dorsal three with long white hairs on the sur- 
face (especially toward apex). Capsule 10-15 mm long, 11-18 mm in diameter, ovoid, 
obovoid, oblate, or subspherical.Seeds 1 mm long,0.4 mm wide, oblong, honey-colored, 
faintly striate (Murata 1992, figs. 49-50, 70). Chromosome number n = 21 (Lammers & 
Hensold 1992;Lammers 1993a). 

Icones.—Sims (1810) [as L. gigantea]; Bonpland (1816), pl. 46; Lindley (1826) [as L. 
arguta];Mufioz (1966), pl. 128 [as L. tupa]; Navas (1979), pl.42A-C [as L. salicifolia]; Hoffmann 
(1997), pg.218 no.1 [as L. tupa]. 

Distribution, Habitat,and Phenology.—Endemic to xeric north-central Chile between 
latitude 29°S and 35°S, in the regions of dwarf- and xerophytic-shrub and sclerophyllous 
vegetation (cf.Walter 1973).Frequent on dry rocky slopes and cliffs, sometimes in sandy 
soil, from near sea level (though seldom in sight of the ocean) up to 1200 m above sea 
level, often in company with L. polyphylla. Flowering September through February. 

The report of this species from Bolivia (Wimmer 1953) is apparently due to an 
inadvertant switching of labels. Rusby 623 (US) allegedly represents the only known speci- 
men of L. excelsa from Bolivia, while Rusby 634 (US) is sup} lly the only specimen from 
Chile of another lobelioid, Centropogon cornutus (L.) Druce, which is indigenous to Bo- 
livia (cf. Stein 1987). However, duplicates of Rusby 634 at GH and NY are specimens of L. 
excelsa from Valparafso. Clearly, labels were accidentally switched at some point, result- 
ing in the erroneous reports. 

Discussion.—I|n contrast to L. polyphylla, the other woody member of the section, L. 
excelsa is relatively homogeneous in its vegetative morphology. One feature that does 
vary more in this species than in L. polyphylia is habit. Although L. excelsa is ordinarily a 
shrub, one individual was encountered in Prov. Limari in 1989 (Lammers et al. 6382, B, C, 
CONC, F, MU) that was definitely a tree. The plant was fully 6.8 m tall and did not begin to 
branch until 2.5 m above the ground. Its twin boles were 11.5 cm and 9.5 cm in diameter 
just above the base and were of sufficient strength that the tree could be climbed to 
collect flowering branches without damage. 

Lobelia excelsa is unique within the section in its tan or pale yellow (vs. white) latex, 
regularly bibracteolate pedicels, and for the color change of its flowers, which go from 
yellow and orange in bud to brilliant red at full anthesis. The taxonomic distribution and 
biological significance of such color changes were detailed by Weiss (1995). 

Plants bearing pure sulfur yellow flowers with no trace of orange or red (and thus 
no color change) are known, but only from cultivation. The label of a specimen culti- 
vated in Switzerland [anonymous s.n., 29 Jul 1858 (NY)] states “flor sulphurei.” More re- 
cently, Eric Knox (pers.comm.) reported that plants grown at the University of Michigan's 
Matthei Botanical Garden from seed of Lammers et al.6393 consistently bore yellow flow- 
ers over a period of three years.| know for a fact that the plants from which the seed was 
obtained bore normally pigmented flowers. It would be of interest to learn if plants with 
yellow flowers ever occur in natural populations, and if so, under what conditions. 


100 SIDA 19(1) 


Representative specimens. CHILE. Prov. Elqui: aren 14 Jan 1971, Blaise s.n. (SGO); Cuesta de 
Buenos Aires, Kubitzki 278 (VALD); Cuesta de Buenos Aires, Lammers et al. 7604 (ASC, CONC, F,OS, UC); 
erba Loca, Bien 327 (RSA, UC); La Serena, Sparre 2780 (8, SGO); 42 km N of La Serena, Worth & 
Morrison 16328 (BH). Prov. Limari: Monte Redondo, Jiles 372 (CONC); Quebrada La Higuera, Jiles 
1131 (CONC); Cabreria, Jiles 1150 (CONC); Fray Jorge, Sep 1958, Kummerow s.n. (CONC); 1 km N of 
Mantos de Hornillo, Lammers et al.6375 (CONC, F, MU, NY): 17 km N of Mantos de Hornillo, Lammers 
et al. 7664 (BM, CONC, F). Prov. Choapa: 1 km S of Los Vilos, DeVore 1574 (OSH); 0.2 km N of Los Vilos, 
Lammers et al.6330 (C, CONC, F, MEXU, MU, NY); 10 km N of Los Vilos, Lammers et al.6348 (CONC, F, 
NY); 11 km E of Panamerican Hwy on road to Canela Baja, Lammers et al.6353 (CONC, F,GB, MU, 
NY);5 km S of Canela Baja, Lammers et al. 6357 (B, CONC, F, GB, MU);8 km NE of IIlapel, Lammers et al. 
6362 (CONC, F,MU):7 km SE of Illapel, Lammers et al.6364 (CONC, F, NY); 24 km SW of Illapel, Lammers 
et al.6367 (B, CONC, F, MU);6 km S of Puente Amolanas, Lammers et al.6370 (CONC, F, MU, NY): 14. 
km S of Rio Choapa bridge, Lammers et al. 7676 (CONC, F, NY); entre Illapel & Los Vilos, Marticorena & 
Matthei 427 (CONC); Illapel, Montero 2399 (CONC); Los Vilos, Montero 8772 (CONC), Montero 10777 
(CONC); Illapel to Huentelauquen, West 3940 (GH, MO, UC). Prov. Petorca: 3 km S of Zapallar, Gardner 
& Page 5085 (E); Pichicuy, Lammers et al. 6393 (B, CONC, F, MU); 2 km E of Papudo, Lammers et al. 6399 
(B, CONC, F,MU);1 km S of Papudo, Lammers et al.6401 (B, CONC, F, MU);21 km N of Nogales, Lammers 
et al.6408 (CONC, F, MU, NY); 14 km N of Nogales, Lammers et al.6411 (CONC, F MU, NY); 1.3 km N of 
northernmost road to Zapallar, Lammers et al. 7701 (ASC, CONC, F); Pichicuy, Marticorena et al. 177 
(CONC); Papudo, Montero 8023 (CONC); Cerro Iman, Villagran & Meza 750 (SGO), 932 (SGO).Prov. San 
Felipe de Aconcagua: Quebrada de Cabildo, 29 Jan 1950, Pfister s.n. (CONC, OS); Quebrada de Las 
Palmas, Sep 1965, Ramirez s.n. (VALD). Prov. Quillota: 12 km from Tiltil towards Olmue, Gardner & 
Knees 5446 (E); Olmue, Bocher et al. 587 (CQ); 2 km S of San Pedro, Lammers et al.6418 (CONC, F, MU,NY, 
UB); Parque Nacional La Campana, Lammers et al. 6440 (B, CONC, F, MU), 6448 (CONC, F, MU), 6450 
(CONC, F, MU),6452 (CONC, F). Prov. Valparaiso: Valparaiso, Gaudichaud 129 (P);Las Zonas, Harshberger 
1074 (NY, PENN); Quebrada de la Tortuga, 24 Feb 1941, Junge s.n. (CONC, OS); 9 km N of Concon, 
Lammers et al.6417 (CONC); 2.6 km S of Punta de Angeles, Lammers et al. 7754 (CONC, F, UC);3 km N 
of Laguna Verde, es si al. ba (CONC, F); Algarrobo, Landrum 3341 (ASU); Mirasol, Landrum 
3822 (MICH, NY, SGO):Q Vina, Lourteig 2510 (K,S);El Salto, Meyer 9336 (MO, UC);Valparaiso, 
Dec 1862, Philippi s.n. S60 del Mar, Ricardi 5449 (CONC); Valparaiso, Rusby 634 (K, MICH, NY, PH); 
Valaparaiso, Rusby 634 (GH, NY); Agua Potable, Schlegel 387 (CONC); Cerro Valparaiso, Schlegel 967 
(CONC); Quebrada Verde, Schlegel 89 (CONC); El Salto, Skottsberg & Skottsberg 947 (GB, S); El Salto, 
Solbrig et al. 3600 (GH, NY); 3.9 km N of Concon, Spooner & Contreras 4304 (CONC, F, MO, WIS). Prov. 
San Antonio: Quebrada Cordoba, Gentry 68043 (NY); 3 km E of Algarrobo, Lammers et al. 7795 (B, 
BM,CONG,F);N of San Sebastian, 2 Feb 1971,Oehrens s.n. (VALD);El Tabo, Feb 1968, Santos s.n. (VALD). 
Region Metropolitana: Quebrada Dormida, Garaventa 1323 (CONC); Cerro La Cruz, 30 Sep 1932, 
Olathe s.n. (CONC); Dunas de las Cruces, 19 Oct 1950, Pfister & Ricardi s.n. (CONC, OS); Quebrada La 
Plata, Schlegel 1680 (CONC). Prov. Cardenal cue: pico Sint Montero 9286 (CONC); 9 km S of 
Pichilemu, Lammers et al. 7903 (CONC, F MO, NY uario Nil Lammers et al. 7917 (CONC, F, NY); 
4.2 km N of Bucalemu, Lammers et al. 7933 (CONC, F) Bieler Sanz de Cortazar 573 (SGO), Prov. 
Colchagua: Ciruelos, Aravena 47 (SGO); Matanzas, Mar 1878, Ferndndez s.n. (SGO); Nancagua, 11 Jan 
1951, Ricardi s.n. (CONC, OS). Prov. Curicé: Lipimavida, Aravena 34L (SGO); Lipimavida, Spooner & 
Contreras 4332 (CONC, F, WIS); Punta Totorilla,6 Feb 1969, Villagran neva 5.n. (SGO); 1 km antes de 
lloca, Weldt 107 (CONC). Prov. Talca: Quivolgo, Matthei & Bustos 34 (B); Constitucién, Feb 1895, Philippi 
5.n. (SGO). 
CULTIVATION. SWITZERLAND: Hort. Bot. Basil., 29 Jul 1858, anonymous s.n. (NY). 


3. Lobelia tupa L., Sp. P/.929.1753,'trapa.’ Tupa feuillei G. Don, Gen. Hist. 3:700. 1834. Rapuntium 
tupa (L.) C.Presl, Prodr. Monogr. Lobel. 28.1836. Dortmanna tupa (L.) Kuntze, Revis. Gen. P1972. 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 101 


1891.Lobelia feuillei (G.Dor 1) Voss in Siebert t & Voss, Vilm Blumengartn.(ed. 3) 1:577.1 1894,nom. 
illeg. Tyee: CHILE.Mountains, 37°S (Lectorype [designated by Matthews 1988:161]:Feuillée 1714, 
pl. 291), 


Lobelia mucronata Cav., con. 6:11.1801. Tupa cavanillesiana G. Don, Gen. Hist. 3:700. 1834, nom. 
illeg. Rapuntium mucronatum (Cav.) C. Presl, Prodr. Monogr. Lobel. 29. 1836. Tupa mucronata 
(Cav) A.DC.in DC., Prodr. 7:392. 1839. Tupa feuillei var. mucronata (Cav.) Vatke, Linnaea 38:727 
1874. Dortmanna mucronata (Cav.) Kuntze, Revis.Gen. Pl.972.1891.Lobelia tupa var.mucronata 
Say Reiche, Anales Univ. Chile 117:458. 1905. Type: CHILE. Fundo Cuchacucha, Nee s.n. (HOLOTYPE: 

A! [photographs: F! GH! MICH! WI: fragments: F! CONC! W)). 
obelo sera a Reise 1:300. 1834. Tyee: CHILE. Prov. Curico: Hacienda Monte Grande, alt.ca. 
924, Werdermann 572 (notype here designated: BM!; isoneotyres: E! F! GH! M! MO! 
7 UC!). ne original material could be located, so a more recent collection from the same 
general area, which agrees with the protologue, is here designated as the neotype. 

Tupa berteroi A.DC.in DC., Prodr. 7:392.1839,‘berterii, Tupa feuillei var.berteroi (A.DC,) Vatke, Linnaea 
38:727.1874,'berterii.’ Dortmanna berteroi (A.DC.) Kuntze, Revis. Gen. PIl.972. 1891. Lobelia tupa 
var. berteroi (A. DC.) Reiche, Anales Univ. Chile 117:458. 1905, ‘berterii.’ Lobelia mucronata var. 
berteroi (A.DC.) E.Wimm.,Pflanzenr.\V.276b:614. 1953,'berterii.’ Type: CHILE. Rancagua, in sylvaticis 
umbrosis, torrentes et rivulos, Apr 1828, Bertero 389 (HoLorype:G-DC [microfichel]; isorypes: P! SGO!). 
Orthography corrected following the recommendations of Stuessy and Marticorena (1990). 

upa mucronata var. hookeri A.DC.in DC., Prodr.7:392. 1839. Lobelia mucronata f.hookeri (A.DC,) E. 
Wimm., Pflanzenr. lV.276b: 614. 1953. Type: GREAT BRITAIN. Scotland, Glasgow Botanic Garden, 
Aug 1832 (Hotoryee: Hooker 1833, pl. 3207!) 

Tupa montana Phil., Anales Univ. Santiago 43:506. 1873;non C.Wright ex Griseb., Cat. Pl. Cub. 159, 
1866. Dortmanna philippiana Kuntze, Revis. Gen. P|.972.1891.Lobelia tupa var.montana Reiche, 
Anales Univ.Chile 117:459. 1905. Type: CHILE. Haclienda].del Principal, 1870, Philippi s.n. (HOLOTYPE: 
SGO-057202!; isotype: SGO-043568! [photograph: GH!]). 

Tupa feuillei var. macrophylla Vatke, Linnaea 38:726. 1874. Type: GREAT BRITAIN. Lowe's Nursery 
(Lecrorype here designated: Lindley 1833, pl. 1612!).Vatke also cited Philippi 340, Ochsenius s.n., 
and a specimen cultivated at Berlin in 1832, all deposited at B.None of these specimens could 
be located and presumably perished during World War Il (cf Lammers 1994).| have seen du- 
plicates of the Philippi specimen at K, P.and W, but they do not conform to the description 
given in the protologue, while the plate doe 

Dortmanna bicalcarata Kuntze, Revis.Gen.PI.3(2 : 186.1898. Lobelia bicalcarata (Kuntze) A.Zahlbr. 
ex K.Schum., Just’s Bot. Jahresber. 26(1):373. 1900. Lobelia tupa ae mee (Kuntze) E.Wimm., 
Ann. Naturhist. Mus.Wien 56: sc aks ie Sila Gnigueyanites 19 Feb 1892, Kuntze $.n. (HOLO- 
type [Mounted on t | J:h F! GH! MICH! i SGO!],W! [photo- 
graphs: A! F! MICHI)). 

Lobelia mucronata f.ovalifolia E.Wimm., Pflanzenr.|V.276b:614. 1953. Type: GERMANY. H[ort.] Bonn, 
1848, anonymous s.n. (HOLOTYPE: B}). 

Lobelia tupa var. pavonii E.Wimm., Pflanzenr. IV.276c: 881.1968. Type: CHILE. Ruiz & Pavon s.n. (HOLO- 

ype: BMI), 


Robust perennials, 0.5—3 m tall; stems several from the base, normally unbranched, hol- 
low, herbaceous or suffruticose, densely short or long pubescent; latex white. Lamina 
4.5-25.5 cm long, 1.2-7.8 cm wide, ovate, oblong, elliptic, narrowly elliptic, or rarely lan- 
ceolate; upper surface densely short pubescent; lower surface very densely short pu- 
bescent; margin serrulate or minutely serrulate; apex acute or acuminate, sometimes 


102 SIDA 19(1) 


mucronate or cuspidate; base sagittate, the pair of basal lobes decurrent on the stem for 
2-30 mm (rarely cuneate or obtuse and non-decurrent). Inflorescence a 10-65-flowered 
raceme, densely short pubescent; bracts 7-45(-85) mm long, 2-15(—20) mm wide, ovate, 
lanceolate, or rarely linear, the apex acuminate, the base sagittate, the pair of basal lobes 
decurrent on the stem for 2-33 mm (or very rarely free from the stem), rarely obtuse and 
non-decurrent; pedicels 8-30 mm long, ebracteolate. Hypanthium 5-10 mm long, 8-14 
mm in diameter, hemispheric, depressed hemispheric, obconic, or broadly campanulate, 
densely short pubescent.Calyx lobes 2-8 mm long, 1-3 mm wide, triangular or narrowly 
triangular, short pubescent; apex acuminate, long acuminate, or rarely acute. Corolla 31- 
49 mm long, red (very rarely yellow), sparsely or densely short pubescent; tube 19-35 
mm long, 2-4 mm in diameter at middle, arcuate;lobes 10-22 mm long, 1-2.5 mm wide. 
Filament tube 22-33 mm long, red; anther tube 6-9 mm long, 2-3 mm in diameter, grey, 
the dorsal three pubescent with long white hairs or rarely glabrous. Capsule 0.9 cm long, 
1.2 cm in diameter, ovoid. Seeds 0.7 mm long, 0.3 mm wide, broadly ellipsoid, honey- 
brown, minutely foveolate-reticulate (Murata 1995, figs.54-55). Chromosome number n 
= 21 (Vilmorin & Simonet 1927; Spooner et al. 1987; Lammers & Hensold 1992;Lammers 
1993a); the voucher (Sanz 573, SGO) for Sanz de Cortazar’s (1948) report of n = 16 in L. 
tupa is actually a specimen of L. excelsa, and the count is considered erroneous in any 
event (Lammers & Hensold 1992; Lammers 1993a), 

Icones.—Feuillée (1714), pl. 29 [as“Rapuntium spicatum ..."];Cavanilles (1801), pl.516 
Las L. mucronata]; Sims (1825); Sweet (1827-29): Hooker (1833) [as L. mucronata]; Lindley 
(1833); Loudon (1844), pl. 66, fig. 1; Santa Cruz (1932), p.99; Wimmer (1953), fig. 96; Finnis 
(1966), fig. 48; Schultes (1976), p. 153, (1981), p. 122; Matthews (1988); Thomas (1990), pl. II, 
no. 5; Hoffmann (1997), pg. 218, no. 2 [as L. bridgesii]. 

Distribution, Habitat, and Phenology.—Endemic to mesic south-central Chile between 
latitude 32°S and 42°S, in the regions of deciduous forest and evergreen temperate rain 
forest (cf.Walter 1973). Also naturalized on Masatierra (Isla Robinson Crusoe) in the Juan 
Fernandez Islands sometime prior to 1824 (Matthei et al. 1993). Common on roadsides, 
fields, streambanks, grassy slopes, and forest margins, at elevations from near sea level 
up to 400 m (rarely up to 940 m). Flowering late October to early April. 

Discussion.—Lobelia tupa varies considerably in the nature of its leaf and bract bases, 
variation which Wimmer (1953) used to recognize several taxa. Typically, the leaf margins 
continue as a pair a broad triangular auricles for some distance below the point of at- 
tachment of the midrib at the node, forming a sagittate base that is adnate to the stem. 
Wimmer (1953, 1968) treated plants with relatively short auricles, particularly among the 
bracts, as var. tupa or (if the leaves were lanceolate) var. pavonii. Those with very long 
auricles were var. montana or (if the auricles of the bracts were free from the inflores- 
cence rachis) var. bicalcarata. Otherwise similar plants with obtuse or cuneate non-de- 
current leaf bases were segregated as L. mucronata. Plants that were intermediate be- 
ta and L. tupa, i.e., those with very shortly decurrent auricles, were treated 
as L. mucronata var. berteroi. The intermediacy of this variety is highlighted by the fact 
that Vatke (1874) and Reiche (1905, 1910) previously had assigned it to L. tupa. 


na 


tween | 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 103 


Careful study of natural populations showed that the length of the basal auricles 
varied considerably within populations and even within individual plants. This was ap- 
parent even in herbarium material.In Gay 1468 (SGO), the auricles were only 3 mm long 
in the basal-most leaves, but 15 mm long within the inflorescence. As such, the varieties 
of L. tupa recognized by Wimmer (1953, 1968) do not merit recognition. 

The case of L. mucronata is more complex. Though not emphasized by Wimmer, it 
differs from L. tupa not only in it non-decurrent leaves, but also in its very sparse inflores- 
cence (cf. Hooker 1833).Here,only 10-15 flowers form before the apical meristem aborts; 
the flowers appear to be largely in bloom all at once and to take on a nodding rather 
than spreading or ascending posture. However, all other features of the plants, including 
the flowers and the seeds, are identical to ‘ tupa. PUnthenniots 2 specimens referable to 
L. mucronata have been collected within the g levational range of L. tupa. 

| was unable to locate plants in nature ie ade this description, and have 
seen only a few specimens besides the types cited above. While it is possible that L. 
mucronata is an extremely rare or possibly extinct species, perhaps adapted to some 
unusual edaphic niche, other hypotheses must be considered. The handful of specimens 
examined may simply represent plants of L. tupa whose apical meristems were dam- 
aged at a critical point of development by some insect or pathogen, resulting in aber- 
rant growth. They may represent an occasional genetic variant, perhaps a simple Men- 
delian recessive, of that species. Alternatively, these plants may be relicts close to the 
evolutionary divergence of L. excelsa and L. tupa, or recent La of these two species. 
Although they are not sympatric today, their ranges do appr ne another (see above) 
and may have overlapped in the past. The most recent gathering of plants referable to L. 
mucronata (Montero 736, CONC) was collected in 1928 in Prov. Colchagua, in the zone of 
closest approach between these two species. 

Until such time as sufficient new data permit support or rejection of some of these 
hypotheses,| am loathe to recognize L. mucronata as a distinct species. Because it occurs 
within the geographic range of L. tupa and because its flowers and seeds are indistin- 
guishable from those of that species, it is here relegated to synonymy. 

Populations of L. tupa also show clinal variation in pubescence. The hairs on leaves 
and stems generally become longer and softer from north to south, while the dorsal 
surface of the anther tube becomes increasingly pubescent from south to north. The 
corolla of L. tupa typically is red, though it does not change color (Weiss 1995) as in L. 
excelsa.| have seen just one specimen (Reiche s.n., Jan 1902, SGO) in which the corolla is 
yellow (“flores flavescente”; cf. Reiche 1905, 1910). 


Representative specimens. JUAN FERNANDEZ ISLANDS. Isla Masatierra: in fruticetis apricis 
collium, Bertero 12473 (CONC); Quebrada Pangal, Marticorena et al. 9172 we iM >); a Anson, 
Meyer 9580 (MO, RSA, UC); San Juan Bautista to Pangal, Pacheco & Valdeb 1 (B, CONC); Valle 
Colonial, Sparre 7 (CONC, S); colony, Skottsberg & Skot 3 183 (GB, S, UPS); between Pangal and La 
Centinela, Stuessy & Crawford 6306 (CONC, OS); between hosteria and San Juan Bautista, Stuessy & 
Crawford 6310 (CONC); path to Quebrada Pangal, Stuessy et al. 6200 (CONC, OS). 
CHILE. Prov. Cachapoal: Rancagua, Frodin 494 (UPS);Termas de Cauquenes, 3 Nov 1952, Pfister 
n. (CONC). Prov. Colchagua: Las Penas, Barrientos 1605 (CONC); Cerro Echaurrina, Montero 736 


104 SIDA 19(1) 


(CONC).Prov. Curicé: Lipimavida, Spooner & Contreras 4332 (CONC, F,WIS). Prov. Talca: Constitucion, 
Feb 1895, Philippi s.n. (SGO); Talca, 13 Oct 1897, Philippi s.n. (SGO); Constitucién, Nov 1891, Reiche s.n. 
(SGO). Prov. Linares: Rio Achibuena, Gereau & Taylor 5171 (ASU, F); Termas de Catillo, Montero 6276 
(CONC). Prov. Nuble: a Quirihue despues de Trehuaco, Schlegal 756 (CONC). Prov. Concepcion: 
Parque Hualpen, Carrasco 333 (CONC); Concepcion, DeVore 1278 (OSH, UC); Saedaen Elliot 144 
(BM, NY);Punta Hualpén, Hutchinson 248 (F,GH, UC); 1 km E of Coronel, Lammers et al.6316 (C,CONC, 
F MEXU,MU,NY);20 km S of San Pedro, Lammers et al.6323 (B, CONC, F, MU); Playa El ae Lammers 
et al. 6329 (C, CONC, F, GB, MEXU, MU, NY); 10 km W of Florida, Lammers et al. 6460 (B, CONC, F, set 
Concepcion, Landrum 8388 (F); San Vicente, Pennell 12867 (GH, NY, PH, SGO); Talcahuano, Dec 1 
Philippi s.n. (SGO); Talcahuano, Skottsberg & Skottsberg 1377 (GB, S); San Pedro, Stuessy et al. 6681 i 
Prov. Arauco: Contulmo, kel 40765 (CONC); 3 km E of Contulmo, Lammers & Baeza 6510 (CONC 
F MU); Laraquete, 20 Dec 1949, Ricardis.n. (CONC, OS);4.7 km N of bridge over Lébu, Spooner 4483 (F, 
WIS). Prov. Malleco: inane Lammers & Baeza 6508 (CONC, F, MU);9 km W of Purén, Lammers & Baeza 
6509 (F, MU); Nahuelbuto, Rahn & @dum 4735 (C); Contulmo, Jan 1902, Reiche s.n. (SGO); entre Puren 
y Contulmo, Sparre & Smith 156 (CONC, OS). Prov. Cautin: Puerto Saavedra, Aravena 30 (CONC); 
Trovolhue, Montero 7904 (CONC); Carahue, Sparre 3381 (SGO). Prov. aie Niebla, Bricker 227 (ASU); 
Valdivia, Bridges 661 (BM, E, RSA); 20 mi. NE of Valdivia, Eyerdam 10687 (F, NY, SGO, UC, WTU); Niebla, 
Garaventa 5533 (CONC); Huiecolla, Gardner & Knees 4158 (E, K); Valdivia, Gunckel 21 (CONC); 1 km W of 
hwy on road to Corral, Lammers & Baeza 6463 oe F, MU); 38 km E of Corral, Lammers & Baeza 
6464 (F, MU); 31 km E of Corral, Lammers & Baeza 6466 (F, MU); 7 km N of Chaihuin Bajo, Lammers & 
Baeza 6478 (CONC, F, MU); 10 km E of Mehuin, Lammers & Baeza 6496 (F, MU); Mehuin, Lammers & 
Baeza 6503 (B, F, MU); 6 km S of San Jose de la Mariquina, Lammers & Baeza 6506 (F, MU); Corral, 
Rudolph 6097 (VALD); Angachilla, Dec 1960, Santos & Retamal s.n. (ISC); Niebla, Schmitz 91 (VALD). 
Prov. Osorno: Alencapi, Rudolph 6091 (VALD); Tres Esteros, Rudolph 6092 (VALD), 6094 (VALD), 6095 
(VALD); La Barra del Rio Bueno, Sparre 4558 (S, SGO). Prov. Chiloé: Ancud, Pennell 12493 (F, GH, NY, 
PH), 25 Dec 1951, Pfister & Ricardi s.n. (CONC). 

CULTIVATION. U.S.A. California: Berkeley, Bracelin 1450 (GB). GERMANY: Hamburg, 1834, 
anonymous 5.n. (S). 


4. Lobelia bridgesii Hook. & Arn., J. Bot. (Hooker) 1:278. 1834. Rapuntium bridgesii (Hook. & 
Arn.) C. Presl, Prodr. Monogr. Lobel. 28. 1836. Tupa bridgesii (Hook. & Arn.) A.DC.in DC., Prodr. 
7:394. 1839. Dortmanna bridgesii (Hook. & Arn.) Kuntze, Revis. Gen. PI. 972. 1891. Type: CHILE. 
Valdivia, near El Castello de Amargos, Bridges 663 (HoLorype: KI; soryees: BMI! E! K[2]! NY! RSA! W! 
[photographs: A! F! MICH})). 


Tupa blanda D.Don in Sweet, Brit. Fl. Gard. (ser. 2) 4:p!.308. 1835. Rapuntium blandum (D.Don) C. 
Presl, Prodr. Monogr. Lobel. 27. 1836. Lobelia blanda (D. Don) Endl., Cat. Hort. Acad. Vindobon. 
1:437. 1842. Dortmanna blanda (D. Don) Kuntze, Revis. Gen. PI. 972. 1891. Type: GREAT BRITAIN. 
Kent, Sundridge Park, raised from Chilean seeds by Mr. Malleson (Lectotype here designated: D. 
Don 1835, pl.308!).As no original material was located, the plate published with the protologue 

is here designated as the lectot 
Rapuntium eae a Prodr. near Lobel. 27.1836. Lobelia lucaeana (C. Presl) A.DC.in 
DC., Prodr. 7:383.18 rtmanna lucaeana (C.Presl) Kuntze, Revis. Gen. PI.972. 1891. Tyee: GER- 
MANY. Cultum in nee botanico berolinensi, semina ex America meridionali orta diguntur, 
Sep 1835,anonymous 5.n. (HOLOTYPE: PR! 


= 


Robust perennials, 0.5-2.5 m tall; stems several from the base, normally unbranched, 
hollow, herbaceous or suffruticose, glabrous; latex white. Lamina 9-24 cm long, 1.8-4.5 
cm wide, lanceolate, glabrous; margin minutely serrulate; apex long acuminate, aristate 
or cirrhose; base sagittate, the pair of basal lobes decurrent on the stem for 12-32 mm. 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 105 


Inflorescence a 15-55-flowered raceme; bracts lanceolate, 15-60 mm long, the base sag- 
ittate, the pair of basal lobes decurrent on the stem for 6-22 mm; pedicels 14-30 mm 
long, ebracteolate, glabrous or sparsely pubescent with stiff spreading hairs. Hypanthium 
4-8 mm long,6-10 mm in diameter, hemispheric or obconic, glabrous. Calyx lobes 5-10 
mm long, 2-4 mm wide, narrowly triangular, glabrous; apex narrowly acuminate, aristate 
Corolla 25-36 mm long, pink, glabrous; tube 13-19 mm long, 2.5-4.5 mm in diameter at 
middle, suberect; lobes 12-16 mm long, 1.5-2.5 mm wide. Filament tube 14-17 mm 
long; anther tube 6-7 mm long, 2-3 mm in diameter, gray, the dorsal three sometimes 
with scattered long spreading hairs, and/or all five pubescent with short appressed hairs. 
Capsule 16-17 mm long, 13-14 mm in diameter, ovoid to subglobose. Seeds 0.5 mm 
long, 0.4 mm in diameter, broadly ellipsoid, golden tan, minutely foveolate-reticulate. 
Chromosome number n = 21 (Lammers & Hensold 1992; Lammers 1993a). 

[cones.—D. Don (1835) [as T. blanda); Hooker (1839); Lemaire (1843) [as T. blanda]; 
Loudon (1844), pl. 66, fig. 2; Lammers (1993b). 

Distribution, Habitat, and Phenology.—Endemic to the immediate vicinity of Bahia 
San Juan in Prov.Valdivia (latitude 39°49’S), in the evergreen temperate rain forest region 
(cf.Walter 1973) of south-central Chile, at elevations from near sea level up to 200 m.Not 
infrequent there, on grassy slopes, roadsides, and forest margins, sometimes in company 
with L. tupa. It is most easily found along the road that runs from the main highway 
south of Valdivia to the little coastal village of Corral. Also collected once (from intro- 
duced plants?) in Prov. Cautin and in Prov.Osorno or Llanquihue. Flowering mid-Decem- 
ber through early March. 

Discussion.—Wimmer (1953) distinguished L. blanda (including L. lucaeana) from L. 
bridgesii on the basis of subtle differences in the size and shape of the leaves. Study of 
natural populations convinced me that this variation was of no taxonomic significance. 
This species is unique in the section (and perhaps in the subgenus) for its pink corolla (cf. 
Lammers 1993b). 


Representative ae CHILE. Prov. Cautin: Conguillio, Delgado s.n. (VALD). Prov. Valdivia: Corral, 
Brooke 6985 (BM); 24 km SE of Corral, Gardner & Newton 17 (E); Corral, Gay 1469 (SGO); Amargos, 
Gunkel 29 (BM, F); Corral, Gunckel & Junge 618 (BH, CONC);La Aguada, Gunckel 5018 (NY); Castillo San 
oe a San Martin, eee (VALD); San Carlos, 1987, Krause s.n. Ne SGO); 31 km E of Corral, 

ammer & Bae 5 (B, CONC, F, MU):4 km E of Corral, Lammers & Baeza 6470 (CONC, F, GB, MEXU, 
MU); 2.5 km S of eon Lammers & Baeza 6485 (CONC, F, MU, NY); 36 km E of Corral, Lammers et al. 
7856 (CONC, F, UC); Amargos, Montero 1341 (CONC); Corral, Mar 1878, Philippi s.n. (SGO); Amargos, 
Sparre & Smith 397 (CONC, OS, Universidad de Talca); Corral, Werdermann 1938a (B,M,S); hills above 
Corral, West 4880 (GH, MO, UC). Prov. Osorno or Llanquihue: Monte Sobre, Lago Llanquihue, 1939, 
Santa Cruz s.n. (BH). 
CULTIVATION. GERMANY: Munich bot.gard.,9 Apr 1863, Kummer s.n. (M),30 Aug 1867,Kummer 
s.n. (M),5 Jul 1869, Kummer s.n. (M 


a 


PUTATIVE HYBRID 
Lobelia excelsa Bonpl. x L. polyphylla Hook. & Arn. Tupa kingii Phil. Anales Univ.Santiago 
90:189. 1895,’kingi.’ Tyee: CHILE. Valpara(so, Poppe's [Poeppig's] Hill, Dec 1868, King s.n. (ho- 
lotype: SGO-057170! [photograph: GHI)). 


106 SIDA 19(1) 


Presumably a shrub; stems woody, glabrous. Lamina 4.8-11.2 cm long, 1.3-2.4 cm wide, 
oblong, glabrous; margin minutely serrulate, particularly toward apex; apex acute; base 
cuneate. Flowers aggregated into a 18-flowered inflorescence; bracts 28-38 mm long, 
4-9 mm wide, oblong, minutely pubescent; pedicels 15-18 mm long, ebracteolate, mi- 
nutely pubescent. Hypanthium 5-6 mm long, 8-10 mm in diameter, hemispheric or 
broadly campanulate, minutely pubescent. Calyx lobes 5 mm long, 1 mm wide, narrowly 
triangular, minutely pubescent; apex acuminate. Corolla 40 mm long, apparently dark 
reddish-purple, minutely pubescent; tube 23 mm long, 2.5 mm in diameter at middle, 
curved; lobes 17 mm long, 1 mm wide. Filament tube 21 mm long, dark reddish-purple; 
anther tube 6 mm long, 2 mm in diameter, pale straw-colored, the dorsal three with 
scattered long white hairs on the surface toward apex. Fruit and seeds not seen. Chro- 
mosome number unknown. 

Discussion.—Though Tupa kingii was treated as a synonym of L. polyphylla by Reiche 
(1905, 1910) and Wimmer (1953), the type appears to represent a hybrid between that 
species and L. excelsa. It resembles L. excelsa generally, particularly in the size and shape 
of the leaves, but differs in its well-demarcated inflorescence (vs. flowers solitary and 
axillary) of darker flowers on ebracteolate pedicels, features characteristic of L. polyphylla. 
Various quantitative features of the flowers are intermediate in size: pedicels 15-18 mm 
long (vs. 7-17 mm in L. polyphylla and 12-45 mm in L. excelsa); hypanthium 8-10 mm in 
diameter (vs.4-8 mm in L. polyphylia and 8-15 mm in L. excelsa); corolla 40 mm long (vs. 
15-25 mm in L. polyphylla and 38-65 mm in L. exce/sa), with tube 23 mm long (vs. 7-17 
mm in L. polyphylla and 25-42 mm in L. excelsa) and lobes 17 mm long (vs.4—12 mm in L. 
polyphylla and 12-33 mm in L. excelsa); and filament tube 21 mm long (vs.9-15 mm in L. 
polyphylla and 29-47 mm in L. excelsa), with anther tube 6 mm long (vs. 4-7 mm in L. 
polyphylla and 8-11 mm in L. excelsa).As noted above, this specimen is the sole evidence 
| have seen of hybridization between these two sympatric species. 


ACKNOWLEDGMENTS 


Field work in Chile during 1989 (including visits to CONC, SGO, VALD, and the Univer- 
sidad de Talca) was made possible by grant #87 from the Willard Sherman Turrell Her- 
barium Fund of Miami University, Oxford, Ohio.| extend my most heartfelt gratitude to 
my many collaborators at the Universidad de Concepcidn, who accompanied me in the 
field and in general facilitated every aspect of my visits to their wonderful nation, espe- 
cially Clodomiro Marticorena, Roberto Rodriguez, Oscar Matthei, Max Quezada, Marcello 
Baeza, and Patricio Penailillo. | gratefully acknowledge the administrations and staffs of 
the following herbaria for making specimens in their care available, via loans or during 
visits: A, ASC, ASU, B, BH, BM, C, CAS, CONC, E, F, GB, GH, ISC, K, M, MA, MEXU, MICH, MO, MU, 
NY, OS, OSH, P, PENN, PH, PR, RSA, S, SGO, TEX, TUB, U, UB, UC, Universidad de Talca, UPS, US, 
VALD, W, WIS, and WTU. Finally, special thanks to my colleague Neil Harriman for his cri- 
tique of an earlier draft of this manuscript, and to Eric Knox and an anonymous reviewer 
for their helpful comments on a subsequent draft. 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 107 


REFERENCES 


BentHaM, G. 1876. Campanulaceae. Pp. 541-564. In: G. Bentham and J.D. Hooker, Genera 
plantarum, vol. 2.Reeve and Co., London. 
Biacow, N.W.(ed.) 1972. Martindale. The extra pharmacopeia, ed.26. Pharmaceutical Press, 
London. 
Bonptano, A. 1813. Description des plantes rares cultivées € Malmaison et 4 Navarre. P. 
Didot, Paris. 
CANDOLLE, A. de. 1839. Lobeliaceae. Pp. 339-413, 784-787. In: A.P. de Candolle, Prodromus 
systematis naturalis regni vegetabilis, vol. 7. Treuttel et Wurtz, Paris. 
Cavanites, J. 1801. lcones et descriptiones plantarum, vol.6. Regia Typographia, Madrid. 
CHITTENDEN, FJ. (ed.). 1923. Perennial lobelias at Wisley, 1921. J. Roy. Hort. Soc. 48:239-240, 
Don, D. 1834. Lobelia polyphylla. Leafy lobelia. P!.242.|n: R. Sweet, The British flower garden 
(ser. 2), vol. 3. James Ridgway and Sons, London. 
. 1835. Tupa blanda. Blush-flowered tupa. Pl. 308. In: R. Sweet, The British flower 
garden (ser. 2) vol. 4. James Ridgway and Sons, London. 
Don, G.1834.A general history of dichlamydeous plants, vol.3.J.G.and F. Rivington, London. 
Feuittee, L.1714, Journal des observations physiques, mathematiques et botaniques, vol. 2. 
Pierre Giffart, Paris. 
FINNIS, V. 1966. Lobelia tupa. J. Roy. Hort. Soc. 91:132 + fig. 48. 
Hitt, R.K. 1970. Pyrrolidine, piperidine, pyridine, and imidazole alkaloids. Pp. 385-429. In: 
S.W. Pelletier, ed. Chemistry of the alkaloids. Van Nostrand Reinhold, New York. 
Hoffmann J., A.E. 1997. Flora silvestre de Chile, zona araucana, ed. 4. Fundacion Glaudio 
Gay, Santiago. 
Hooker, W.J. 1833. Lobelia mucronata. Sharp-pointed Lobelia. Bot. Mag. 60:pl. 3207. 
. 1837.Lobelia polyphylla. Many-leaved lobelia. Bot. Mag. 64:pI.3550. 
. 1839. Lobelia bridgesii. Mr. Bridges’ lobelia. Bot. Mag. 65:pl. 3671. 
and G.A.W. Arnott, 1830. List of the places visited by the expedition under the 
command of Captain Beechey,R.N. Pp.i-ii.In: The botany of Captain Beechey’s voyage. 
Henry G. Bohn, London. 
Huxtey, A.J. (ed.) 1992. Lobelia. Pp. 104-106. In: The new Royal Horticultural Society dictio- 
nary of gardening, vol. 3. MacMillan, London. 
IBANEZ, J. 1955. Fanerogamia. Editorial Universitaria, Santiago 
Knox, E.B., S.R. Downie, and J.D. Paver. 1993. Chloroplast genome rearrangements and the 
evolution of giant lobelias from herbaceous ancestors. Mol. Biol. Evol. 10:414-430, 
Kunze, O. 1891. Revisio generum plantarum, vol. 2. Arthur Felix, Leipzig. 
Lammers, T.G. 1993a. Chromosome numbers of Campanulaceae. Ill. Review and integra- 
tion of data for subfamily Lobelioideae. Amer. J. Bot. 80:660-675. 
. 1993b, Lobelia bridgesii, Campanulaceae. Kew Mag. 10:70-75 + pl. 220. 
. 1994. Typification of the names of Hawaiian Lobelioideae (Campanulaceae) 
published by Wilhelm Hillebrand or based upon his specimens. Taxon 43:545-572. 


108 SIDA 19(1) 


. 1997. Phylogeny, biogeography, and systematics of the Wahlenbergia 
fernandeziana complex (Campanulaceae: Campanuloideae). Syst. Bot. 21:397-415. 
1999. A new Lobelia from Mexico, with additional new combinations in world 
Campanulaceae. Novon 9:381-389. 
and L. Giass. 1998. Morphological variation in the Lobelia polyphylla complex 
(Campanulaceae) of Chile [abstract]. Amer. J. Bot.85 (6, suppl.): 140. 
and N. HensoLp. 1992. Chromosome numbers of Campanulaceae. II. The Lobelia 
tupa complex of Chile. Amer. J. Bot. 79:585-588. 
Lemaire, C. 1843. Tupa élégant. Hort. Universel 4:259-262 + plate. 
Lewis, W.H. and M.PF. Ewvin-Lewss. 1977. Medical botany. John Wiley and Sons, New York. 
Linotey, J. 1826. Lobelia arguta. Fine-toothed lobelia. Edwards’ Bot. Reg. 12:pI.973. 
. 1830. Lobelia purpurea. Purple lobelia. Edwards’ Bot. Reg. 16:pl. 1325. 
. 1833. Lobelia tupa. The tupa-poison plant. Edwards’ Bot. Reg. 19:pl. 1612. 
Linnaeus, C. 1753. Species plantarum. Laurentius Salvius, Stockholm. 
. 1762. Species plantarum, ed. 2.Laurentius Salvius, Stockholm. 
Louoon, J.W. 1844. The ladies’ flower-garden of ornamental perennials. Willim Smith, 


London. 
Mariani R., C. 1965. Témas de hipnosis. Andrés Bello, Santiago. 
Marticorena, C.and M. Quezapa. 1985. Catalogo de la flora vascular de Chile. Gayana, Bot. 
42:1-157. 
and R. Rooricuez (eds.) 1995. Flora de Chile, vol. 1. Universidad de Concepcion, 
Concepcion. 
MattHel, O.,C. MARTICORENA, and T.F. Stuessy. 1993.La flora adventicia del archipielago de Juan 
Fernandez. Gayana, Bot.50:69-102. 
MartHews, V. 1988. Lobelia tupa. Kew Mag.5:157-161. 
Munoz P, C.1960.Las especies de plantas descritas por R.A. Philippi en el siglo XIX. Univer- 
sidad de Chile, Santiago. 
1966. Sinopsis de la flora chilena, ed. 2. Universidad de Chile, Santiago. 
Murata, J. 1992. Systematic implications of seed coat morphology in Lobelia 
(Campanulaceae-Lobelioideae). J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 15:155-172, 
.1995.A revision of the infrageneric classification of Lobelia (Campanulaceae- 
Lobelioideae) with special reference to seed coat morphology. J. Fac. Sci. Univ. Tokyo, 
Sect. 3, Bot. 15:349-371. 
Murito, A. 1889. Plantes médicinales du Chili. Exposition Universelle, Seccion Chilienne, 
Paris. 
Navas B., L.E. 1979. Flora de la cuenca de Santiago de Chile, vol. 3. Universidad de Chile, 
Santiago. 
Preirrer, L. 1874. Nomenclator botanicus, vol. 2.T. Fischer, Kassel. 
PHiuippl, R.A. 1873. Descripcidn de las plantas nuevas incorporadas ultimamente en el 
herbario chileno. Anales Univ. Chile 43:479-583. 
. 1895. Plantas nuevas chilenas. Anales Univ. Chile 90:187-191. 


LAMMERS, REVISION OF LOBELIA SECT. TUPA 109 


Pres, C.B. 1836. Prodromus monographiae Lobeliacearum. Theophilus Haase, Prague. 
Rarraur, R.F. 1970. A handbook of alkaloids and alkaloid-containing plants. Wiley- 
Interscience, New York. 
Reiche, K. 1905. Estudios criticos sobre la flora de Chile. Campanulaceas. Anales Univ. Chile 
117:206-208, 451-464, 481-482. 
.1910.Flora de Chile, vol.5. Cervantes, Santiago. 
SANTA Cruz, A. 1932.La trupa o tabaco del diablo. Revista Chilena Hist. Nat. 36:98-100. 
SANZ DE Cortazar, C. 1948. Observationes chromosomales en seis especies chilenas. Agric. 
Técn, 8(1): 28-35. 
ScHuttes, R.E., 1976. Hallucinogenic plants. Golden Press, New York. 
. 1981. lconography of New World hallucinogens. Arnoldia 41:79-125, 
.1990.The virgin field in psychoactive plant research. Bol. Mus. Paraense Emilio 
Goeldi, Sér. Bot. 6:7-82. 
and A. HorrmMann. 1980.The botany and chemistry of hallucinogens, ed.2.Charles 
Thomas, Springfield MA. 
Sims, J. 1810. Lobelia gigantea. Gigantic lobelia. Bot. Mag. 32:pl. 1325. 
. 1825.Lobelia tupa. Mullein-leaved lobelia. Bot. Mag. 52:pl. 2550 
Spooner, D.M., T.F. Stuessy, D.J. Crawrorb, and M. Siiva O. 1987. Chromosome numbers from 
the flora of the Juan Fernandez Islands. |]. Rhodora 89:351—356. 
Stace, H.M. and S.H. James. 1996, Another perspective on cytoevolution in Lobelioideae 
(Campanulaceae). Amer. J. Bot. 83:1356-1364. 

STEIN, B.A. 1987. Systematics and evolution of Centropogon subgenus Centropogon 
(Campanulaceae: Lobelioideae). Ph. D. dissertation, Washington University, St. Louis. 
Stuessy, T.F. and C. Marricorena. Orthography of some epithets honoring Bertero in the 

vascular flora of the Juan Fernandez Islands and continental Chile. Gayana, Bot.47:77- 


——* 


SweeT,R. 1827-29. Lobelia tupa. Mullein-leaved lobelia. Pl. 284. In:The British flower garden, 
vol. 3. James Ridgway and Sons, London. 

THomas, G.S. 1990, Perennial garden plants or the modern florilegium, ed. 3. Timber Press, 
Portland. 

THompson, S.W. and T.G. Lammers. 1997. Phenetic analysis of morphological variation in the 
Lobelia cardinalis complex (Campanulaceae: Lobelioideae). Syst. Bot. 22:315-331, 

Varke, W. 1874. Notulae in Campanulaceas herbarii regii berolinensis. Linnaea 38:699-735. 

ViLMorIN, R. be and M. Simonet. 1927. Nombre des chromosomes dans les genres Lobelia, 
Linum, et chez quelques autres especes vegetales. Compt.-Rend. Hebd. Séances Mém. 
Soc. Biol. 96:166-168. 

Voss, A. 1894. Campanulaceae. Pp. 560-578. In: A. Siebert and A. Voss, Vilmorins 
Blumegarterei Beschreibung Kultur und Verwendung, ed. 3, vol. 1. Paul Parey, Berlin. 

Water, H. 1979. Vegetation of the earth, ed. 2. Springer-Verlag, New York. 

Weiss, M.R. 1995. Floral color change: a widespread functional convergence. Amer. J. Bot. 
82:167-185. 


110 SIDA 19(1) 


Wimmer, FE. 1953. Campanulaceae-Lobelioideae, Il. Teil. Pp. i-viii, 261-814. In: R. Mansfeld 
(ed.), Das Pflanzenreich IV.276b. Akademie-Verlag, Berlin. 

«1968. Campanulaceae-Lobeliocideae supplementum et Campanulaceae- 
Cyphioideae. Pp. i-x, 815-1024. In: S. Danert, ed. Das Pflanzenreich IV.276c. Akademie- 
Verlag, Berlin. 


A NEW SPECIES OF TREPADONIA 
(ASTERACEAE: VERNONIEAE) FROM PERU 


Harold Robinson Hamilton Beltran 
Department of Botany M de Historia Natural 
National Museum of Natural History Universidad Nacional Mayor de San Marcos 
Smithsonian Institution Av.Arenales 1256, Apartado 14-0434 
Washington, DC 20560-0166, U.S.A. Lima U 
ABSTRACT 


pad ppositifolia (A V is described from Peru and a key is presented for 
the two species of the genus. 


RESUMEN 


Se describe Trepadonia ee a (Asteraceae: Vernonieae) de Peru y se presenta una Clave para 
las dos especies del géne 


Trepadonia was established as a genus separate from the broad concept of Vernonia 
(Jones 1980) by Robinson (1994) based on the one Peruvian species, Vernonia mexiae 
(Robinson 1981). The genus is based on the totally scandent habit and the 90°-angle 
branching of the primary branches of the inflorescence. The species also has distinctive 
racemose branchlets in the inflorescence. A second species, Trepadonia oppositifolia is 
described herein, also from Peru, differing by having opposite leaves, cymiform inflores- 
cence branchlets, and more florets in the heads. The new species occurs in southern 
Peru, and if distributions of cohabitants such as the bamboo Guadua are indicative, the 
new species may be found eventually in nearby western Brazil. 


ee Lureaeions H.Rob.& H. Beltran, sp.nov. (Fig. 1). Type: PERU: Dpto. Cuzco:Provincia 
cion, Echarati, Cashiriari-3 Well Site, 5 km S of Camisea River; 11°52'57.1 S,72°39'6.1 
vi ae m, ie forest mixed with “paca” Guadua sarcocarpa, vine, petals pale purple, clear- 
ing, 2 Sep 1998; P Nufiez, H. Beltran nie de la Colina, J. Tenteyo et al. 23967 (HoLotyee: US; 
isorype USM, CUZ). 


A T.mexiae in foliis oppositis ramulis inflorescentii cymiformis et floribus 18-23 in capitulo differt. 
Scandent, to 8-9 m high; branches striate, glabrous. Leaves opposite; petioles mostly 
1.0-1.5 cm long, pake dilated and reddish; lamina ovate, 11-13 cm long, 5-7 cm wide, 
base rounded, margins ntire to slightly undulate,apex acuminate, adaxial surface bright 


green, glabrous, sbaial surface paler green, puberulous with minute hairs, venation pin- 
nate, ca. 9-10 pairs of widely spreading secondary veins. Inflorescence rather thyrsoid- 
paniculate, with primary and secondary branching mostly spreading at 90°-angles, 
branchlets cymose. Heads separate, mostly sessile, homogamous; involucre campanu- 
late, 3-4 mm high, 4-5 mm wide; involucral bracts ca. 28, gradate in 3-4 series; outer 
bracts ovate, 2 mm long, 1 mm wide, puberulous outside, brown at apex, inner bracts 


SIDA 19(1): 111-113. 2000 


112 SIDA 19(1) 


Fic. 1. Trepadonia oppositifolia H. Rob. & H. Beltran. Live plant 


oblong-lanceolate, 4 mm long, 1 mm wide, glabrous. Florets 18-23; corollas purple, gla- 
brous, ca.5 mm long, tube 2 mm long, throat ca. 1 mm long, 1 mm wide, lobes erect, 
lanceolate,ca.2 mm long.Cypselas 2 mm long,0.5 mm wide, 10-costate, with many short 
appressed setulae; pappus bristles white, ca. 38, 4 mm long, scabrid, squamae of outer 
series ca. 0.9-1.2 mm long, scabrid. Pollen ca. 37 mm in diameter in fluid, tricolporate, 
non-lophate. 

Pararype: PERU. Deto. Cuzco: Provincia La Convencion, Echarati, Cashiriara-3 Well Site, 5 km south of 
Camisea River; 11°52'57.1 S, 72°39'6.1 W, 700 m, 2 Sep 1998, P Nunez, H. Beltran, W. Nauray, R. de la 
Colina, J. Tenteyo 23842 (CUZ, US). 

KEY TO THE SPECIES OF TREPADONIA 


la. Leaves alternate, blade oblong-ovate; branchlets of inflorecence racemiform; heads 


with 8-10 florets T. mexiae 
lb. Leaves opposite, blades ovate; branchlets of inflorescence cymiform; heads with 
18-23 florets T. oppositifolia 


Leaves of the Vernonieae are usually alternate. Opposite and verticillate leaves are com- 
paratively rare in the tribe, being most common in the Neotropical subtribe 
Piptocarphinae and the African genus Bothriocline Oliv. ex Benth. In the subtribe 
Vernoniinae, to which Trepadonia belongs, opposite or verticillate leaves have previously 
been known only in one Jamaican species of Lepidaploa and one Colombian variety of 
another species of Lepidaploa (Robinson 1999). The character is almost always variable 
within the genera in which it occurs. Only in the Andean genus Joseanthus H.Rob.,of the 


Piptocarphinae, with five species, are all the species opposite-leaved. 


ROBINSON AND BELTRAN, A NEW SPECIEIS OF TREPADONIA 


ACKNOWLEDGMENTS 
We thank F. Dallmeier and A.Alonso of the SI/MAB Program, responsible for the Biodiversity 
Assessmant and Long-term Monitoring, Lower Urubamba Region, Peru, for inviting the 
junior author to participate in the botanical expedition that collected the new species. | 
wish to thank Marjorie Knowles for work on the text and illustration preparation. 


REFERENCES 


Jones, $.B. 1980.Family Asteraceae: [Tribe Vernoniae].|n:J.F. Macbride & Collaborators. Flora 
of Peru. Fieldiana, Bot. n.s. 5:22-69. 


Rosinson, H. 1981. Six new species of Vernonia from South America. Phytologia 49: 
261-274. 
__ «1994, Cololobus, Pseudopiptocarpha, and Trepadonia, three new genera from 
South America (Vernonieae: Asteraceae). Proc. Biol. Soc. Wash. 107:557—568. 
.1999.Generic and subtribal classification of American Vernonieae. Smithsonian 
Contr. Bot. 89:i-Iv, 1-116. 


114 SIDA 19(1) 


BOOK REVIEW 


WILHELMINA FEEMSTER JASHEMSKI. 1999. A Pompeian Herbal: Ancient and Modern Medici- 
nal Plants. (ISBN 0-292-74060-3, pbk.). Univ. of Texas Press, RO. Box 7819, Austin, TX 
78713-7819,US.A.$17.95, pbk., $35.00, hbk. 123 pp., 15 color and 7 b&w photos, 36 
b&w line drawings. 


While excavating ruins at Pompeii,author Wilhelmina Jashemski noticed several workmen collect- 
ing a variety of plants for medicinal purposes."As the days passed and my workmen continued to 
collect plants, | began to wonder if the plants they were collecting for medicine were the same 
ones the ancient Romans had used for cures.” That question germinated into A Pompeian Herbal,a 
collection of 36 aah common in Pompeii juxtaposing nee medicinal uses with an- 
cient ones.Not only doe idatet ontinuity of and in the Pompeian 
area, but it also serves as an example of exhaustive and ic research. My initial skepticism of 
reading an herbal- not written by a botanist, was soon replaced with a sense of relief that we, as 
readers and researchers, are lucky enough for it be written by an archaeologist. For only an archae- 
ologist, perhaps, would pose the question and have the fortunate circumstances to be doing ar- 
chaeological excavations while simultaneously collecting ethnobotanical eae from local 
Pompeians. 
In the introduction the author explains her fieldwork and methodology and gives appropri- 
ate background information on Pompeii, local medicinal plants, and the ancient literature of Pliny 
e Elder, Dioscorides, Theophrastus and others. Gleaning data from a variety of sources is what 
ae the book sl Serene and unique. Information on the contemporary uses of me- 


dicinal plants came workmen of the site- the people, often times bringing plants to 
the author and describing preparations and what ailments the ise treat (fegato, or liver prob- 
lems—treated with Verbanum sinuatum L.—are common). For the ancient uses of the plants 


ashemski consults ancient literature, discussing both devi et and medicinal citations. This 
information is strongly supported by the scant, yet important, archaeological evidence. Excava- 
tions on sites that were covered with the resulting pyroclastic flow of Mt. Vesuvius provided invalu- 
able archaeo-botanic evidence normally unavailable in the climate of Pompeii. The lava provides 
enough heat to carbonize plant remains such as seeds, roots, fruits, and stems, however it’s unfa- 
vorable to preservation of pollen, limiting identification to family names but not of genus and 
species. Archaeological evidence revealed the planting patterns of ancient Pompeians- that 
as the workers revealed, were the same patterns they followed in their own gardens today. Most 
sles aks Hans been excavated at ee over the years have neglected the palecethnobotanic 
king this bo i all the more valuable as it provides insight into ancient Pompeian life 
that before now was obso 
Following the aoe comes the core of the book, the herbal. The design of the herbal 
is one of this book's major strong points. For each plant the author provides the scientific, English, 
and Italian names and a physical Ae of the plant and its habitat. Drawing on information 
from local villagers the author describes how the plant is prepared as a medicinal and what ail- 
ments it is used to treat. From there the author cites the ancient literature giving reference to the 
plant as a medicinal and its mythological connotations, Information on each plant is presented vis- 
a-vis a full-page black and white plant portrait giving the herbal an elegance any classicist is sure 
to appreciate. Completing the herbal is a bibliography of ancient literature and notes on citations, 
a bibliography of secondary sources, a list of peer general and Greek indices. 
The potentially Bee readership should stand testament to the book's worth. Being under 
100 pages and less than Se ee mena en lin alternative medicine, 
folklore, and Pompeii as well as the de polaris ang BIC NacOlog/st. The book is not designed 
for college courses, but should prove for those researching Old World 
paleoethnobotany.—Kevin D. Janni. 


SIDA 19(1): 114. 2000 


RHYNCHOSPORA ZACUALTIPANENSIS AND ELEOCHARIS 
MOOREL TWO NEW CYPERACEAE FROM MEXICO 


Mark T. Strong M.S. Gonzalez-Elizondo 
Department of Botany, MRC/166 Herbario Cll CIIDIR - IPN 
mithsonign Institi tion Apdo. Po al 738 
10th and Constitution Avenue Durango, ne 34000 
Washington, DC 20560-0166 U.S.A. MEXICO 
strong.mark@nmnh.si.edu sgonzalez@omanet.com.mx 
ABSTRACT 


> 


ioe zacualtipanensis M. Stron ong & S. Gonzalez, from east- 
ern Mexico, are described and illustrated. cai ee to ue: allied and morphologically 
similar aoccie are discussed. 


RESUMEN 
Se describen e ilustran Rhynchospora zacualtipanensis M. Strong y Eleocharis moorei M. Strong & S. 
Gonzalez, del oriente de México, y se discuten sus relaciones con especies cercanas y 
morfoldgicamente similares. 


Rhynchospora Vahl nom. cons. is a genus of approximately 250 species worldwide. It is 
most diverse in the Western Hemisphere, particularly warm-temperate North America 
and the neotropics. Espejo Serna and Lopez Ferrari (1997), recognized 51 taxa (including 
infraspecific taxa) from Mexico. 

Rhynchospora, a very vegetatively diverse group, contains plants that range from 
small diminutive annuals less than 10 cm tall to large rhizomatous perennials up to 3 m 
tall. The leaves are primarily basal, 2-several cauline, linear to filiform, dorsiventrally com- 
pressed, and often have scabrous margins. The inflorescence ranges from a single soli- 
tary spikelet or capitate head of spikelets at the summit of the culm to large decom- 
pound panicles of many spikelets, often composed of a terminal and a series of 1-several 
lateral, remote or contiguous, simple to compound, corymbiform or cymose partial 
panicles from the upper sheathing bracts. The floral morphology is generally uniform 
and can be characterized as follows: spikelets have spirally arranged scales, with (1-)2- 
many basal scales empty (sterile), the fertile scales are borne above these and each sub- 
tends a perfect flower, or often the terminal 1-2 scales are staminate only with an abor- 
tive pistil;each flower has (1—)2—3 stamens;the style is 2-branched or entire;and achenes 
are biconvex or lenticular and are often transversely rugose or rugulose, each bearing at 
its apex the persistent triangular, triangular-lanceolate or discoid base of the style. 

RI I Iti |.Strong, sp.nov.(Fig. 1). Tyee: MEXICO. Hivaico Stare: Zacualtipan 
District, ca.3 mi [48 km] from Zacualtipan on road to Tianguistengo, 2100 m, pine woods 
and sphagnum bags,in sphagnum, 4 Jul 1947, Moore 3327 (HoLotyre: BH; isotyres: MICH, UC, US 
3386987). Possible isotypes at G and M [These herbarium acronyms were also recorded in 


7 


SIDA 19(1): 115-122. 2000 


SIDA 19(1) 


116 


) A Hahit. B Secti £ loaf hlad 


(left) to apex (right). C. Section of leaf at sheath orifice showing junction of sheath and blade. D. Detail of terminal 


le.G. Achene 


F. Spikelet sca 


E 


STRONG AND GONZALEZ, NEW SPECIES OF CYPERACEAE FROM MEXICO 117 


H.E. Moore's collection notebook as prospective herbaria for the distribution of duplicates. 
However, recent searches made by curators at these institutions were unsuccessful]. 


Planta perennis caespitosa glabra rhizomate brevi crasso. Culmi erecti vel ascendentes, 20-60 cm 
alti, 0.5-1.7(-2) mm lati, obtuse trigoni vel subteretes. Folia basilaria et inferne caulinaria; vaginae 
eligulatae; laminae 10- =o cm lon ngae, OS: -2 mm latae vel 3 mm 1 latae ubl Pialide, v- formae vel 
subcomplicatae vel sul | ulata 
paucispiculata, st terminalis major, superiores 2 approximatae laterales 1-2, pedunculatae, 
pedunculi ad 6 ongi. Spiculae 3.5-4.2 mm longae, 1.2-1.5(-1.6) mm latae, anguste ovoideae 
vel aor ee hie 3-4-floriferae, 2 nucigerae; squamae 5-7, ovatae vel ovato-ellipticae, 
mucronatae vel breviter ari ti bi fertil 5-)2.6-3.1 mm longae, 
2-2./(-3) mm latae, squamae steriles breviores. Stamina tres, antheris 1.5—-3 mm longis. Stylus 
profunde bifidus. Achaenia 1.8-2.1 mm longa (sine styli basi), 1.2-1.4 mm lata, ee biconvexa 
spadicea, leviter transverse tee. styli basis anguste triangularis, 0.8-1 mm longa, 0.3-0.7 mm 
lata; setae 6, antrorse barbatae, basi saepe setosae, achaenium leviter breviores ad ee 


Caespitose perennial; rhizome short, thickened and knotty. Culms erect or ascending, 
20-60 cm tall, 0.5-1.7(-2) mm wide, obtusely trigonous to subterete, finely ribbed, often 
channelled along ide distal Ne glabrous Leaves basal and lower cauline;sheaths 


as 


eligulate, coarsely veined abaxially, green to pale brown, often reddish brown proximally, 
glabrous, the inner band membranous, finely veined, reddish brown, with concave ori- 
fice at apex; blades somewhat stiff, 10-30 cm long, 0.5-2 mm wide (to 3 mm wide when 
flattened) widely to narrowly v-shaped or crescentiform in cross section, sometimes 
subinvolute or tightly folded, often subtrigonous towards apex, finely ribbed abaxially, 
smooth adaxially, at least medially, green, glabrous, the margins antrorsely scabrous or 
sometimes essentially smooth proximally. Inflorescence composed of a terminal and 1- 
2, small, lateral cymose-fasciculate partial panicles, the lateral panicles on slender pe- 
duncles to 6 cm long, the lowest panicle remote from the two upper subcontiguous 

luncles flattened, finely eed anole) scabrous on margins, glabrous; bracts 
oftiic panicle branches setaceous, t ling the peduncles, particularly the lower, 


leaf-like; spikelet pedicels subtrigonous or SMEG CORUES in cross section, the margins 
often antrorsely scabrous; spikelets narrowly ovoid to ovoid-lanceoloid, 3.5-4.2 mm long, 
1.2-1.5(-1.6) mm wide, with 5-7 scales, 3(-4)-flowered, maturing 2 achenes, the scales 
spreading with maturing achenes;scales ovate to ovate-elliptic, indistinctly finely veined, 
glabrous, margins entire, the midcosta prominent, pale brown to brown, prolonged be- 
yond the acute to acuminate apex as a mucro or short awn; fertile scales (2.5-)2.6-3.1 
mm long, 2-2.7(-3) mm wide; sterile scales 2-3 at base of spikelet, smaller than fertile 
ones, 2-2.2(-2.5) mm long, 0.8-1.2 mm wide. Flowers bisexual; stamens 3, the anthers 
1.5-3 mm long, basifixed, thecae parallel, longitudinally dehiscent, apiculate at apex; style 
2-branched from below middle, ca. 2/3 length of style, the branches glabrous, minutely 
scaly;achenes biconvex, obovate or obpyriform, achene body 1.8—2.1 mm long (exclud- 
ing style base), 1.2-1.4 mm wide, light brown to yellowish brown, finely transversely ru- 
gulose; style base narrowly triangular, 0.8-1 mm long, 0.3-0.7 mm wide, pale brown, of- 
ten truncated at apex; bristles 6, antrorsely barbed, reddish brown, often setose at very 


118 SIDA 19(1) 


base, slightly shorter than to equaling or exceeding the length of the achene body with 
several sometimes equaling the tip of the style base. 

Habitat and distribution —Rhynchospora zacualtipanensis is known only from the 
type collection made in the southern extent of the Sierra Madre Oriental mountain range 
at approximately 20° 43' 00"N, 98° 39' 00"W, ca.4.8 km north of the town of Zacualtipan. 
It occurs in pine woods and sphagnum bogs at about 2100 m. 

Rhynchospora zacualtipanensis is a species of R. subgenus Rhynchospora. It keys to 
R. section Stenophyllae Kukenthal in Kukenthal’s (1950) monumental worldwide treat- 
ment of the subfamily Rhynchosporideae. Section Stenophyliae included Gale's (1944) 
Rhynchospora section Rhynchospora series Glaucae (C.B. Clarke) Gale (in part), and series 
Rariflorae Gale (in part). Raynchospora zacualti to be best placed in R. sec- 
tion Glaucae C. B. Clarke as circumscribed bi Guaglianone (1979). The obovate or 
obpyriform, yellow-brown achene, with a longitudinally striate surface interrupted by 
transverse corrugations (finely transversely rugulose) is characteristic of species in that 
section. However, the narrow (0.5-2 mm), stiff, thickened, often v-shaped or crescentiform 
leaf blades of R. zacualtipanensis are uncharacteristic of the wider (1-5 mm), flattened 
leaf blades typical of species in R.section Glaucae.R. zacualti is can be distinguished 
at once by this feature from closely related species such as R. brownii ssp. americana 
Guaglianone,R. pungens Liebmann, and R. rugosa (Vahl) Gale. Generally, R.zacualtipanensis 
has longer bristles and a longer style base on R. brownii and R. pungens. The bristles of 
R. zacualtipanensis typically equal or exceed the length of the achene body with several 
sometimes equaling the tip of the style base, while in the latter two species, there are 
generally several that are shorter than the achene body and rarely do any equal the tip 
of the style base. The style base of R. zacualtipanensis ranges from 0.8-1 mm in length 
while those of R. brownii ssp.americana, R. pungens, and R. rugosa range from 0.5-0.8 mm 
in length. In a treatment of Mesoamerican Rhynchospora by Thomas (1992), R. 
zacualtipanensis falls out at couplet 79[b] (R. rugosa) in the part of the key that treats 
species of R. section Glaucae. R. zacualtipanensis can be distinguished from the wide- 
ranging R. rugosa by the morphology of the leaf blades (as previously discussed) and 
further distinguished by its shorter spikelets (3.5-4.2 mm vs.3.5—5 mm long) and smooth 
margins of the style base (often antrorsely scabrous proximally in R. rugosa). 

Eleocharis R. Brown is a widely distributed genus of ca. 200 species with 43 species 
recorded by Espejo Serna & Lopez Ferrari (1997) from Mexico. The taxonomy of the ge- 
nus is difficult,in part because there are very few morphological characters represented, 
e.g., the inflorescence is reduced to a single spikelet and lacks foliar involucral bracts; 
and leaves are all basal and reduced to tubular sheaths (Gonzalez & Peterson 1997). 


Eleocharis moorei M. Strong and S. Gonzalez, sp. nov. (Fig. 2). Type: MEXICO. Hinatco Stare: 
District Zacualtipan, ca. 3 mi [4.8 a from Zacualtipan on road to Tianguistengo, 2100 m, 
pine woods and sphagnum bogs, in sphagnum, 4 Jul 1947, H.E. Moore, Jr. 3332 (HoLotyee: BH; 
isorypes: MICH, UC, US 2594748 ne isotypes at B,G,and M [These herbarium acronyms 
were also recorded in H.E. Moore's collection notebook as prospective herbaria for the dis- 


aS 


119 


STRONG AND GONZALEZ, NEW SPECIES OF CYPERACEAE FROM MEXICO 


h 


J 


from the type). A. Habit. B. Sections of 


summit of leaf sheath with acute, mucronate apex (right) and ventral summit of leaf sheath with slightly concave 


\ 


iM Strong & S. Gonzalez (drawn 


Fic. 2. El 


+ rk 


+ 


L 


+] £ 


+ 


t). D. Achene. E.C 


<e (left), C Inf] 
\ / 


* 


point. 


120 SIDA 19(1) 


tribution of duplicates. However, recent searches made by curators at these institutions were 
unsuccessful]. 
Planta perennis. Rhizoma ca. (1.5—-)2.5-3.5 mm crassum, repens vel oblique ascendens cum squamis 
atropurpureis instructum.Culmi erecti,4-20 cm alti, (0.2-)0.4-1 mm latae;apice vaginis subtruncatis 
ve SUDO IAS vel oes Spiculae 3.5- - mm longae, 2.5-3 mm latae, ovatae vel lanceolatae, 


lystichae, 2-2.6 mm longae, carinatae, ovatae 

oblongae, seta maturitate divergentibus ad apicem. sarin tres, antheris (0.6-)1.1 mm 
ee Stylus trifidus. Achaen A 6mm longa eal styl basi), 0.75—1 mm lata, a obovata vel 
obpyriformia, obtuse trigona vel k m abaxialem cristata ostatis, lutea 
vel rubro-brunnea, conspicue reticulata; styli basis pyramidal deltata, leviter complanat, 
basaliter crassa trilobata leviter decurrens; setae 4—/, ret ,ach | el 


superantes. 


Perennial from an elongated, creeping or slightly ascending rootstock ca. (1.5—)2.5-3.5 
mm thick, covered by conspicuous, elongate, sheath-like, dark purple, fibrous scales. Culms 


caespitose, or sometimes solitary along the rhizome, erect, 4-20 cm tall, (0.2-)0.4-1 mm 
wide, angular to terete, soft, flattened when dried, finely sulcate, pale green;sheaths mem- 


branous, purple, or stramineous and purple-striolated at the base, green to translucent 
distally, subtruncate to slightly oblique or acute at apex dorsally, emucronate or with a 
0.1-0.2 mm long mucro, the orifice margin not or slightly darkened; upper sheaths with 
the ventral orifice concave; lower sheaths with the ventral orifice truncate to somewhat 
convex. Spikelets ovoid to ovoid-lanceoloid, 3.5—-9 mm long, 2.5—3 mm wide, usually acute 
to subacuminate at apex; rachilla 0.3-0.4 mm wide, 9-35-flowered; scales polystichous, 
2-2.6 mm long, 0.5-0.8 mm wide in lateral view, divergent distally, membranaceous 
subtranslucent, ovate to oblong, obtuse at apex, carinate, with a narrow pale green to 
stramineous midvein, the sides dark purple to black distally, the margins and apex very 
narrowly hyaline, (at most 0.1-0.2 mm wide); 1-2 basal scales shorter, 1-2.3 mm long, 1— 
1.8 mm wide, ovate, obtuse, ecarinate, with a broad midvein of the same color and ap- 
pearance as the culm, the sides purple, black, or hyaline, and the margins and apex hya- 
line, 0.3-0.5 mm wide; flowers bisexual; stamens 3, the anthers (0.6-)1.1 mm long, the 
connective shortly prolongated at apex; style 3-branched. Achene 1.4—1.6 mm long (in- 
cluding the style-base), 0.75-1 mm wide, very obtusely triangular or biconvex with a 
costula on the abaxial face, the angles costulate, widely obovate to obpyriform, yellow to 
reddish-brown, finely and shallowly cellular-reticulate, often conspicuously so, the cells 
rectangular or nearly square, longitudinally arranged; style base pyramidal to deltoid, 
dorsally subflattened, 0.4-0.5 mm long and as wide at base, sometimes slightly wider 
than the apex of the achene, trilobate at the base, the lobes slightly decurrent on the 
angles of the achene, whitish to brown, sometimes with a narrow ridge at junction with 
the body of the achene; bristles 4—7,reddish brown or yellowish, retrorsely barbed 1/2 to 
2/3 their length, equaling to exceeding the achene. 


Paratype. MEXICO. VERACRUZ S tera al sur de Huayacocotla, 17 km del borde con Hidalgo, 2100 
m, Bosque de encino, suelo arcilloso, hierba 5 cm, perenne, regular, 13 Jul 1977,J.Fay and J.l. Calzada 
883 (F, US 2879357) [distributed as Eleocharis acicularis (L.) R.Br). 


STRONG AND GONZALEZ, NEW SPECIES OF CYPERACEAE FROM MEXICO 121 


Habitat and distribution.—Eleocharis moorei is known from only two localities that are in 
close proximity to each other in the southern extent of the Sierra Madre Oriental moun- 
tain range in the states of Hidalgo and Veracruz. These lie at approximately 20° 33' 00"N, 
98° 29' 00"W to 20° 43' 00"N, 98° 39' 00"W, the Hidalgo locality near Zacualtipan being 
approximately 25-30 km northwest of the Veracruz locality near Huayacocotla. £. moorei 
occurs in sphagnum bogs in pine woods, and in oak forest, at about 2100 m. 

The name of the new species honors its collector, Harold Emery Moore, Jr. (1917- 
1980), world renowned Arecaceae specialist, professor, and director of the L.H. Bailey 
Hortorium at Cornell University, Ithaca, New York from 1960-1980. 

Eleocharis moorei belongs to E. series Eleocharis (E. series Palustriformes of Svenson 
1929, 1932, 1939, 1957), subseries Truncatae Svenson, in the subgenus Eleocharis. How- 
ever, its achene with costulate angles and trilobate style base distinguishes it from all 
other species in the group. Achenes of £. tricostata Torrey are prominently costulate, but 
in cross section they are trigonous, while those of E. moorei are very obtusely triangular 
or biconvex with a costula on the abaxial face. The achenes of E. tricostata also differ in 
bearing a much shorter and narrower style base that is neither trilobate nor decurrent, 
and they lack bristles. Furthermore, the sheaths and scales of E. tricostata differ as well in 
being lighter-colored, and the spikelets are generally longer and obtuse to subrounded 
at apex while those of £. moorei are acute to subacuminate. £ moorei differs from E. 
compressa Sullivan in having scale tips not whitened and attenuate; larger achenes with 
a different tubercle; and more abundant bristles, which are longer and coarser. From E. 
montevidensis Kunth it differs in the elongate, sheath-like, dark purple and fibrous scales 
covering the rhizome; the soft, flattened culms; the membranous sheaths; the spikelets 
usually acute; the larger, biconvex or very obtusely triangular achene with costulate angles; 
and the big tubercle which is trilobate at the base. From the variable E. tenuis (Willdenow) 
Schultes, E. moorei differs in the coarser, flattened culms; the lowest spikelet scale being 
shorter than the middle scales; the larger, biconvex to obtusely trigonous achenes with 
much less pronounced reticulation; and the larger and coarser bristles. 


ACKNOWLEDGMENTS 


We would like to thank Dan Nicolson (US) and Paul Goetghebeur (GENT) for reviewing 
the manuscript;an anonymous reviewer for helpful comments that improved the manu- 
script; Bob Dirig (BH) for collection and biographical information on H.E. Moore; curators 
at B, G, M, MICH, and UC for searching their collections for H.E. Moore specimens; and 
Cathy Pasquale for the fine drawing of Rhynchospora zacualtipanensis. 


REFERENCES 


Espejo Serna, A.and A.R. Lopez Ferrari. 1997.Las Monocotiledéneas Mexicanas. Una Sinopsis 
Floristica. 1. Lista de Referencia, Parte V. Cyperaceae. Consejo Nacional de la Flora de 
Mexico, A.C., Universidad Aut6noma Metropolitana, y Consejo Nacional para el 
Conocimiento y Uso de la Biodiversidad. México, DF. 


122 SIDA 19(1) 


Gate, S. 1944. Rhynchospora, section Eurhynchospora, in Canada, the United States, and 
the West Indies. Rhodora 46:255-278. 
Gonzatez Euzonpo,M.S.and PM. Peterson. 1997.A classification of and key to the supraspecific 
taxa in Eleocharis (Cyperaceae). Taxon 46:433-449,. 
GUAGLIANONE, R. 1979. Sobre Rhynchospora rugosa (Vahl) Gale (Cyperaceae) y algunas 
especies afines. Darwiniana 22:255-311. 
KUKENTHAL, G. 1950. Vorarbeiten zu einer monographie der Rhynchosporideae. Bot. Jahrb 
75(1):90-126; (2):127-195. 
Svenson, H.K. 1929. Monographic studies in the genus Eleocharis. Rhodora 31:121-135. 
. 1932. Monographic studies in the genus Eleocharis. Rhodora 34:193-203; 
215-227. 
. 1939.Monographic studies in the genus Eleocharis. Rhodora 41:1-13; 43-77. 
. 1957. Eleocharis (Cyperaceae). In: North American Flora 18:509-540. 
Tuomas, W.W. 1992. A synopsis of Rhynchospora (Cyperaceae) in Mesoamerica. Brittonia 
44:14-44. 


DISTINCTION BETWEEN VITIS BLANCOI AND 
V. CINEREA VAR. TOMENTOSA (VITACEAE) 


Barry L.Comeaux Jiang Lu 
1634 Hwy 356 Center for EUG Sciences 
Sunset, LA 70584, U.S.A. Florida Agricult / ‘| | University 


Tallahassee, FL 32307, U.S.A. 


ABSTRACT 


Vitis blancoi Munson as treated by Munson included two, unrelated taxa with separate distribu- 
tions. Plants from southern Mexico are treated as V. blancoi Munson emend. Comeaux, and are 
assigned to series Occidentales Munson. The northern group of plants from northern Mexico and 
extreme southern Texas belong to series Cinerascentes Planchon, and are here designated as V. 
cinerea (Engelm.in Gray) Engelm. ex Millardet var. tomentosa (Planchon) Comeaux. Revised descrip- 
tions and specimen citations are provided. 


RESUMEN 


Vitis blancoi Mason tal como We aratgae por Munson incluye dos taxa no relacionados con 
distribuc | Mexico son tratadas como V. blancoi Munson emend. 
Comeaux, y se asignan a las serie Occidentales Munson. El grupo norteno, de plantas del norte de 
Mexico y extremo sur de Texas, pertenece a la serie Cinerascentes Planchon,y se designa aqui como 
V. cinerea (Engelm. in Gray) Engelm. ex Millardet var. tomentosa (Planchon) Comeaux. Se ofrecen 
descripciones y citas revisadas de especimenes. 


Thomas Volney Munson Munson (1843-1913) was the recognized authority of his day 
on the indigenous species of North American Vitis and respect for his knowledge has 
continued into recent times (Renfro 1983).Munson studied the North American species 
of Vitis for nearly fifty years and through these efforts he produced a comprehensive 
treatment of the genus contained in his classic work, Foundations of American Viticulture 
(Munson 1909), 

Despite Munson’'s immense knowledge of the genus his interpretation of V. blancoi 
(Munson 1909)included two unrelated taxa with separate distributions. The plants from 
northern Mexico and extreme southern Texas belong to series Cinerascentes Planchon 
and the more southerly distributed individuals belong to series Occidentales Munson. 

The objectives of this study were: (1) to provide a chronological historical account 
of the taxa involved; (2) to emend Munson’s description of V. blancoi; (3) to thoroughly 
describe and provide justification for accepting as distinct V. cinerea var. tomentosa 
(Planchon) Comeaux,a currently unrecognized taxon from northern Mexico and extreme 
southern Texas; (4) and to document the distributions of these taxa. 


NOMENCLATURAL REVIEW 
Munson (1890a; 1890b) first reported obtaining specimens of V. blancoi in 1887 from 


SIDA 19(1): 123-131. 2000 


124 SIDA 19(1) 


Luciana Blanco of Guadalajara, Mexico, for whom he named the species. Blanco discov- 
ered it growing along streams in the nearby Sierra Madre Mountains in the southern 
state of Jalisco and repeatedly sent Munson specimens of the plants. 

Munson (1909) provided an elaborate description of V. blancoi; along with a photo- 
graph of a specimen labeled as from Guadalajara. Significantly, Munson stated that in 
1895 he received additional specimens from C.M. Stuart collected near Montemorelos, 
in the state of Nuevo Leon in northern Mexico. Vitis blancoi was placed in series 
Cinerascentes Planchon by Munson, since he felt it was closely related to V. caribaea DC 
(V. tiliifolia Humb.& Bonpl.ex Roem. & Schult.), also assigned to that series. 

Subsequent treatments of V. blancoi have varied regarding its status as a species 
and classification within the rank of series. Bailey (1895-97) interpreted V. blanco as syn- 
onymous under V. tiliifolia, but he later (Bailey 1934) recognized it as distinct and related 
to series Labruscoideae Planchon, which included V. labusca L. and other large-fruited 
(15-25 mm dia.) species. Standley (1920-26) also considered V. blancoi as a synonym of 
V. tiliifolia, while Galet (1988) treated V. blancoi as distinct and placed it along with V. 
tiliifolia in series Caribaea Galet. 

Bailey (1934) made a substantial contribution towards clarifying the correct status 
of V. blancoi.He pointed out that Planchon’s (1887) description of V. berlandieri, was based 
on two forms. Plants with glabrous leaves as in Berlandier 2412 (holotype PH!) from New 
Mexico or Texas were treated as the typical form, V. berlandieri var. berlandieri, and plants 
with tomentose leaves as in Berlandier 3116 (WIS!) from the state of Nuevo Leon were 
referred to var. tomentosa Planchon. The latter trinomial has not been treated in any 
subsequent publications. Significantly, Bailey suggested that the tomentose form might 
be V. blancoi,and, also mentioned that in the Munson distribution of specimens in 1888 
another name was used, but he did not provide to his readers the earlier name. 


MATERIALS AND METHODS 


Field work by the senior author was carried out in 1986 and 1994 in Texas and in 1986, 
1987 and 1990-92 in Mexico with collections made in the following states: Coahuila, 
Colima, Guerrero, Jalisco, México, Michoacan, Nuevo Leon, Oaxaca, Puebla, San Luis Potosi 
and Tamaulipas. Herbarium studies were conducted from 1980-92 during visits to the 
following: BRIT, JAL, JALU, MEXU, MO, TAMU, UF and UT. 


RESULTS AND DISCUSSION 


Recent studies revealed that the northern and southern plants treated by Munson as V. 
blancoi are, in fact, two unrelated taxa with separate distributions. The southern plants 
are separated morphologically from the northern taxon (Table 1) by their relatively large 
stipules (2-5 mm long), which are often obscured by arachnoid trichomes. Also, indi- 
viduals in the states of Colima (Comeaux 4207, 5074, 5075 and 5076 BRIT), Jalisco (Comeaux 
5078 BRIT) and Puebla (Bruff 1577 MEXU) have leaves with rufescent pubescence, while 
all collections observed of the northern taxon have only white pubescence. Large stipules 
indicate that this species is not related to V. tiliifolia and other members of series 


COMEAUX AND LU TOMENTOSA 125 


Taste 1. Comparison of certain characters between V. blancoi Munson and V. cinerea var. tomentosa 
(Planchon) Comeaux. 


Characters V. blancoi V. cinerea var. tomentosa 


I, Leaves pubescent 


adaxial surface pr, gt?,.gs gt, gs 
abaxial surface eM lcamcloua og ts, pb 
2. Pubescence color white to rusty white 
3. Stipule length 2-5mm 1-3 mm 
4. Pistil length 1.5mm 1.0mm 
5. Fruit size 1-seeded 1-seeded 
(diameter in mm) average 7.4 average 6.7 
range 6.0-10.0 range 5.5-8.0 
sample size 42 sample size 37 
2-seeded 2-seeded 
average 9.4 average 7.8 
range 7.0-12.0 range 6.0-9.0 
sample size 41 sample size 19 
3-seeded 3-seeded 
average 10.1 average 9.1 
range 9.0-12.0 range 8.5-10.0 
sample size 23 sample size 4 
4-seede 
average 10.9 
range 9.0-13.0 
sample size 13 
eede a 
iia 
ae size | 


' pr = puberulent 


ntos 
eb ne 


Cinerascentes, which have short stipules (approximately 1-3 mm long). The fruit size (6— 
13. mm dia.) shows that it is not a relative of V. labrusca and other large-fruited (15-25 
mm dia.) species of series Labruscoideae. Instead, the above characters and others, such 
as relatively early flowering time, stems round in cross section, and medium size fruit 
ripening in midseason, suggest that the southern taxon referred to as V. blancoi has clos- 
est affinities with the western series Occidentales Munson. 

The northern Mexican vines considered by Munson as V. blancoi are, as suggested 
by Bailey (1934), the same as Berlandier 3116, which was designated by Planchon as V. 
berlandieri var. tomentosa Planchon.|n Munson’s (1909) description of V. blancoi most of 


126 SIDA 19(1) 


the attributed characters apply to both northern and southern plants, however, features 
such as small stipules and tiny flowers (see pistil length) are found only in the northern 
grapes (Table 1),and are characteristic of series Cinerascentes. The northern taxon differs 
from other members of series Cinerascentes by a syndrome of characters including: to- 
mentose abaxial leaf surfaces, a general absence of short, straight trichomes on leaves 
and stems, entire leaf margins, U- or lyre-shaped basal sinuses, and relatively short fruit 
clusters (5.6-14.7 cm) It is the overall combination of characters that delimits this taxon, 
and the absence of one or two characters in an individual does not exclude it from the 
group. 


TAXONOMIC TREATMENT 


Vitis blancoi, as first published by Munson (1890a; 1890b), included plants from southern 
Mexico having large stipules (2-5 mm long) and relatively large fruit (6-13 mm dia.).This 
taxon initially named V. leucobrya by Munson, as evidenced by a specimen (Munson s.n. 
UF!) distributed in 1888, was later renamed V. blancoi as seen on a specimen from 1889 
(Munson s.n. MO!). The binomial V. leucobrya, is not valid since it was never published; 
therefore V. blancoi is the correct name. 

Munson’s (1909) description of V. blancoi also included plants from northern Mexico 
and southern Texas having small stipules (1-3 mm long) and relatively small fruit (5.5— 
10.0 mm dia). Planchon (1887) named this taxon V. berlandieri var. tomentosa. In view of 
the recent treatment (Moore 1992) of V. berlandieri as V. cinerea var. helleri (Bailey)Moore, 
and its intergradation with the northern group of plants treated by Munson as V. blancoi 
in the Del Rio, TX, area, this taxon should be treated as V. cinerea (Engelm.in Gray) Engelm. 
ex Millardet var. tomentosa (Planchon) Comeaux. 


KEY TO SELECTED TAXA 


. Stipules 2-5 mm long; pistils 1.5 mm long; southern Mexico V. blancoi 
. Stipules 1-3 mm long; pistils 1.0 mm long; northern Mexico and United States 
2. Leaves tomentose on abaxial surfaces, entire to minutely toothed, basal sinuses 
U- or lyre-shaped;leaves and stems without short (0.2 mm long), straight, pointed 
trichomes; inflorescences (including peduncles) 5.6—-14.7 cm long; stems nearly 
terete in cross section V. cinerea var. tomentosa 
2. Leaves pubescent to glabrous on abaxial surfaces, toothed, basal sinuses V- 
shaped; stems with or without short, straight, pointed trichomes; inflorescences 


par 


= 


(including peduncles) 5.5-20 cm long; stems angular in cross section 3 
Leaves glabrous to glabrate on abaxial surfaces; berries glaucous V. cinerea 
var. helleri 

3. Leaves pubescent on abaxial surfaces; berries glaucescent ____ V. cinerea var. cinerea 


Vitis ue Munson emend. Comeaux, Amer. Forests 3:374-375. 1890; USDA Pom. Bull. 
1890. Type: MEXICO. Jauisco: From the Sierra Madre Mountains 30 mi from Guadalajara, 
ee by Luciana Blanco, 1887, Munson s.n. (Neoryee, here designated: MO!). 


Vines to 15 m,stems on current season growth typically tomentose (glabrous in certain 
individuals from high elevations) faintly striated; branchlets terete; internodes 2.3-10. cm 


COMEAUX AND LU, VI TOMENTOSA 127 


long; nodes not encircled with red pigmentation; pith interrupted at nodes by a 
diaphgram 2-3 mm thick; lenticels absent; growing tips normally tomentose varying to 
pubescent, with white, tan or rufescent arachnoid trichomes, not enveloped by young 
leaves.Leaves cordiform to long-cordiform or nearly deltoid, usually 3-lobed, with lateral 
lobes acute to acuminate, infrequently divergent, apex acute to long-acuminate, base 
cordate to nearly truncate with the basal sinus varying from U-shaped, to V-shaped and 
also lyre-shaped, lateral sinuses acute; margins serrate to nearly entire, with teeth to 3 
mm long (typically 2 mm long), occasionally ciliate, with veins frequently extending be- 
yond the teeth, midrib with 5-8, usually 6 pairs of prominent veins; lamina glabrous to 
puberulent on adaxial surfaces, felted, tomentose to pubescent (puberulent on high 
elevation specimens from Morelos) on abaxial surfaces, with the pubescence consisting 
of mostly arachnoid trichomes, sometimes including simple, straight, pointed trichomes 
that are ordinarily confined to primary veins, 4.2-16.5 cm wide, 7.4-20.5 cm long; peti- 
oles tomentose to puberulent, 2.6-10.5 cm long; stipules clear to brown, especially at 
the base, sometimes obscured by dense pubescence, 0.5-1 mm wide, 2-5 mm long, 
caducous. Tendrils and inflorescences absent every third node, tendrils bifurcate or tri- 
furcate, to 18 cm long. Inflorescences 2.2-9.5 cm long, peduncles 1.9-4.8 cm long, shoul- 
der 2.3-3.5 cm long, sometimes absent or replaced by a tendril. Flowers of the function- 
ally pistillate plants with pistils 1.5 mm long. Fruit a berry, black, glaucous, 7-13 mm in 
diameter, with a pleasant flavor when fully ripe. Seeds brown, pyriform, 3.0-6.0 mm long. 
Distribution.—Widely distributed in the southern deciduous forests and the pine/ 
evergreen oak forests (De Miranda 1989) of southern Mexico south of the twenty-sec- 
ond parallel,in the states of Colima, Guerrero, Jalisco, México, Michoacan, Morelos, Oaxaca 
Puebla and San Luis Potosi. Typically found along streams and similar sites at high eleva- 
tions (1,050-2,450 m). 
Above description based on the following specimens: MEXICO. Colima: 2.6 km S of Jalisco state 
line via Hwy 55, 1,300 m, 29 Jun 1986, Comeaux 4207 (BRIT); 23.7 km N jct. Hwy 55 to Manzanillo via 
toll Hwy 55 to Guadalajara, 1,150 m, 27 Jul 1991, Comeaux 5074, 5075, 5076 (BRIT).Guerrero: 17.1 km 
W jct. Hwy 95 at Chilpancingo, 1,550 m, 18 Jul 1992, Comeaux 5189 (BRIT); 18.9 km W jct. Hwy 95 at 
Chilpancingo, 1,500 m,18 Jul 1992, Comeaux 5190, 5197 (BRIT); 16 km S of Taxco via Hwy 23, 1,200 m, 
18 Jul 1992, Comeaux 5193 (BRIT); Mpio. Chilpancingo de los Bravos, a 28 km al W de Chilpancingo, 
27 Mar 1982, Martinez 226 (MEXU); Mpio. Cutzamala de Pinzon, 1 km al E de Ventarron, 17 Mar 1973 
600 m, Medrano 5593 (UT);W ince sore Oct 1944, Sharp 441413 (MEXU). Jalisco: Nevado 
de Colima, below La eres m, 20 Nov 1968, Boutin and Brandt 2378 (MEXU); Mpio. of Autlan de 
Navarro, 2,160 m, 17 Aug 1980, Breedlove Le (MEXU); 1 km N of Colima state line via old Hwy 44, 
1,200 m, 27 Jul 1991, Comeaux 5078 (BRIT); Charandas, Mazamitla, 2,400 m, 14 May 1972, Luna 3176 
(MEXU); Cerro Viego, vereda al de la Bola del Viego, Mpio. Jocotepec, 2,450 m, 24 Apr 1986, Machua 
550 (JAL); Cerro Viejo, cauce enfrente de Huejotitan, Mpio. Jocotepec, 2,050 m, 2 May 1986, Machua 
1000 (JAL);6 km N de Huapala, Mpio. de Jilotlan, 1,390 m,5 Apr 1988, Mendoza 3708 (MEXU); 3 km al 
so de las Coloradas, Mpio. de Jilotlan, 1,720 m, 5 Apr 1988, Mendoza 3730 (MEXU); Near eee 
2,100 m, 2 Aug 1985, Rodriquez and liga 167 (MEXU). México: Valle de Bravo, 25 May 1971, Boege 
1806 (MEXU);9.8 km N of Guerrero st state line via Hwy a 1,550 m, 18 Jul 1992, res eens (BRIT); 
10.7 km N of Guerrero state line via Hwy 55,18 Jul 199 neaux 5195 (BRIT);N of Ixtapan on toll rd. 
4.2 km S of jct.rd.to Villa Guerra, 2,000 m, 18 Jul 1992, (eS S06 (BRIT); “ of Ixtapan on toll rd. 


128 SIDA 19(1) 


3.8 km S of jet. rd. to Villa Guerra, 2,100 m, 18 Jul 1992, Comeaux 5197 and 5198 (BRIT); N of Ixtapan 
on toll rd.1.6 km S$ jct.rd.to Villa Guerra, 2,300 m, 18 Jul 1992, Comeaux 5199 (BRIT); N of Ixtapan on 
toll rd.0.8 km S jet. rd.to Villa Guerra, 2,300 m, 18 Jul 1992, Comeaux 5200 (BRIT); N of Ixtapan on toll 
rd. 1.0 km N jct.rd.to Villa Guerra, 2,300 m, 18 Jul 1992, Comeaux sl (BRIT); Nepantla, 2,000 m, 17 
May 1953, Matuda 28368 (MEXU); Amatepec y cercanias, 2,000 m, 11-12 Apr 1954, Matuda 30668 
(MEXU). Michoacan: Rincon, 4 Apr 1909, Arsene 3009 (MEXU); : ens Testarazo, cerca 
Tacambaro, 2,100 m, 28 May si a 7595 (MEXU); Cucuchuchu, Mpio.de Tzintzuntzan, 2,2 
m,23 Apr 1979, Caball o.de Quiroga, 24 Mar 1986, ie. 917 (MEXU);5 km E . 
Zitacuaro via Hwy 15, 2,000 m, 24 jul 1991 ,Comeaux 5046 (BRIT); 4.2 km E of Zitacuaro via Hwy 15 
2,000 m, 24 Jul 1991, Comeaux 5047 (BRIT); 3.8 km E of Zitacuaro via Hwy 15, 2,000 m, 24 Jul 
1991,Comeaux 5048 (BRIT); 3.5 km E of Zitacuaro via Hwy 15, 2,000 m, 24 Jul 1991, Comeaux 5049, 
5050 (BRIT); 5.3 km W of Zitacuaro via Hwy 15, 2,000 m, 24 Jul 1991, Comeaux 5057 (BRIT);5.6 km W 
of Zitacuaro via | IWY 15,1,950 m,24 Jul 1991 ,Comeaux 50527, 5053, 5054 (BRIT); 7.7 km W of Zitacuaro 
via Hwy 15, 1950 m, 24 Jul 1991, Comeaux 5055 (BRIT); 25.8 km W of Zitacuaro via Hwy 15,1,850 m 
24 Jul 1991, Comeaux 5056 and 5057 (BRIT); 42 km W of Zitacuaro via Hwy 15, 2,000 m, 24 Jul 1991, 
Comeaux 5058 (BRIT); 10.7 km W of Hidalgo via Hwy 15, 2,150 m, 25 Jul 1991, Comeaux 5059, 5060 
(BRIT); 10.8 km W of Hidalgo via Hwy 15, 2,150 m, 25 Jul 1991, Comeaux 5067 (BRIT); 11.0 km W of 
Hidalgo via Hwy 15, 2,150 m, 25 Jul 1991, Comeaux 5062 (BRIT); 25.9 km W of Hidalgo via Hwy 15 
2,300 m, 25 Jul 1991, Comeaux 5063 (BRIT); 60.2 km W of Hidalgo via Hwy 15, 2,450 m, 25 Jul 1991, 
Comeaux 5064 and 5065 (BRIT); 61.1 km W of Hidalgo via Hwy 15, 2,450 m, 25 Jul 1992, Comeaux 
5066 and 5067 (BRIT); 20.6 km E jct. Hwy 15 and 126/43 near Morelia, 2,400 m, 25 Jul 1991, Comeaux 
5068 (BRIT); 31.7 km E jct. Hwy 15 and 126/43 near Morelia, 2,300 m, 25 Jul 1991, Comeaux 5069 
(BRIT); 31.0 km E jct. Hwy 15 and 126/43, 2,300 m, 25 Jul 1991, Comeaux 5070 (BRIT); 13.1 km E jet. 
Hwy 15 and 126/43, 2,000 m, 25 Jul 1991, Comeaux 5077 (BRIT); 7.2 km E jet. Hwy 15 and 120, W of 
Morelia, 2,200 m, 26 Jul 1991, Comeaux 5072 (BRIT); 32.8 km E of Zamora via Hwy 15, 2,000 m, 26 Jul 
1991, Comeaux 5073 (BRIT); Ladera Norte del Cerro El Cacique, zona 9, Mpio. Zitacuaro, 2,410 m, 11 
Oct 1980, Contreras 1214 (MEXU); Rincon, 2 Apr 1932, Hinton 482 (MEXU); 2 km S de Villa Escalante, 
27 May 1979, Soto Nunez 1428 (MEXU); 16 km al E de Uruapan, 29 May 1979, Soto Nunez 1445 (MEX); 

12 km E de Coalcoman, 28 Mar 1980, 1,675 m, Soto Nunez 2157 (MEXU);En Turundeo, 6 km al NW de 
Tuxpan, carr. México-Morelia, 1,845 m, 3 Apr 1981, Soto Nunez 2930 (MEXU); San Jose Purua, Mpio.de 
Jungapeo, 10 Mar 1984, Soto Nunez 6231 (MEXU); Mpio.de Villa Madera, 2,240 m, 29 Apr 1979, Zavala 
206 (MEXU). Morelos: 1.1 km N jct. Hwy 95 and 115 near Tepoztlan, 2,200 m, 22 Jul 1991, Comeaux 
5033 and 5034 (BRIT); Base of mountain with temple at Tepoztlan, 1,500 m, 23 Jul 1991, Comeaux 
5035, 5036, 5037, 5038, 5039, 5044,and 5045 (BRIT); 12.8 km S of Tepoztlan via Hwy 115, 1,450 m, 23 Jul 

991, Comeaux 5040 (BRIT); NE of Tepoztlan ca.3 km on dirt road, 23 Jul 1991, Comeaux 5041 (BRIT): 
1.3 km E of square at Tepoztlan, 1,450 m, 23 Jul 1991, Comeaux 5042 and 5043 (BRIT); Cuernavaca, 
Nov 1941, Mirand 1906 (MEXU); Mountainside above Cuernavaca, 3 Feb 1899, Pringle s.n. (MEXU); 
Sierra de Tepoxtlan, 2,287 m, 15 Mar?, Pringle s.n. (MEXU); Tepetixtla km 71 carr. México-Cauutla, 
Mpio. de Cauutla, 19 Mar 1978, Sanchez 2142 (MEXU); Pedregal de Santa Maria, 1 Apr 1970, Vazquez 
2415 aera Santa Clara, N.O.de Acatlipa, 2 7 fe ees 3100 (MEXU).Oaxaca: 19.5 
km N of jct. Hwy 190 and 175 at Oaxaca, 1,700 m, 16 Jul 1992, Comeaux 5180 (BRIT); 10 km NE 
Oaxaca, camino a Yxtlan,4 Feb 1966, Delgadillo 213 eu Dto.de Ixtlan, Sierra de Juarez, Ruta 175 
a1 kmalN de Ixtlan de Juarez, 1,800 m, 18 Apr 1962, Lorence 4067 (MEXU); 10 km S de Cuquila, carr, 
Tlaxiaco- ioe Dto. de Tlaxiaco, 1,800 m, 8 Jun 1985, Mendoza 1479 (MEXU); 4 km al W del Vado 
camino n Sebastian de las Gratas, Dto. Sola de Vega, 6 Apr 1984, Torres 4927 (MEXU). Piiebla: 
Zacatlan, an 1943, Bruff 1511 (MEXU); 1.4 km N of Puente Cantarranas Il via Hwy 190 S of Atlixco, 
1,850 m, 15 Jul 1992, Comeaux 5171 (BRIT); At Puente Cantarranas Il S of Atlixco, 1,850 m, 15 Jul 1992, 
Comeaux 5172,5173,5174,5175,5176,5177,51 be ee (BRIT). San Luis Potosi: In Rio Verde, 50 mS of 
Hwy 70 along dirt rd.0.2 km E of main blvd., 1 m, 28 Jun 1986, Comeaux 4205 (BRIT);in Rio Verde, 
2.9 km E jct. Hwy 69 and 70, 1,050 m, 19 Jul ee Comeaux 5207 (BRIT). 


— 


COMEAUX AND LU 129 


Vitis cinerea (Engelm sray)Enc Millardet var.t t (Planchon) Comeaux, 
b. nov Bava be danalenty var.tomentosa Planchon, Compt. Rend. Hebd. Séances Acad. 
Sci.91:425. 1880. Type: MEXICO. Nuevo Leon: Cerro de la Silla, Jun, 1834, Berlandier 3116 (LectotyPe: 

WISI isocectotyPe: MO!). 


Vines to 15 m, stems on current season growth tomentose to pubescent,rarely with 
bristle-like trichomes, branchlets nearly terete to faintly angular; internodes 3-10 cm 
long; nodes faintly banded to without red Pigmentation, pith interrupted at the nodes 
by a diaphragm 2-3 mm thick;lenticels al g tips tomentose, with white, arach- 
noid trichomes, not enveloped by young leaves. Leaves cordiform to long-cordiform, 
ger erally without lobes or less frequently 3-lobed, with lateral lobes acute, apex acute to 
acuminate, base cordate to nearly truncate with the basal sinus varying from mostly U- 
shaped, to lyre-shaped and also V-shaped, lateral sinues acute; margins mostly entire, 
occasionally with teeth to 3 mm long, with or without cilia; midrib with 4—7, usually 5 
pairs of prominent veins; lamina glabrous to puberulant on adaxial surfaces, tomentose 
to pubescent on abaxial surfaces, with the pubescence consisting of mostly arachnoid 
trichomes, sometimes including simple, straight, pointed trichomes, 5.5-11.2 cm wide, 
6.2-12.8 cm long; petioles tomentose to puberulent, 1.4-4.1 cm long; stipules brown, 
0.5—-1 mm wide, 1-3 mm long, caducous. Tendrils and inflorescences absent every third 
node, tendrils bifurcate or trifurcate, to 10 cm long. Inflorescences 4.0-11.0 cm long, pe- 
duncles 1.0-4.2 cm long, shoulder 1.0-5.0 cm long, sometimes absent or replaced by a 
tendril. Flowers of the functionally pistillate plants with pistils 1 mm long. Fruit a berry, 
black, glaucous, 5.5-10 mm in diameter, with a pleasant flavor when fully ripe. Seeds 
brown, pyriform, 3.5-5.5 mm long. 

Commonly found along streams and other moist sites in the semiarid scrub and 
grasslands (De Miranda 1989) east of the Sierra Madre Oriental along the Rio Grande 
from the Del Rio, TX, area south to the Tropic of Cancer (120-1050 m. elev.) Individuals 
from the Del Rio area have characters intermediate between V. cinerea var. helleri, prima- 
rily from the Edwards Plateau region of Texas, and the more southerly distributed var. 
tomentosa. These intermediates establish the existence of clinal variation between the 
above varieties of V. cinerea. 


ae description based on the following specimens: MEXICO. Coahuila: S edge of Morelos via 
wy 57,16 Jun 1986, Comeaux 4032 and 4033 (BRIT); 10.1 km N of Morelos via Hwy 57, 16 Jun 1986, 
eae 4034 (BRIT); 36.8 km N of Morelos via Hwy 57,16 Jun 1986, Comeaux 4035, 4036, 4037 and 
4038 (BRIT); S of Acuna at 3.0 km S jct. Hwy 2 and 29, 17 Jun 1986, Comeaux 4039 (BRIT); S of Acuna 
at 3.5 km S jct. Hwy 2 and 29,17 Jun 1986, Comeaux 4040 and 4041 (BRIT);S of Acuna at 23.0 km S jct 
Hwy 2 and 29,17 Jun 1986, Comeaux 4042, 4043 and 4044 (BRIT); 27.2 km S jct. Hwy 2 and 29 in S. 
Juan de Sabina, 17 Jun 1986, Comeaux 4045 (BRIT); 2.7 km N of Muzquiz via Hwy 53, 17 Jun 1986, 
Comeaux 4046, 4047 and 4051 (BRIT);10.7 km N of Muzquiz via Hwy 53 at river, 17 Jun 1986, Comeaux 
4048, 4049 and 4050 (BRIT). Nuevo Leon: 6.6 km E of Sabinas Hildago via Hwy 34, 16 Jun 1986, 
Comeaux 4026, 4027 and 4028 (BRIT); 1.8 km W of Sabinas Hidalgo via Hwy 34, 16 Jun 1986, Comeaux 
4029 (BRIT); 21.1 km W of Sabinas Hidalgo via Hwy 29, 16 Jun 1986, Comeaux 4030 (BRIT); 22.1 km W 
of Sabinas Hidalgo via Hwy 29, 16 Jun 1986, Comeaux 4031 (BRIT); Monterey, 25.0 km N jct. Hwy 85 
and rd.to Santiago, 18 Jun 1986, silos ne one (BRIT); S of Monterey at 2.2 km N jct. Hw 
85 and rd.to Santiago, 18 Jun 1986 BRIT); at river 19.2 km N of Montemorelos 


n 


130 SIDA 19(1) 


via Hwy 85, 18 Jun 1986, Comeaux 4056 (BRIT); 1.1 km N of Montemorelos via Hwy 85, 18 Jun 1986, 
Comeaux 4057, 4058 and 4059 (BRIT); 3.2 km S of Montemorelos via Hwy 85, 18 Jun 1986, Comeaux 
4060 (BRIT); 37.3 km S of Montemorelos via Hwy 85, 18 Jun 1986, Comeaux 4061 (BRIT); 3.5 km N of 
Linares via Hwy 85, 18 Jun 1986, Comeaux 4062 (BRIT); 27.5 km N of Montemorelos via Hwy 35, 29 
Sep 1987, Comeaux 4946 (BRIT); 20.5 km N of Montemorelos via Hwy 35, 29 Sep 1987, Comeaux 
4947 (BRIT); 19.4 km N of Montemorelos via Hwy 35,29 Sep 1987, Comeaux 4948 (BRIT); 24.2 km N of 
Linares via Hwy 85, 29 Sep 1987, Comeaux 4949 (BRIT); 5.0 km W of Iturbide via Hwy 58,5 Oct 1990, 
Comeaux 4975 (BRIT); 2.4 km E of Iturbide via Hwy 58,5 Oct 1990, Comeaux 4976 (BRIT); 5.4 km E of 
lturbide via Hwy 58,5 Oct 1990, Comeaux 4977 (BRIT); 9.4 km E of Iturbide via Hwy 58,5 Oct 1990, 
Comeaux 4978 (BRIT); 12.6 km E of Iturbide via Hwy 58,5 Oct. 1990, Comeaux 4979 (BRIT); 7.8 km W 
of jct.rd. to Horsetail Falls and Hwy 85,950 m, 22 May 1991, Comeaux 5007 (BRIT);8.8 km W of jct.rd. 
to Horsetail Falls and Hwy 85, 1050 m, 22 May 1991,Comeaux 5002 and 5003 (BRIT);5.6 km E of jct. of 
rd. to Casacade de Chipitin and rd. to Horsetail Falls from Hwy 85, 850 m, 22 May 1991, Comeaux 
5004 (BRIT);9.4 km N of bridge at General Teran via Hwy 35,275 m, 11 Jul 1992, Comeaux 5154 (BRIT); 
1.0 km S of bridge at General Teran via Hwy 85,275 m,11 Jul 1992, Comeaux 5155 (BRIT); 13.0 km N 
of bridge at Linnares via Hwy 85, 275 m, 11 Jul 1992, Comeaux 5156 (BRIT); 4.8 km NW of General 
eee heals San Pedro, 21 Apr. 1962, Ruiz 63 (TAMU). Leelee 31.4kmN of Rio mae near 
Hw 19 Jun 1986, Comeaux 4063 and 4064 (BRIT); 18.1 km N of Rio Corona via Hwy 85, 1 

1986, pone 4065 (BRIT);At Rio Corona jct. Hwy 85, tun 1986, Comeaux 4066 (BRIT); 21.9 oy ‘ 
of Tropic of Cancer via Hwy 85, 19 Jun 1986, Comeaux 4067 (BRIT); 5.6 km N of Rio Purificacion via 
Hwy 85, 29 Sep 1986, Comeaux 4950 (BRIT); 0.3 km S of Rio Purificacion via Hwy 85, 29 Jun 1990 
Comeaux 4951 (BRIT); 10.9 km N of Tropic of Cancer via Hwy 180,120 m,5 May 1991, Comeaux 4980 
and — S : 

EXAS. Maverick Co.: 1.6 km S of Quemado via Hwy 277,275 m,21 May 1986, Comeaux 

3861 a a (BRIT). Val Verde Co.: Del Rio, near Val Verde Winery, 300 m, 21 May 1986, Comeaux 
3864, 3865, 3866, 3867, 3868, 3869 and 3870 (BRIT); San Felipe Country Club, Del Rio,8 May 1943, Cory 
41717 (UT 


ACKNOWLEDGMENTS 
The authors are appreciative of the contributions made by P.R. Fantz in the preparation 
of this manuscript. 
REFERENCES 


Bailey, L.H.1895-97. Vitaceae.|n: Robinson, B.L.,Ed.Gray’s synoptical flora of North America. 
American Book Co., New York. Pp.429-432 
Baiey, LH. 1934. The species of grapes peculiar to North America. Gentes Herb. 3:154- 


Gatet, P1988. Cepages et vignobles de France. Tome I. Les Vignes Americaines. 2nd ed. 
Imprimerie Charles Dehan, Montpellier 

De Miranda, E.G.1989.Nuevo atlas porrua de la Republica Mexicana. 8th ed. Editorial Porrua, 
S.A., Mexico, D.F. 

Moore, M.O.1992.Classification and systematics of eastern North American Vitis L.(Vitaceae) 
North of Mexico. Sida 14:339-367 

Munson, T.V. 1890a. A classification of American grapes. Gard. Forests. 3:474-475. 

Munson, I.V,1890b. Classification and generic synopsis of the wild grapes of North America. 
USDA Pom. Bull. 3:1-14 


COMEAUX AND LU TOMENTOSA 131 


Munson, TV. 1896. Series VIIl- Cinerascentes. Rev. Vitic.6:421-425. 

Munson, T.V. 1909. Foundations of American viticulture. T.V. Munson and Son., Denison,TX. 

PLANCHON, J.E. 1887. Monographie des Ampelidees Vrais. In: DC Monographie 
Phanaerogamarum. 5:305-368. 

Renero, R.E. 1983. The Thomas Volney Munson Memorial Vineyard. Grayson County Col- 
lege, Denison, TX. 

STANDLEY, P.-C. 1920-26. Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23. 


132 SIDA 19(1) 


BOOK REVIEW 


Brent Berwin. 1992. Ethnobiological Classification: Principles of Categorizationof Plants 
nd Animals in Traditional Societies. (ISBN 0-691-09469-1, hbk.). Princeton Uni- 
versity Press, 41 William Street, Princeton, New Jersey 08540, U.S.A.$29.00. Illustrated 

with b&w line drawings. 


As more and more anthropology and biology departments begin to collaborate and ethnobiology 
becomes a bona fide program, it’s hard to imagine Ethnobiological Classification not being an es- 
sential text. It is because of this book, and the debates it sparked, that folk biological taxonomies 
have become such a hot topic for a better part of the nineties and worthy of so much further 
investigation in the coming millennium. It is fitting that one of the nation’s leading ethnobiology 
programs, the University of Georgia, Athens, is run by the author and his wife, Elois 

Divided into two major parts, “Plan” and “Process,” Berlin discusses the fatindauons of 
ethnobiological classification inferred from an analysis of descriptions of individual systems and 
the underlying processes involved in the eh ee and evolution of ethnobiological systems in 
general. Essentially this book is a revision and summary of the many articles Berlin co-authored 
with Dennis Breedlove and Peter Raven in the late ie and the articles Us cos to publish 
on his own throughout the 70s, 80s and 90s. These articles lead man 
to chime in on the issue. Eugene Hunn, Cecil Brown, Terence Hays, to name a a either agreed, 
disagreed, or agreed to disagree with Berlin. This debate contributed a great deal to the overall 
knowledge of ethnobiological classification and theory. |t is only when such topics become thor- 
oughly researched and continually challenged that the essence of the subject is revealed and 
worthy of inclusion in a college program 

Part one is divided into four chapters, discussing the “Making of a Comparative Ethnobiology,” 
“The Primacy of Generic Taxa in Ethnobiological ce ered as Nature of HE CHIE Taxa,” and 
“Natural and Not So Natural Higher Order Categories.” Berlin f the folk classi- 
fication debate, intellectualists and utilitarian approaches, 4 eo ane data presentation as 
flection of a changing theory, andar salience, general nomenclatural properties of i 
taxa, and folk taxonomic ranking to name a few. Part two comes in 3 chapters:"Patterned Variation 
in bee aie ce i Nonarbitrariness of Ethnobiological Nomenclature,” and “The 
Substance and Evolution of Ethnobiological Categories.” This is where the book becomes more 
than just a book on ethnobiological classification, but a book that asks further questions and chal- 
lenges those anthropologists who see reality as a “set of culturally constructed, often unique and 
idiosyncratic images, little constrained by the parameters of an outside world.” Berlin looks for varia- 
tion in the patterns of ethnobiological knowledge and the biological ranges of Folk Taxa, discusses 
experiments on sound symbolism, and the typological speculations on the evolution of 
ethnobiological categories. The book ends with a proper bibliography, author index, indices of 
scientific and ethnobiological names, and a general subject index. 

student's perspective the best thing | can say about the book is that it pushed me into 

the library and taught me an essential lesson in research, that often the most important 
article is its bibliography. Not only did | get the necessary background on ethnobiological classifi- 
cation theories, | also looked further and began to understand the questions Berlin was tackling 
and started asking my own. But, his thorough citations of the articles that challenged and/or cor- 
roborated with his ideas made research both enjoyable, but perhaps more importantly, compre- 
hensive.| wasn't left with one guy’s theory on ethnobiological classification, | got the full debate — 
Kevin D. Janni. 


SIDA 19(1): 132. 2000 


BOLBITIS THOMMANKUTHIANA (LOMARIOPSIDACEAE), 
A NEW SPECIES OF FERN FROM INDIA 


Santhosh Nampy 


ee of Botany 
eph’s College 
ees ae 673 008, INDIA 


ABSTRACT 
Bolbitis thommankuthiana, a new species of fern from Kerala, India, is described. 


Key woros: Lomariopsidaceae, Bolbitis thommankuthiana, India, Ferns, Pteridophyta. 


RESUMEN 


Se describe Bolbitis thommankuthiana, una nueva especie de helecho de Kerala, India. 
INTRODUCTION 


In his monograph of the genus Bolbitis Schott, Hennipman (1977) recognized 44 species 
and 13 hybrids. Since that time, 17 new taxa have been described, including two from 
Kerala (Manickam & lrudayaraj 1992; Nayar & Geevarghese 1993). During recent floristic 
explorations in the Western Ghats of Kerala,| encountered an interesting population of 
plants growing on exposed wet boulders and rocks in streams at an elevation of 400 m. 
Since this material does not agree with descriptions of any of the known species of 
Bolbitis, it is described here as new. 


saa thommankuthiana Nampy, sp. nov. (Figs. 1, 2). Tyee: INDIA. Keraca. Idukki Dt: 
ommankuthu, alt. 400 m, 26 Dec 1996, Nampy S 426 (HoLotyPe: CALI; Isotype: K). 


RATHI tic +h | bs ; q ffinis sed ab tat hi i lathratis 


distincte bicoloribus, rhachidi late alata, alis ad rhi tendentib 'sporis fuscioribus exina dense 
plicata, bulbillis foliaribus nullis facile distinguenda. 


Plants 10-13 cm tall. Rhizome 1.5-2 mm thick, short-creeping, sclerenchyma strands 2— 
4,without circumvascular sheath. Rhizome scales ovate-lanceolate, pseudo-peltate, gradu- 
ally tapering to a glandular apex, bicolorous, central cells blackish, margin pale, glandular. 
Fronds clustered, dimorphic, pinnate, scaly along the stipe and occasionally the costa. 
Sterile fronds 6-13 cm long, 2.2-3 cm broad, pinnae 4-6 pairs; terminal pinna conform 
with the lateral pinnae. Stipe 2—3.5 cm with wings ca. 1mm wide. Pinnae sub-opposite to 
alternate, 1-2.3 x 0.5-0.7 cm, 2.5 cm apart, acroscopic side broadly cuneate, basiscopic 
side narrowly cuneate, herbaceous, apex blunt to acute, margin shallowly lobed with 
short setae; lacking foliar bulbils; veins anastomosing to form a row of costal areoles, 
extra costal free veins present. Fertile fronds to 10.5 cm long, pinnae acrostichoid, 4-7 
mm long.Sporangium 261 x 226 um.Annulus 13 or 14 celled. Spores monolete, bilateral 
41 x 31 um, dark with densely folded exine. 


SIDA 19(1): 133-135. 2000 


" wet. 
ve 


yh 12 
Ck} 
RRS 
A 
Ways 
Cy 


¢ 
ee 


SIDA 19(1) 


Lh 


py A. Habit; B. Venation; C Sporangium; D. Rhizome Scale 


NAMPY, A NEW SPECIES OF BOLBITIS FROM INDIA 135 


Fic. 2. SEM nhot h nfth £ Balhitic +h I he Nl A Snore y 2000°R Netail of . cnnn 


t oer r ry t / t t 


logy.—The species is named after the type locality Thommankuthu, a water 
fall at an elevation of 400 m in wet evergreen forests of Kerala. 

Distribution and habitat.—Known only from three localities in the Western Ghats of 
Kerala, usually found growing on boulders and rocks in streams. 


Additional specim ined: INDIA. KERALA. Idukki Dt.:T| hu, 1 Jan 1995, nampeS 
5 (CALI). Kozhikode Dt.: Vellarimala, 11 Aug 1995, Nampy S 74 (CALI). Pathanamthitta Dt. 
Kulathupuzha, 29 Nov 1974, Shyamala CU 9366 (CALI). 


Notes.—Bolbitis thommankuthiana is closely allied to B. semicordata but can readily be 
distinguished by its smaller size, broadly winged rachis, the wings extending to the rhi- 
zome and complete absence of foliar bulbils. In B. semicordata the plants are 15-70 cm 
tall, wings usually below 0.5 mm, which are not extending to the rhizome and with foliar 
bulbils. | have examined the type of B. semicordata (K) as well as other material of this 
species and found that these two species are not connected by any intermediates. 
ACKNOWLEDGMENTS 

|am thankful to the Department of Science and Technology, New Delhi for the BOYSCAST 
Fellowship; the Curators of K and L for allowing me to examine the specimens; B. van 
Heuvan for her assistance in SEM photography; P. Bas, H.R Nooteboom (Rijksherbarium) 
and Principal, St. Joseph's College, Kozhikode for providing facilities and A.Radcliffe-Smith, 
(Kew) for Latin diagnosis and A.K. Pradeep, University of Calicut for various help. 


REFERENCES 
HeNNIPMAN, E. 1977. A monograph of the genus Bolbitis (Lomariopsidaceae). Leiden Uni- 
versity Press. 
Manickam, V.S.and V.IRuDAvARAJ, 1992. Pteridophytic flora of the Western Ghats—South India. 
Bishen Singh & Mahendrapal Singh, New Delhi. 
Nayar, B. K. and K.K. GeevarGHese, 1993. Fern flora of Malabar. Indus Publishing Company, 
New Delhi. 


136 SIDA 19(1) 


BOOK REVIEW 


H. Resit Akcakaya, Mark A. Burman, and Lev R. Ginzeurc. 1999 Applied Population Ecology. 
(ISBN 0-87893-028-0, pbk.and CD-ROM), Sinauer Associates, Inc.23 Plumtree Road, 
Sunderland, MA 01375, U.S.A. (publish@sinauer.com). $41.95. 280 pages/64 illustra- 
tions, text with CD-ROM) 


This book is designed to be used as a text for undergraduates studying biology, wildlife or natural 
resource management, or environmental science. It functions with the RAMAS® EcoLab 2.0 soft- 
ware, which is provided with the book on a CD-ROM. Its “aim is to teach quantitative methods that 
are necessary to develop a basic understanding and intuition about ecological processes, without 
intimidating or discouraging students who do not have extensive mathematical backgrounds.” 
The aim has certainly been achieved. 

The book is divided into 8 chapters, including: 1) Population Growth, 2) Variation, 3) Popula- 
tion Regulation, 4) Age Structure, 5) Stage Structure, 6) Metapopulations and Spatial Structure, 7) 
Population Viability Analysis, and 8) Decision-Making and Natural Resource Management. Each 
chapter includes the introduction of several major concepts, discussions of each, followed by exer- 


cises and further reading. The information is very clearly | | and richly illustrated with tables 
and ued The exercises, — the RAMAS Ecolab, provide students with the opportunity for data 
manipulation and analysis, and will be useful when they need to set up their own models based on 


data they have ares 

This textbook, concomitant with its CD-ROM, provides a powerful package that will make any 
population biology course relevant to everyday situations a practicing biologist must face. It pro- 
vides not only tools to facilitate mastery of the course material, but that will also serve as a frame- 
work upon which students can formulate their own hypoethical models and test them using ac- 
tual population data.| was sel asi We ed with the Sialeians on Mele epelet ns which are 
precisely the most common phenomenon botanists face 
Conservation and Development sites. ne the text does not sAdicss ‘the fovel molecular ap- 
proaches to conservation of infraspecific genetic diversity, the text’s success thus far ensures that it 
will have another edition, where we fully expect to find that additional element. Other than that 
one oversight, | find the book to be complete, clearly presented and certainly a bargain given the 
long-term utility of the software provided on the CD ROM. | unhesitatingly recommend it to all 
who must teach a population biology course.—John J. Pipoly Il, Botanical Research Institute of Texas. 


SIDA19(1 ): 136. 2000 


A NEW COMBINATION FOR SARCOSTEMMA 
CYNANCHOIDES VAR. HART WEGII (ASCLEPIADACEAE) 


Alexander Krings' 


Zilker Botanical Garden 

2220 Barton Springs Rd. 

Austin, 1X 78746, U.S.A. 
In order to validate names for a checklist of the vines of the Chihuahuan desert, a new 
combination for a taxon traditionally placed in Sarcostemma R. Br. (Correll & Johnston 
1979; Kartesz 1994) is proposed. Recent cladistic analyses by Liede (1996) failed to sup- 
port the monophyly of Sarcostemma sensu Holm (1950), instead showing support for 
the monophyly of genera as circumscribed by Schlechter (1914). Based on her analyses, 
Liede (1996) suggested, among other circumscriptions, that Sarcostemma be restricted 
to the non-campanulate Old World taxa and that Funastrum E. Fourn. should apply to 
the non-campanulate New World taxa. The taxon in question, Sarcostemma cynanchoides 
Decne. var. hartwegii (Vail) Shinners, was originally described as Sarcostemma lineare 
Decne. in 1840. However, the existence of an earlier homonym, Sarcostemma lineare 
Spreng., 1822, (based on a Venezuelan collection), required the use of another name for 
the Mexican taxon. Although once employed through widespread misapplication, Holm 
(1950) notes that the name Sarcostemma heterophyllum Torr. is unavailable for the sub- 
specific taxon of Sarcostemma cynanchoides as the type (Wright 1679, New Mexico, USA) 
is Sarcostemma crispum Benth. Holm (1950) decided to employ the epithet “hartwegii" 
rather than create a new name. Thus, the name Philibertella hartwegii Vail—validly pub- 
lished in 1897, based on the same collection as the 1840 Decaisne name (Hartweg 217, 
Leon, Mexico)—is the appropriate basionym of the most recently used names, 
Sarcostemma cynanchoides var. hartwegii (Vail) Shinners and Sarcostemma cynanchoides 
ssp. hartwegii (Vail) R.W. Holm, as well as the proposed, new combination: 


Funastrum cynanchoides (Decne.,) Schltr. var. hartwegii (Vail) Krings,comb.nov.Philibertella 
artwegii Vail, Bull. Torrey Bot. Club 24:308. 1897. Tyee: MEXICO. Leon: Hartweg 217, 1839 (NY). 
Funastrum hartwegii (Vail) Schltr, Repert. Soec. Nov. Regni Veg. 13:285. 1914. Sarcostemma 
cynanchoides Decne. var. hartwegii (Vail) Shinners, Sida 1:361.1964. Sarcostemma cynanchoides 
Decne.ssp. hartwegii (Vail) R.W.Holm, Ann. Missouri Bot. Gard. 37:530. 1950. Sarcostemma 
Decne. in Benth., Pl. Hartw. 25. 1840 (non Spreng. 1822). 


— 


ineare 


Distribution.—Funastrum cynanchoides var. hartwegii occurs between 30-1500 m in dry, 
sandy, or gravelly soil in Trans-Pecos Texas, New Mexico, Arizona, and Utah, as well as the 
Mexican states of Baja California Norte, Baja Califonia Sur, Chinuahua, Coahuila, Durango, 
Guanajuato, Jalisco, Querétaro, Sinaloa, Sonora, and Zacatecas. 


"Current address: Herbarium, Department of Botany, North Carolina State University, Campus Box 7612, Raleigh, 
NC 27695-7612, U.S.A. 


SIDA 19(1): 137-138. 2000 


138 SIDA 19(1) 


ACKNOWLEDGMENTS 


| thank M. Fishbein and an anonymous reviewer for their thoughtful comments on the 
manuscript. 


REFERENCES 


Corrett, D.S and M.C. JoHNsTon. 1979. Manual of the vascular plants of Texas. University of 
Texas at Dallas Press, Richardson. 

Kartesz,J.1994.A synonymized checklist of the vascular flora of the United States, Canada, 
and Greenland. Timber Press, Portland. 

Hoim, R.W. 1950. The American species of Sarcostemma R. Br. (Asclepiadaceae). Ann. 
Missouri Bot. Gard. 37:477-560. 

Liebe, S. 1996. Sarcostemma (Asclepiadaceae)-a controversial generic circumscription 
reconsidered: Morphological evidence. Syst. Bot. 21:31-44. 

ScHLECHTER, R. 1914. Philibertia H.B. et Kth. und Funastrum Fourn. Repert. Spec. Nov. Regni 
Veg. 13:279-287. 


RHYNCHOSPORA LEPTOCARPA (CYPERACEAE), 
AN OVERLOOKED SPECIES OF THE 
SOUTHEASTERN UNITED STATES 


Bruce A. Sorrie 


3076 Niagra-Carthage Road 
Whispering Pines, NC 28327, U.S.A. 


ABSTRACT 


Based on morphological analysis, Rhynch | ,long synonymized under R. capitellata, 
is reinstated as a distinct species. Although ion related 1 to ane Bttey R. lepiocaipa aitete lh ina 
number of mensural and qualitative characters. Furtherm 

ern coastal plain and it inhabits a narrow range of a t communities, very unlike those of R. 
capitellata. Based on the relati f finding R.leptocarpa in suitable habitat, it is believed not to 
be rare, but merely overlooked. 


RESUMEN 


En base a analisis morfolégicos Rhynchospora leptocarpa, que fue sinonimizada a R. capitellata, se 
vuelve a considerar como una especie diferente. Aunque claramente relacionada con esta Ultima, 
R. leptocarpa difiere en cierto numero de caracteres cuantitativos y cualitativos. Ademas, su 
distribucion esta confinada al sur de la hae costera y vive en un numero Pissniuiglae de 

comunidades vegetales, muy diferentes de las de R. capitellata ace alo facil q ar. 
leptocarpa en ic habitats adecuados, se cree que no es rara,sino que aero ha side fanroradsy 


INTRODUCTION 


Rhynchospora leptocarpa (Chapm.ex Britt.) Small was first recognized as a distinct entity 
by AW.Chapman, longtime resident of the Florida panhandle and author of Flora of the 
Southern United States (Chapman 1860, 1883, 1897). Curiously, Chapman did not pub- 
lish the name, although he had prepared a manuscript (fide Britton 1892). In the three 
editions of his Flora, Chapman does not mention this entity.In 1892 Britton published it 
as R.glomerata (L.) Vahl var. leptocarpa Chapm.and ascribed to it a range of South Caro- 
lina, Florida, and Alabama. Later, Blake (1918) unraveled the R. glomerata tangle by split- 
ting off the generally more northern R. capitellata (Michx.) Vahl from the more southern 
R. glomerata. Blake treated the entity as R. capitellata var. leptocarpa (Chapm. ex Britt.) 
Blake and provided a brief Latin diagnosis. Blake gave its range as Virginia to Florida and 
Mississippi.In the second edition of his Flora, Small (1933) raised it to full species status 
(without explanataion or discussion) and provided the only complete English descrip- 
tion available. Small gave its range as South Carolina to Florida and Mississippi. In her 
monograph of the section Eurhynchospora, Gale (1944) reduced it not to varietal status, 
but to complete synonymy under R. capitellata. Although she treated two other botani- 
cal forms of &. capitellata, Gale omitted any discussion of taxon leptocarpa, possibly due 
to the paucity of known specimens at that time. 


SIDA 19(1): 139-147. 2000 


140 SIDA 19(1) 


Subsequent authors, most notably Clewell (1985), Godfrey and Wooten (1979), Jones 
and Coile (1988), Kartesz (1994), Radford et al. (1968), and Wunderlin (1998), do not rec- 
ognize R. leptocarpa at any level.None of the state Natural Heritage Programs within the 
range of R. leptocarpa recognizes it or lists it as rare, except for North Carolina. The only 
references to Rhynchospora leptocarpa in the past half century are those of Anderson 
(1995) and Sorrie et al. (1997). Anderson collected specimens in the Florida panhandle 
and treated leptocarpa at the species level, but suggested that the R. capitellata complex 
needed further study. Robert Kral (pers. comm.) also has suggested that R. leptocarpa 


may warrant renewed status. 

My attention was first brought to the problem by Julia Larke, who collected plants 
in 1992 on Fort Bragg Military Reservation, North Carolina, as part of a rare species sur- 
vey. Although these plants keyed cleanly in Godfrey and Wooten (1979) and Gale (1944), 
her plants did not closely resemble plants | knew from New England as R. capitellata. 
Field and herbarium studies over the past seven years have convinced me that the two 
are distinct species. 


MATERIALS AND METHODS 


Specimens of Rhynchospora leptocarpa were collected in the field during the course of 
botanical survey work in Alabama, Florida, Mississippi, and North Carolina. These have 
been supplemented by speci examined at the following herbaria: DUKE, FLAS, FSU, 
GA, GH, IBE, NCSC, NCU, SWSL, US, USA, VDB. From across the range of each species, five 
achenes from each of 10 plants were measured. Measurements were made of the fol- 
lowing achene characters: length of achene body, tubercle length, and length of peri- 
anth bristles relative to tubercle (e.g., the length that the bristles exceed the tip of the 
tubercle).M vere also made of the width of spikelet clusters (n = 50).Counts 
were made of the number of spikelet clusters per culm and the number of glomerules 
per inflorescence node (n = 50). 

Notes on habitat were made at over 40 sites in ie ie supplemented by her- 
barium label data. Range maps were prepared from rature, atlases, herbarium 
specimens, and field work. 


Herbarium acronyms follow Holmgren et al.(1990), except“bas” (personal herbarium 
of Bruce A. Sorrie). 


RESULTS AND DISCUSSION 


Distinguishing characters 
Mensural data are presented in Table 1.These data suggest that Rhynchospora leptocarpa 
differs from R. capitellata in at least four important ways. 

1) Number of spikelet clusters per culm.—Rhynchospora leptocarpa possesses 5-8 Clus- 
ters per culm, occasionally 4; R. capitellata possesses 3-5, rarely 6 

2) Glomerule width.—Each spikelet cluster is composed of 1-3 head-like glomer- 
ules, which are much broader in R. capitellata. In addition, glomerules of R. leptocarpa 


SORRIE TOCARPA, AN OVERLOOKED SPECIES 141 


Taste 1. Comparison of selected characters of R hospora leptocarpa and R. capitellata, based on 
50 counts or measurements. Measurements in irilimersie 


Character R. leptocarpa R. capitellata 

number of spikelet 4-8 (K=6.2) 3-5 (K=4.1) 

clusters per culm 

glomerule width 4-8 (X=6.8) 6-13 (X=9.1) 

achene body length 1.6-1.8 (X=1.7) 1.4-1.8 (X=1.6) 

tubercle length 0.8-1.1 (X=1.0) 0.9-1.4 (-1.6) (X=1.1) 

ratio of achene body/ 125 1.45 

tubercle length 

longest bristle relative bristle exceeds tip bristle varies from shorter 

to tip of tubercle by 0.3-1.0 mm than tip (0.4 mm) to 
(X=0.8 mm longer) longer (0.3 mm) 


*=0.05 mm shorter) 


assume a turbinate shape, rather than the broadly ovoid to hemispherical shape of R. 
capitellata. 

3) Achene body and tubercle length._—Although the combined length of body and 
tubercle is similar in the two species, the proportions differ. Body length of R. leptocarpa 
averages longer than capitellata, but tubercle length averages shorter. 

4) Bristle length.—A striking feature of R.leptocarpa is that the bristles always exceed 
the tubercle (in rare instances, one of the six bristles may be equal or slightly shorter). 
Moreover, the longest bristle usually exceeds the tubercle by at least 0.5 mm. In R. 
capitellata, bristles normally are slightly shorter than the tubercle, but the longest bristle 
may exceed it by 0.2 or even 0.3 mm. 

There are qualitative characters which also serve to distinguish the two species. 
The first three are strikingly different and will facilitate field identification. 

1)—Plants of R. leptocarpa form dense tussocks, from which radiate many lazy culms 
(prostrate to ascending).This habit of the culms appears to be a constant feature, inde- 
pendent of flooding or fire events (pers. obs.). By contrast, R. capitellata although more- 
or-less cespitose, does not form dense tussocks and the relatively few culms are erect to 
strongly ascending. Anderson (1995) reports 30-40 culms for R./eptocarpa in Florida, and 
| have occasionally noted similar numbers in North Carolina, but my sense is that 15-30 
is the norm.Herbarium specimens almost invariably consist of a lesser number of culms, 
because whole plants are rarely collected. 

2)—Foliage and culms of R./eptocarpa are light green, often glaucescent; those of R. 
capitellata are medium to dark green. These differences are retained in dried specimens 
and seem to be independent of light levels. For example, R. leptocarpa growing in clear- 
ings created by logging or military activity show the same pale coloration as those grow- 
ing in shade. Similarly, R. capitellata retains the same dark coloration in dense alder thick- 
ets as when growing in open sites (pers. obs). 


142 SIDA 19(1) 


3)—Spikelet clusters of R. leptocarpa are pale brown; those of R. capitellata are dark 
brown. 

4)—Number of glomerules per spikelet cluster. In R. leptocarpa, glomerules occur 
singly or in pairs at intervals along the culm; in R. capitellata, glomerules are paired or 
tripled—there clearly are 2-3"heads‘arising from separate or branched stalks. Combined 
with greater glomerule width, the overall effect is to give R. capitellata avery n \uch broader 
look to the clusters. 

5)—Mature achene color is pale brown with a pale central bulge (umbo) in R. 
leptocarpa; dark brown with a more contrasting pale bulge in R. capitellata. 

6)—The base of the achene narrows into a shorter stipe in R. /eptocarpa than in R. 
capitellata, as depicted by Blake (1918). 


Habitat 

The widespread Rhynchospora capitellata occupies a broad range of moist to wet, mostly 
open pare from moist dere to wet marshes, creek banks, pond and lake shores, 
beaver ponds, peat bogs, and roadside ditches. By contrast, R. leptocarpa is very limited in 
its habitat preference, occurring almost exclusively in sohagnous streamhead commu- 
nities within longleaf pine ecosystems. These communities receive water from seepage 
which discharges from adjacent uplands and form the head ends and upper portions of 
drainage creeks.In the Carolinas, they are termed “streamhead pocosins,"due to the den- 
sity of shrubs in them.Farther south and on the Gulf Coastal Plain, they are called“baygalls,” 
due to the presence of “bay” species: sweet bay, Magnolia virginiana L, red bay, Persea 
palustris (Raf.) Sarg.;loblolly bay, Gordonia lasianthus (L.) Ellis. When embedded in a mesic 
ravine, as occurs frequently on the Gulf Coastal Plain, the baygall community is called a 
“steephead.” All of these habitats are typically forested, but the narrowness of such 
streamheads normally allows for filtered sunlight to penetrate from the sides. There, R. 
leptocarpa occurs in the semi-shade of trees and tall shrubs, and especially in small open- 
ings created by blowdowns, fire,animal trails, and human disturbance. Downstream, where 


flooding becomes too frequent or of long duration, &. leptocarpa is absent. Table 2 lists 
common associates at streamhead sites. 

In addition, Rhynchospora leptocarpa occasionally inhabits sphagnous margins of 
beaver ponds and human impoundments, especially where tributaries enter.One speci- 
men from southern Mobile County, Alabama, was collected from a “pitcher plant bog,” 
presumably from the edge where such bogs typically grade into streamheads. Ander- 
son (1995) reports R. leptocarpa from the upper edge of a tidal marsh in panhandle Florida; 
it is the only record from a near-maritime habitat. 


Range 

Rhynchospora capitellata is a wide-ranging species of the northeastern United States 
and southeastern Canada. Figure 1 portrays this range, with emphasis on the southern 
margins. Rhynchos| capitellata is common southward through the montane and pla- 
teau regions of the Carolinas, north Georgia, Tennessee, northeastern Alabama, southern 


SORRIE, TOCARPA, AN OVERLOOKED SPECIES 143 


— 2.Common associates of Rhynchospora leptocarpa in streamhead habitats. Species are listed 
alphabetically within each group, not according to frequency. 


TREES HERBS 

Acer rubrum L. Arundinaria tecta (Walter) Muhl. 
Chamaecyparis thyoides (L.) B.S.P. Carex collinsii Nutt. 

llex opaca Aiton Fri lon decangulare L 

Liriodendron tulipi ifera L. Mayaca fluviatilis Aubl. 

Mag! olia virgil viana L. Osmunda cinnamomea ie, 

Nyssa biflora Walter Panicum luc 

Persea palustris (Raf) Sarg. Ala aan ifolia (Michx.) Morong 
Pinus elliottii Engelm. var. elliottii Pingu iCula pri imuliflora Wood & Godfrey 
Pinus serotina Michx. Platan llata (Michx.) Luer 
SHRUBS AND VINES Rhynchospora 1 chalarocephal a Fern. & Gale 


Rhynchospora gracilenta Gra 

Ne aes stenophylla Chapman 
Sphag 

Viola a mul ifo ia 

Woodwardia areolata (L (L.) T. Moore 


Clethra alnifolia L. 
Cliftonia nae (Lam.) Britton ex Sarg. 


Rhododendron aoa n (L .) Torrey 
Smilax laurifolia 

Toxicodendron aa (L.) Kuntze 
Vaccinium formosum H.C. Andrews 
Vibrirntim nnidiim L. 


Missouri,and Arkansas. However, it abruptly becomes rare southward and is absent from 
the rest of the southern Atlantic and East Gulf Coastal Plains. 

In southern Arkansas, R. capitellata appears to be fairly common (Smith 1988), but | 
question whether all spcimens are correctly identified, since three specimens have re- 
cently been annotated to R. glomerata: Bradley Co., Demaree 24539 (NCU); Hempstead 
Co., Demaree 54102 (NCU); Pulaski Co., Merril! 183 (GH). 

In Louisiana, Thomas and Allen (1993) map R. capitellata in five parishes, but two 
cited specimens are actually R. glomerata: Natchitoches Parish, Thomas 725217 (NLU); 
Rapides Parish, Thomas 40834 (NLU). In addition, specimens from Grant, Morehouse, 
Ouachita, Rapides, Sabine, and Vernon parishes (FSU, NCU, NLU) prove not to be R. 
capitellata.A Union Parish specimen is equivocal—the achene size is closer to R.glomerata, 
but shape is closer to R. capitellata: Lewis 3308 (NLU).| have not seen specimens from Red 
River and Winn parishes that were cited by Thomas and Allen. 

Two east Texas specimens are correctly identified as Rhynchospora capitellata: 
Henderson Co., Correll 26688 (GH); Smith Co., Moore, Jr. 888 (GH). Both were collected in 
hillside seepage bogs.A Leon County collection, also from a bog, proves to be R.glomerata 
var.angusta Gale, an overlooked taxon of the West Gulf Coastal Plain:Barkley 13505 (GH, US). 

Rhynchospora leptocarpa is a strict inhabitant of the southern Atlantic and East Gulf 
Coastal Plains (Fig. 1). It occurs from the vicinity of New Orleans, Louisiana, eastward to 
Tallahassee, Florida and adjacent Georgia, then reappears in the Sandhills region of the 


SIDA 19(1) 


144 


25°] 


MILES 


R. 
anitellata 


P 


Fic. 1: Ranges of Rhynchospora leptocarpa and R. capitellata in the eastern United States and Canada. Triangles 


leptocarpa. 


{Dp 


fR canitellata 


Nat 


UU 


Carolinas. Current knowledge suggests that R. leptocarpa rarely occurs on the outer por- 


tion of the Atlantic Coastal Plain. This distribution pattern—a Gulf Coastal Plain compo- 


nent and a disjunct Carolina Sandhills component—occurs in a number of other spe- 


cies (Sorrie et al. 1997).Among the latter is Raynchospora macra 


C.B.Clarke) Small, which 


( 


inhabits open seepage slopes adjacent to streamheads and baygalls. 


SORRIE, RHYNCHOSPORA LEPTOCARPA, AN OVERLOOKED SPECIES 145 


Rhynchospora leptocarpa and R. capitellata are not currently known to be sympatric, 
but further field work may bridge the narrow gap that separates therm. Speciation may 
have occurred during the Pleistocene, when glaciation cycles forced the ancestral entity 
(presumably a species of cool temperate climates) deep onto the coastal plain. Once 
there, this entity survived in open wet habitats analogous to those farther north, and 
also by adapting to localized cool-water environments in shady streamheads. Retreat of 
glaciers allowed the ancestral entity to return northward, while the newly adapted en- 
tity of southern streamheads remained. Several glacial cycles may have been involved 
to complete speciation. Alternatively, the ancestral entity may have occupied a southern 
range; during glacial cycles peripheral populations became adapted to a broad suite of 
cool-water habitats as they migrated northward. 


Repieseniauve specimens: The rollewinigie are dae specimens from throughout the range 
of RI | collections. ALABAMA: Baldwin Co.: acne 
in wet wooded stream with Peltandra sagi ihe lia, upper Majors Creek drainage, 26 Jun 1993, L.C. 
Anderson 14393 (FSU). Geneva Co.: W of route 87,6.7 mi.S of Sea spegnots a nee . 
ae of Pea River, 3 Aug 1996, B.A. roe NCU), Mobile Co.: pine woods near the 
879, C. Mohr s.n. (US); Theodore, pitcher plant bog on Range ee Road, 22 Nov 1980,M. oe 

Sn. aes FLORIDA: Gadsden Co.: Quincy, A.W. Chapman s.n. (GH). Liberty Co.: hammock on road 

12 near Bristol, 22 Jul 1940, a ds.n. (FLAS). Santa Rosa Co.: 5 mi. N of Milton, local in 
dense sweet bay-Cliftonia hammock, 24 Jul 1965, S. McDaniel 6607 (IBE); Eglin Air Force Base, bay 
swamp near head of steephead ravine, Weaver Creek, 22 Sep 1989, Orzell and Bridges 12567 (NCU). 
Walton Co.: springy woodland, bay head between Mossy Head and Deerland, 1956, R.K. Godfrey 
55250 (GA, GH); swamp near De Funiak Springs, 6 Jul 1897,A.H. Curtiss 5926 (FLAS, FSU, GA, GH, NCU, 
US).Washington Co.: acid seep forest ravine above sandy limesink depression pond, 12 Sep 1989, 
S. Orzell and E. Bridges 11883 (NCU). GEORGIA: Decatur Co.: wet woods near Whigham, 10 Aug 

1901, R.M. Harper 1185 (US). Liberty Co.: Fort Stewart, bayhead E of FS 9, locally common in full 
shade, stems lax, 9 Jul 1992, R. Carter 10114 with P Bower (VDB). Thomas Co.: low wet woods along 
small stream, near Thomasville, 14 Aug 1936, D.S. Correl! 6484A (DUKE). Worth Co.: densely shaded 
sphagnum pocket in swamp, Sylvester, H.K. Svenson 6928 (GH). LOUISIANA: Orleans Parish: New 
Orleans, Ingalls s.n. (NY) [cited by Gale 1944]. MISSISSIPPI: Simpson Co.: Saratoga, 3 Aug 1903,5.M. 
Tracy 8616 (GH, US). Stone Co.: University of Mississippi Forest Lands, S of Wire Road and E of route 
15, mossy headwaters of Little Railroad Creek, 21 Aug 1997,B.A. Sorrie 9445 (GH, NCU).NORTH CARO- 
LINA: Hoke Co.: Fort Bragg, McPherson Impact Area buffer zone, sohagnous mucky soil of 
streamhead pocosin, 25 Oct 1991, BA. Sorrie 6060 (bas, GH); Fort Bragg, in troop trail through branch 
of Nicholson Creek, growing prostrate through sphagnum, 28 Sep 1992, BA. Sorrie 7002 (bas, NCU). 

Moore Co:: Sandhills Game Land, E of SR 1104, in sohagnum of old oadbee through tributary of 
Deep Creek, 4 Sep 1997,B.A. Sorrie 9482 (NCU, SWSL). Richmond Co.: Sandhills Game Land, W of SR 
1003, tributary of Naked Creek, large tussocks where fire burned into pocosin,9 Sep 1997,B.A. Sorrie 
9490 (bas, FSU, GA, GH, NCU, VDB). Scotland Go. Camp Mackall (U.S. Army), boggy depression bor- 
dering floodplain of Drowning Creek, 5 Aug 1992, 1.0. Larke s.n. (VDB); Sandhills Game Land, head- 
waters of Little Muddy Creek, frequent in streamhead pocosins with abundant cane, burned Janu- 
ary 1994,17 Aug 1994,B.A. Sorrie 8131 (bas, GA,GH, NCU, VDB). SOUTH CAROLINA: Aiken Co.: Aiken, 
1866, H.W. Ravenel s.n. (GH). Barnwell Co.: pond side, Savannah River Nuclear Operations Area, W.T. 
Batson and W.R. Kelly s.n. (NCU). Stee Co.: swift stream in Carolina Sandhills Refuge, 4.2 m 
of jct.of SC 109 on SC 145, 26 Jul 1994, R. Kral 83865 (VDB). Darlington Co.: Hartsville, flats across 
from paper mill, 5 Jul 1909, We Coker s.n. (NCU). 


146 SIDA 19(1) 


Rarity 

Despite the relative paucity of historical specimens,| believe that Rhynchospora leptocarpa 
is not rare but merely overlooked. In addition to the above collections, | have encoun- 
tered it at 40 sites from North Carolina to Mississippi. Raynchospora leptocarpa is not 
difficult to find in proper habitat. It inhabits a narrow niche, but one which is widely 
distributed over much of the coastal plain. However, these streamheads are usually very 
shrubby and tangled with Smilax laurifolia, thus uninviting to botanists. The period when 
it is most easily detected—July to aks September—is one of formidable heat and hu- 
midity. Add insects, poisonous plants (Toxicodendron vernix (L.) Kuntze), 
and resemblance to several other Rhynchospora (especially R. chalarocephala), and there 
is small wonder that R. leptocarpa remained little known. For many of the same reasons, 
Lindera subcoriacea remained unknown until recently (Wofford 1983), but subsequently 
has been documented from over fifty sites in eight states. The two species co-occur at 
several sites in the Carolina Sandhills. Even with continued fire suppression and conver- 
sion of longleaf ecosystems, searches in appropriate habitat are likely to yield many more 


populations of Rhynchospora leptocarpa. 


ACKNOWLEDGMENTS 


Field work in North Carolina was facilitated by The Nature Conservancy, the North Caro- 
lina Natural Heritage Program, Fort Bragg Military Reservation, and the state Wildlife Re- 
sources Commission. Field work in Mississippi was facilitated by Alan Albritton of the 
UMISS Forest Lands, Julie Moore, Steve Leonard, and Ron Wieland. In Alabama, Bill Finch 
guided me to several high quality longleaf sites. Richard LeBlond contibuted many pleas- 
ant hours afield and provided perceptive comments on this difficult genus. | thank staff 
of the following herbaria for access to collections: DUKE, FLAS, FSU, GA, GH, IBE, NCSC, 
NCU, NLU, SWSL, US, USA, USCH, VDB. | am indebted to R. Kral and an anonymous re- 
viewer for substantial improvements to the manuscript. 


REFERENCES 


Anberson, L.C. 1995. Noteworthy plants from north Florida. VI. Sida 16:581-587. 
Bake, S.F.1918. Notes on the Clayton herbarium. Rhodora 20:21-28 
Britton, N.L. 1892. A list of species of the genera Scirpus and Rhynchospora occurring in 
North America. Trans. New York Acad. Sci. 11:74-93. 
CHAPMAN, A.W. 1860. Flora of the southern United States. American Book Company, New 
York, NY. 
. 1883. Flora of the southern United States. Second Edition. American Book 
Company, New York, NY. 
. 1897, Flora of the southern United States. Third Edition. American Book Com- 
pany, New York, NY. 
Crewe, A.F.1985.Guide to the vascular plants of the Florida Panhandle. University Presses 
of Florida, Tallahassee. 


SORRIE, RHYNCHOSPORA LEPTOCARPA, AN OVERLOOKED SPECIES 147 


Goprrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United 
States. Vol. 1. Univ. of Georgia Press, Athens. 

Hatch, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas 
Agric. Exp. Sta. Texas A & M University, College Station. 

Houmeren, PK., N.H. Houmeren, and L.C. Barnett (eds.). 1990. Index Herbariorum. Part 1:The 
Herbaria of the World, 8th edition. New York Botanical Garden, Bronx, NY. 

Jones, S.B,, Jr.and N.C. Coie. 1988. The distribution of the vascular flora of Georgia. Dept. of 
Botany, University of Georgia, Athens. 

Kartesz,J.1.1994.A synonymized checklist of the vascular flora of the United States, Canada, 
and Greenland. Second ed., two vols. Timber Press, Portland, Oregon. 

Raororb, A.E., H.E. AHLes, and C.R. Beit. 1968. Manual of the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill. 

Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas. 2nd ed. 
Privately printed. 

Sorrie, B.A.,B. van EerDEN, and M.J.Russo. 1997.Noteworthy plants from Fort Bragg and Camp 
Mackall, North Carolina. Castanea 62:239-259. 

Tomas, R.D.and C.M. ALLEN. 1993. Atlas of the vascular flora of Louisiana. Vol. 1.Louisiana 
Dept. Wildlife & Fisheries and The Nature Conservancy, Louisiana Field Office, Baton 
Rouge. 

Worrorb, B.E. 1983. A new Lindera (Lauraceae) from North America. J. Arnold Arbor. 64: 
325-331. 

Wunber.in, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, 
Gainesville. 


148 SIDA 19(1) 


BOOK REVIEW 


— Primack. 2000.A Primer of Conservation Bi 1 Edition. (ISBN 0-87893- 
32-3, pbk.). Sinauer Associates, Inc.23 Plumtree Road, Sunderland, MA 01375- 0407, 
tien (publish@sinauer.com) $ 31.95 pbk. 319 Pp. 95 illustrations. 


This book has as its stated intention, “to provide a brief but thorough introduction to the major 
concepts and problems of the field,” and it certainly fulfills its mission. The book is divided into 5 
chapters, including: 1) Conservation and Biological Diversity, 2) Threats to Biological Diversity, 3) 

onservation at the Population and Species Levels, 4) Conservation at the Community Level,and 5) 
Conservation and Sustainable Development. Within each chapter, concepts are introduced in sec- 
tions in bold print, and include numbered tables and figures. Ancillary, explanatory material is pre- 
sented in numbered Boxes in the first chapter only. More of such boxes to further explain concepts 
in the other chapters would be welcome 

The level of complexity of the text reinforces its purpose as a basic, introductory undergradu- 
ate textbook for an upper level course.| would recommend that students have basic biology and 
ecology as prerequisites so that time can be spent on supplementary articles or perhaps a short 
term paper! ine nal aa listed at the end of each chapter and suggested readings inte- 
grate the i pts presented within each chapter into its respective overall theme very 

well. 

The book does an excellent job of incorporating elements of anthropology and economics 
into the basic biological framework that supports research and implementation of modern ap- 
proaches to solving problems in conservation biology. In addition, oc appendix with “Selected 
Environmental Organizations and Sources of Information," extensive bi phy, and index make 
it easy find concepts throughout the 

estion for improvement would be to update the two Boxes of Chapter one, where 


box intends to explain the hierarchy used in taxonomy but unfortunately implies that organisms 
are grouped by their overall similarity instead of by their hypothesized evolutionary relationships. | 

ould recommend explaining the hierarchy in terms of genealogy, so that it — not che 
ve what students will learn about phylogenetics in any subsequent system Irs will 
take. It may also be valuable to introduce the Sie of eke allopati and protic 
stasipatric) speciation, perhaps somewhere in the chapt e| nd 
species levels. With even a limited discussion of these un slinaed to but not ciiseared upon in 
Box 1.2 of Chapter One, other concepts such as the Founder and Bottleneck Effects may be better 
understood and the profound effects of habitat fragmentation better appreciated. In future edi- 
tions, the tremendous contribution that molecular studies, including DNA fingerprinting techniques, 
and intraspecific genetic variation work, have given to modern conservation studies, should be 
added 


— 


espite these few comments, | know of no finer text to introduce the subject of Conserva- 
tion Biology to undergraduates, at a global level and with such a synthetic approach. The text, with 
its durable back and high-quality pages, its figures conveniently placed, its references copious, suc- 
cinct yet thorough summaries and suggested readings, all ma ake this text easy to read and opti- 
mally student-friendly.| highly recommend it for everyone planning to teach such a course —John 
J. Pipoly Ill, Botanical Research Institute of Texas, 509 Pecan Street, Fort He TX 76102-4060, U.S.A. 


SIDA DA19(1): 148. 2000 


OCCURRENCE, DISTRIBUTION, AND ECOLOGY OF 
ALOCASIA, CALADIUM, COLOCASIA, AND XANTHOSOMA 
(ARACEAE) IN THE SOUTHEASTERN UNITED STATES 


Brett E.Serviss 


Department of Biological Sciences 
Mississippi State University 
Mississippi State, MS 39762, U.S.A. 
bes3@ra.msstate.edu 


Sidney T.McDaniel 


Department of Biological Sciences 
Mississippi State University 
Mississippi State, MS 39762, U.S.A 


Charles T. Bryson 
USDA, ARS 
Southern Weed Science Research Unit 


Stoneville, MS 38776, U.S.A. 
cbryson@ag.gov 


ABSTRACT 


Eight species in four genera of Aroideae (Araceae) are introduced, cultivated, and variously estab- 
lished in the southeastern United States. Keys, illustrations, descriptions, distribution and ranges, 
habitat preference, flowering and fruiting periods, and variation in both vegetative and sexual re- 
productive characteristics are provided for the genera, species, and varieties. Additional informa- 
tion on the cultivation of all species and the weedy attributes of Colocasia and Xanthosoma are 
discussed. 


RESUMEN 


Ocho especies de cuat de Aroid ‘ ) estan introducidas, cultivadas, y establecidas 
en el sureste de los Estados Unidos. Se ofrecen claves, ilustraciones, descripciones, distribucion y 
areas, habitat ee periodos de floracion y fructificacién, y variacidn en las caracteristicas 

getativas y los géneros, especies, y variedades. Se discute informacién adicional 
Snes ag aultive ae eee las especies y las caracterfsticas como malas hierbas de Colocasia y 
Xanthosoma. 


INTRODUCTION 


The Araceae (including Lemnaceae; Mayo et al. 1995) or arum family is represented world- 
wide by 105 genera and about 3,200 species (Croat 1998; Mayo et al.1997).At present 16 
genera (excluding Acorus that it is now placed in the Acoraceae) and approximately 31 
species occur in the continental United States and Alaska.Of these 31 species, only about 


SIDA 19(1): 149-174. 2000 


150 SIDA 19(1) 


one-half are native; the remaining exotics are established to various degrees as compo- 
nents of our flora.Alocasia, Colocasia,and Xanthosoma are among those non-native natu- 


ralized (or potentially so) genera and are represented by seven species in the United 
States. The origination of these genera from various tropical regions and the characteris- 
tics of the naturalized species discussed herein, imply an inherent plasticity for adapta- 
tion to different climatic conditions and habitat types.in the southeastern United States, 
differences in seasonal temperature range, habitat type, and floristic species composi- 
tion all combine to provide some environmental deviation from the conditions in which 
these species evolved. Five of the eight species are experiencing some degree of range 
expansion since introduction, and three varieties of Colocasia esculenta (Colocasia 
esculenta var. antiquorum, Colocasia esculenta var. aquatilis, and Colocasia esculenta var. 
nymphaeifolia) and Xanthosoma sagittifolium are rapidly becoming invasive. Intraspe- 
cific morphological variability within Colocasia, and an overall similarity in appearance 
between the several varieties of Colocasia esculenta and certain species of Alocasia and 
Xanthosoma allow for continual uncertainty in field identification of this group.Ambigu- 
ity in terms of species identification and potential for invasiveness of these introduced 
Araceae necessitate a better understanding of their morphological and ecological vari- 
ability and character in the United States. 

Currently, Alocasia macrorrhizos and Xanthosoma sagittifolium are restricted in their 
naturalized ranges to central and southern Florida and southern Texas, while Colocasia, 
in addition to occurring throughout Florida and most of eastern Texas, also occurs in 
scattered locations throughout the Gulf coastal region. Colocasia esculenta var. aquatilis, 
Colocasia esculenta var.nymphaeifolia, and Colocasia esculenta var.antiquorum are estab- 
lished as far north as east central Texas, northern Louisiana, and throughout the lower 
one-half of Alabama, Georgia, and Mississippi. These three varieties of C. esculenta are 
invasive weeds of semi-aquatic and aquatic conditions throughout the Gulf coastal re- 
gion. Xanthosoma sagittifolium, while not established as a serious weed, is becoming 
more prevalent in peninsular Florida and southern Texas. 

Colocasia was introduced into the United States as a food crop for its edible tubers 
and for ornament, while Xanthosoma and Alocasia were introduced for their value as 
ornamentals with minimal consideration as a food source (Neal 1991;Vaug! 
1997). Several varieties of Colocasia are grown commercially for tuber production, as the 
tubers are a source of food and propagules for ornamental use. We would like to clarify 
and delineate among the intergeneric, intraspecific,and ecological differences between 
the species and varieties of Alocasia, Caladium, Colocasia,and Xanthosoma in the United 
States. 


Geiccler 


TAXONOMY 


Alocasia, Caladium, Colocasia and Xanthosoma are currently classified in subfamily 
Aroideae (Mayo et al. 1997). These four genera are not monophyletic as once thought 
when formerly placed in the old subfamily Colocasioideae, thus paleotropical Alocasia 
and Colocasia remain in tribe Colocasieae and subtribe Colocasiinae, while the neotropical 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 151 


genera of Caladium and Xanthosoma are separated into tribe Caladieae and subtribe 
Caladiinae (French et al. 1995; Mayo et al. 1997). Subfamily Aroideae (based primarily on 
Croat 1988, 1998;Grayum 1994; Mayo 1997) are perennial, predominately terrestrial, rarely 
hemiepiphytic or epilithic herbs, which produce white or yellow to orange latex sap. 
Plants caulescent to acaulescent, hypogeal, stems sometimes aerial, erect to decum- 
bent, often rhizomatous and tuberous; leaves petiolate, ovate, ovate-lanceolate, ovate- 
elliptic, or triangular-ovate, variously peltate or not, basally sagittate, hastate, or cordate; 
petioles elongate and fleshy, with the terminal petiole housing the newest unemergent 
leaf; venation usually reticulate, sometimes parallel-pinnate. New leaves arise singly from 
the apical portion of the stem and emerge from a cavity within the terminal petiole. 
Laticifers present, simple, articulated, or anastomosing; trichosclereids absent. Plants 
monoecious (occasionally polyg ious),and ianth parts absent; 
at anthesis the spadix is usually open, with or without a ‘oninel ppnenaix and a region 
between the staminate and pistillate flowers comprised of synandrodes (sterile stami- 
nate flowers), stamens are connate and arranged in synandria; fruit a 1-numerously seeded 
berry, endosperm copious or not; predominately insect pollinated by various taxa of 
beetles, flies,and bees. Our species of Aroideae perenniate from subterranean tubers and 
or rhizomes. Production of these perenniating structures allows for efficient and prolific 
vegetative reproduction. 

Many Aroideae form asexual colonies of genetically identical individuals. This is the 
typical method of reproduction for representative genera in the US and one of the pri- 
mary factors allowing for naturalization. Seed production also occurs if environmental 
conditions are conducive, but this is rare in the US. Throughout the plant, but concen- 
trated in the tubers, are deposits of poisonous capsules of pointed calcium oxylate crys- 
tals. Plants are poisonous unless cooked or prepared in such a way as to remove or neu- 
tralize the crystals. These can cause numbing and irritation of the skin and mucus 
membranes and problems with digestion if foliage and tubers are eaten raw or incom- 
pletely cooked (Vaughan & Geissler 1997). 


KEY TO ALOCASIA, CALADIUM, COLOCASIA, AND XANTHOSOMA 
IN THE SOUTHEASTERN UNITED STATES 
1. Leaves predominately green and tinged with purple coloration to completely 


purple, or green with white, pink, or red coloration (leaves rarely uniformly green in 
Caladium). 


2. Leaf margins not crenatet nly slight ly ‘lamina ariously mottled with white, 

pink, or red coloration, or some combination thereof Caladium 
2. Leaf margir 1S variously crenate or not, leaves uniformly purple or a combination 

of purple and 


3. Newly formed leaves blue-violet, lamina becoming green as they enlarge and 
age, midvein, primary lateral veins, and petioles violet or becoming yellow 
Xanthosoma 


3. Newly formed and older leaves green, petioles green and often glaucous 
4. Veins of upper leaf surface conspicuously raised above the blade and leaf 
margins strongly crenate; leaves not or only slightly peltate Alocasia 


SIDA 19(1) 


152 
4. Veins of upper leaf surface flush with blade to slightly raised above it, the 
entire upper surface of the leaf appearing flat or leaf margins weakly 
crenate; leaves eeuleuy peltate Colocasia 
1. Leave tially green, glaucous or glaucescent on the lower surface 
5. Plant arising from well-developed, subterranean tubers. 
6. Basal 2 lobes of leaves apically acute; leaves not peltate Xanthosoma 
6. Basal 2 lobes of leaves apically rounded; leaves peltate orn 
7. Veins of upper leaf surface conspicuously raised above le blade and leaf 
margins strongly crenate; leaves not peltate to peltate Alocasia 
7. Veins of upper leaf surface flush with blade to slightly raised above it, the 
entire upper surface of the leaf appearing flat and leaf margins weakly 
crenate; leaves peltate Colocasia 
5. Plants lacking well-developed tubers, or if tubers present then much reduced 
Plants producing long slender rhizomes above and or below the ground; stems 
arising from a slightly tuberous caudex 
9. Basal 2 lobes of leaves rounded; rhizomes usually emerging and running 
for most of their length above or below ground Colocasia 
9. Basal 2 lobes of leaves acute; rhizomes usually completely running below 
round Xanthosoma 
Peltandra 


8. Plants not rhizomatous and caudex not tuberous 


NOTE: Peltandra is included in this key because of morphological similarities and possible confu- 
sion with the other genera 


pinae 


SPECIES ORIGIN, OCCURRENCE, RANGE, AND DESCRIPTION 


ALOCASIA 
Alocasia (Schott) G. Don in Sweet, Hort. Brit. ed. 3.631. 1839.nom.cons; Krause in Engler, 
Pflanzenr.4.23E.71:71.1920. Tyee species (conserved):Alocasia cucullata (Lour.) G.Don,Arum 


cucullatum Lour 
Colocasia sect. Alocasia Schott in Schott & Endlicher, Melet. Bot. 18.1832. Ensolenanthe Schott in 
Bonplandia 9. 368. 1861 
Xenophya Schott in Ann. Mus. Bot. Lugduno-Batavum 1:124. 1863. 
Schizocasia Schott ex Engler in Bot. Jahrb. 1:185. 1880-1881. 
Panzhuyuia ZY. Zhu in J. Sichuan Chinese Medicinal School 4(5):49, 1985. 


Perennial caulescent or acaulescent herbs usually from a creeping tuberous-rhizome 
caudex, with or without elongate rhizomes, reproducing asexually by secondary tubers, 
tuber offshoots, and or rhizomes. Above ground stems erect to decumbent; leaves ovate 
to triangular-ovate, basally sagittate or rarely slightly peltate; spadix pedunculate, arising 
axillary and subtended by 1 or 2 acute to acuminate bracts, st te and pistillate flowers 
separated by a section of synandrodes (sterile staminate flowers), appendix present; 
spathe present, longer than spadix, constricted basally to form a tube which encloses 
the spadix until anthesis; the basal portion remaining tightly closed; staminate flowers 
comprised of truncate synandria of 3-8 stamens; pistillate flowers 1-4 carpellate, 1-4- 
locular, stigma weakly 3-lobed, ovules 1-numerous, placentation basal; fruit a 1-numer- 
ous seeded berry sometimes with portions of the stigma persistent; ovules 6-10, 
orthotropous, hemiorthotropous, hemianatropous, or anatropous, funicles short; seed 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 153 


subglobose to ellipsoid, testa smooth or scabrous, endosperm copious. 
About 70 species native to Indomalaysia and throughout southern Asia. Three spe- 
cies occur in the US. Alocasia macrorrhizos and Alocasia odora are both established. 
Etymology.—colocasia an arum, and alocho, a prefix meaning bedfellow or spouse; 
depicting the morphological similarity between Alocasia and Colocasia. 


KEY TO SPECIES OF ALOCASIA 


1. Leaves and foliage green (foliage rarely purple in A. macrorrhizos, if so, then leaves 
not peltate). 
2. Mature leaves not peltate; leaves on juvenile plants sometimes slightly peltate 
macrorrhizos 
2. Mature leaves obviously peltate 2. A.odora 
1. Leaves and foliage purple, petioles sometimes green; leaves peltate 3.A. plumbea 


1. Alocasia macrorrhizos (\.) G. Don var. macrorrhizos in Sweet, Hort. Brit, ed. 3.631. 
1839. Hook.f. Fl.Br.Ind.6:526. 1893. Petch, Ann. Roy. Bot. Gard, Peradeniya 7:53.1919. 
Krause in Engler, Pflanzenr. 4. 23E. 17:84. 1920. (Fig. 1). Arum macrorrhizon L., Sp.P1.965. 
1753.Caladium macrorhizon R.Br. Prod. 336. 1810.Alocasia macrorrhiza (L.) Schott in Schott & 
Endl., Melet.Bot. 18.1832. Alocasia macrorrhizon (L.) G.Don in Sweet, Hort. Brit,,ed. 3.631,1839. 
Colocasia macrorrhiza Kunth, Enum. Pl. 3:39. 1841. 


Arum peregrinum L., Sp. Pl. 966. 1753. 
Arum mucronatum Lam., Encyc. 3:12. 1786. 
Arum indicum Lour., Fl. Cochinch. 536, 1790. aes indica (Lour.) Kunth, Enum. PI. 3:39. 1840. 
Alocasia indica (Lour.) Spach, Hist. Nat. Veg. a 
Arum cordifolium Bory, Voy. lles. Afrique 1:376. 1 
Caladium giganteum Blume, Cat. Gew. Buitenz. 1 i 1823. 
rum punctatum Desf., Cat. Hort. Par., ed. 3.385. 1829, 
Caladium glycyrrhizum Fraser in Hook. Bot. Misc. 1:259. 1830. 
Calla badian Blanco, Fl. Filip.658. 1837. 
Calla maxima Blanco, FI. Filip.658. 1837. 
Arum grandifolium Blanco, op. Cit., ed. 2.1845. 
Alocasia alba Schott in Oestr. Bot. Wochenbl. 2:59, 1852. 
Caladium indica Hort. ex C. Koch, Berl. Allg. Gartenz. 136. 1857. 


Caulescent herbs up to 5 m in height from a tuberous caudex; stems to 1.5 m tall and 20 
cm in diameter, becoming thickened and caudex-like, erect or often falling over with 
age and resting on the ground, basally covered with dry, chartaceous, persistent cataphylls, 
adventitious roots sometimes present, rhizomes present or absent, when present vari- 
able in length, elongate, white to whitish-green; leaves 35-160 cm long and 30-90 cm 
wide (often smaller in dimensions if on young, damaged, or spring emergent plants) 
ovate to ovate-triangular, apically subacute to short acuminate, basally sagittate, young 
leaves sometimes peltate, margins strongly undulate, primary lateral veins 4-8 per side, 
midvein and lateral veins prominently raised above the lamina; petioles 70-130 cm in 
length, lamina and petioles green; inflorescence of 1-numerous spadices, peduncles 20— 
50. cm long, spathe 11-35 cm long, 3.5-6.2 cm wide, chartaceous, apically withering with 
age, greenish-white to green or glaucous, becoming peach or light orange with age, 


RNG (hg, 


te 
ke 


& 


, é 
QU 


Fig. 1. Alocasia macrorrhizos (L.) G. Don var. macrorrhizos. Young plant, general habit. 


SIDA 19(1) 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 155 


spadix 11-35 cmin length, yellowish to cream, appendix 7-21 cm long, fertile staminate 
region 2-7 cm, sterile staminate region 0.3-2 cm long, fertile pistillate region 1.2-5 cm 
long, pistillate flowers emit the odor of ether prior to fertilization; berries ovoid, orange 
to scarlet, 8-10 mm long and 5-8 mm in diameter, usually containing 5-6 ovules. 

Some species of Alocasia, most importantly A. macrorrhizos and A. odora, are impor- 
tant food crops in tropical regions, particularly Asia, the south Pacific, and to a lesser 
extent, Africa (Bailey 1997; Neal 1991). The actual crop and source of nutrition is the large, 
edible, subterranean tubers. The tubers provide high levels of dietary starch (approx. 25% 
by weight), ascorbic acid, low amounts of vitamin A, B complex vitamins, and small 
amounts of protein (Bailey 1997; Neal 1991; Vaughan & Geissler 1997). Through use as 
crops and ornamentals A. macrorrhizos and other species of Alocasia have been distrib- 
uted and established throughout the subtropics and tropics. 

Alocasia macrorrhizos was introduced into the United States from tropical Asia and 
occurs sporadically in central and southern Florida and southern Texas. While widely 
cultivated in the southern US, it is only truly established in central and southern penin- 
sular Florida, where climatic conditions permit essentially year-round growth and little 
to no enforced dormancy. Its ability to reproduce vegetatively has aided naturalization. 
Its rate of expansion is reduced in the absence of disturbance and where soils are with- 
out adequate moisture and drainage. Once established, Alocasia macrorrhizos appears 
to tolerate a wide range of condition variability (drought, periodic flooding, variable light 
intensity, and occasional hard freezes) and interspecific interference. The rate of range 
expansion by A. macrorrhizos may at least in part be impeded because it usually colo- 
nizes vegetatively by secondary or daughter tuber formation. This method of establish- 
ment is not normally as conducive to invasion as is rhizome production in Colocasia and 
Xanthosoma.The apparent inability of A. macrorrhizos to produce much fruit (in the United 
States) for dispersal and potential long-range colonization also probably contributes to 
its limited expansion. 

In the United States flower and fruit production typically occur from late summer 
into winter, though flowering can occur at any time of year. One to several inflorescences 
are produced per plant at any one time. Alocasia macrorrhizos is self compatible but 
probably requires out-crossing between different genotypes for full fertility (obviously 
environmental conditions are also important here).In the US most colonies arise from a 
single genotype, which may explain the apparent lack of fecundity in our plants. The lack 
of appropriate insect pollinators is yet another consideration.When fruit is produced the 
number of berries are usually few (10-15 per spadix). 

Alocasia macrorrhizos prefers moderate to well-drained but continuously moist, 
moderately to highly acid soil. It also seems to require at least moderate disturbance for 
establishment. |t can be found along wet woodland edges, thickets, roadsides, old fields, 
open areas of waste ground, vacant lots, or persisting and spreading at home sites. Once 
established it spreads vegetatively, by secondary tubers which produce many small plant- 
lets or to a limited extent along tuberous rhizomes. These plantlets originate and grow 


156 SIDA 19(1) 
for an extended period of time remaining attached to the main plant and may or may 
not eventually separate. Alocasia macrorrhizos is semi cold tolerant and can withstand 
limited exposure to freezing temperatures, regenerating foliage afterwards. A form with 
purple foliage very similar in color to Alocasia plumbea is sometimes cultivated and oc- 
casionally reverts back to the wild-type green coloration. 

Common names.—pai, taro, cunjevoi, giant taro, ape giant, elephant-ear 


2. Alocasia odora (Lodd.) Spach, Hist. Nat. Veg. Phan. 12:46. 1846; (Roxb.) K.Koch, 1.¢. 1854. 
Arum odorum Roxb., Hort. Beng. 65. 1814; Roxb., Fl. Ind. 3:499.1832; Lodd,, Bot. Cab. 5: t. 416. 
1820; Wight, Ic. Pl. Ind. Or. 3(6): t. 797. 1844. Caladium odoratum Lodd., Bot Cab. t.416. 1820. 
Caladium odorum (Lodd_) Lindl., Bot. Reg. 8: t.641. 1822. Colocasia odora Hort. ex Brongn. in 
Nouv. Ann. Mus Par. 3. 145.1834. 
Alocasia commutata eet in Oestr. Bot. WochembIl. 409. 1854. 
aladium od t.ex C. Koch, Berl. Allg. Gartenz. 20. 1857. 
Extremely similar to Alocasia macrorrhizos but differing by having peltate leaves and 
foliage often glaucescent. Also similar to Xanthosoma sagittifolium but basal leaf lobes 
are apically rounded, whereas those of X. sagittifolium are apically acute.Alocasia odora is 
native to southeast Asia and the Philippines and is cultivated for food and ornament 
throughout the tropics and for ornament in the extreme southern US. Alocasia odora is 
spontaneous in southern Florida and much of the material reported or determined as A. 
macrorrhizos is in fact A. odora. Alocasia odora is likely only a subspecies or variety of A. 
macrorrhizos. Ecology is similar to that of A. macrorrhizos. 
Common names.—taro, elephant-ear, ape giant. 


Ee 


3. Alocasia plumbea Van Houtte in Fl. Des. Serres, Sér. 2.6. 93:t. 2206. 1875. 


Alocasia indica Schott in Oestr. Bot.Wochenbl. 410. 1854. 
Alocasia metallica Schott in Oestr. Bot. Wochenbl. 410.1854. 


Caulescent herbs 1-3 m in height from a tuberous caudex; stem to 1 m tall, basally cov- 
ered with dry,chartaceous, persistent cataphylls, rhizomes present or absent, when present 
variable in length, elongate, white to white-green, often tinged with pink or purple col- 
oration; leaves 20-100 cm long and 15-65 cm wide, ovate-triangular, apically subacute 
to short acuminate, basally sagittate, often peltate, margins undulate, primary lateral veins 
4-8 per side, midvein and lateral veins prominently raised above the lamina, petioles 
40-110 cm in length, lamina adaxially dark green-purple, abaxially variable-purple, purple- 
green, red-brown, or yellow-green; inflorescence of 1-numerous spadices, peduncles 25- 
50cm long, spathe 10-20 cm long, 3—7.5 cm wide, chartaceous, apically withering with 
age, purple-pink to pink, becoming peach or yellow with age and often violet-striate, 
spadix 10.5-19 cm in length, yellowish to crear, appendix 6.5-13 cm long, fertile stami- 
nate region 1.5-2.5 cm, sterile staminate region 0.3-0.7 cm long, fertile pistillate region 
1.5-2.5 cm long, pistillate flowers emit a slight odor of ether prior to fertilization; ovary 


containing 3-5 ovules; fruit not observed. 
Similar in form and habit to A. macrorrhizos but differing in its smaller and less ro- 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 157 


bust stature, purple to purple-green foliage (petioles often pink or pinkish-white with 
purple or red spots when on young plants, or plants growing under shade conditions), 
and pink to lavender spathe. It is not currently naturalized in the US but is often culti- 
vated in extreme southern Florida, Louisiana, and Texas, and readily persists subsequent 


to cultivation. 
Common names.—purple taro, purple ape. 


CALADIUM 
Caladium Vent., Mag. Enc. 4(16):463. 1800-1801;Vent., Descr. Pl. Nouv. Jard. Cels 30. t. 30. 
801;Vent., Arch. Bot. (Leipzig) 2(3):347.1801;Vent., J. Bot. (Schrader) 4(6):316. 1801. 
Engler, Pflanzenr. 4. 23E. 71:23. 1920. Lecroryee species: Caladium bicolor (Ait) Vent. Arum 
bicolor Aiton. 


Cyrtospadix C. Koch, Ind. Sem. Hort. Berol. 1858. 
Arosma Rafin., Fl. Tellur. 3:66. 1886. 
Aphyllarum S.Moore in Trans. Linn. Soc. London, Bot. Ser. 2, 4:501. 1895. 


Perennial acaulescent herbs from a creeping tuberous rhizome, reproducing asexually 
by daughter tubers. Above ground stems erect; leaves ovate to triangular-ovate, basally 
sagittate, peltate; spadix pedunculate, arising axillary and subtended by 1 or 2 acumi- 
nate bracts, staminate and pistillate flowers separated by a section of synandrodes (ster- 
ile staminate flowers), appendix absent; spathe present, longer than spadix, constricted 
basally to form a tube which encloses the spadix until anthesis; the basal portion re- 
maining tightly closed; staminate flowers comprised of angular synandria of 3-5 sta- 
mens; pistillate flowers 1-4 carpellate, 1-2 or incompletely 3-locular, stigma discoid, ovules 
I-numerous, placentation subbasal, pseudoaxile, or parietal; fruit a 1-numerous seeded 
berry;ovules 1-20, anatropous, funicles short:seed ovoid to ellipsoid, testa variously cos- 
tate, endosperm copious. 

About 12 species all originating in the neotropics. Several species cultivated in the 
US, currently none established, es C. bicolor is widely cultivated. 

Etymology.—Latin:keladi, of Malaysian origin, vernacular name for several Colocasioid 


a 


and Caladioid genera. 


Caladium bicolor (W. Ait.) Vent., Mag. Enc. 4(16):464. 1801;Vent., Descr. Pl. Nouv. Jard. Cels 
30. t.30.1801;Vent. Arch. Bot. (Leipzig) 2(3):348.1801;Vent., J.Bot. (Schrader) 4(6):319, 
1801. Engler, Pflanzenr. 4. 23E. 71:31. 1920. (Fig. 2). Arum bicolor W. Ait., Hort. Kew. 3:316 
1789 


Caladium surinamense Miq., Delect. Sem. Hort. Amst. 3. 1853. 
Caladium regale Lem., Illustr. Hort. 9. t. 316. 1862. 
Caladium x hortulanum Birdsey, Cult. Aroids 42.1951. 


Acaulescent herbs from 0.2-1 m in height from a tuberous rhizome, tuberous caudex 
well-developed, plant without elongate stoloniferous rhizomes; leaves 5-35 cm long 
and 5-21 cm wide, ovate to ovate-triangular or ovate-elliptic, apically rounded to acute, 
basally sagittate, peltate, margins subundulate, angular, or entire; primary lateral veins 


158 SIDA 19(1) 


Fic. 2. Caladium bicolor (W. Ait.) Vent. General habit. 


3-8 per side, lamina ranging in color from entirely green to green and variously mottled 
with white, pink, or red or some combination thereof, sometimes lamina predominately 
white or red, midvein and lateral veins raised above the blade on abaxial side, petiole 
10-50 cm in length, green to red, purple, or black, often green and splotched with purple, 
sometimes pink basally; inflorescence of 1-numerous spadices, peduncles 28-40 cm long, 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 159 


spathe 7-25 cm long, chartaceous, broadly acute, green externally, white to white-green 
on the inner surface; spadix 9.5-14 cm long, white to yellowish, appendix absent or much 
reduced, region of fertile staminate flowers 3.5-5.5 cm long, region of sterile staminate 
flowers 1-2 cm long, region of fertile pistillate flowers 3.5-5 cm long, pistillate flowers 
essentially without odor, ovaries with 1-numerous ovules; berries white. 

A highly variable species comprised of numerous varieties separated primarily on 
the basis of foliage coloration. Commonly cultivated in the southeastern US for orna- 
ment but not currently documented as naturalized. Caladium bicolor is a prolific seed 
producer and is spontaneous in new growth forests and moist disturbed areas through- 
out the tropics. It should be expected along moist disturbed sites, woodland edges, ham- 
mocks, and fields in southern Florida. 

Common names.—caladium, elephant's-ear, angel-wings, mother-in-law plant. 


COLOCASIA 
Colocasia Schott in Schott and Engl., Melet. Bot. 18. 1832, nom. cons; Krause in Engler, 
Pflanzenr. 4. 23E. 71:62. 1920. Type species (conserved): Colocasia antiquorum Schott, Arum 
colocasia L., Colocasia esculenta (L.) Schott. 
Leucocasia Schott in Oestr. Bot. Wochenbl. 7:34. 1857. 

Perennial caulescent to acaulescent herbs from a creeping tuberous-rhizome; tubers 
and rhizomes usually present; asexual reproduction by means of secondary or daughter 
tubers, and or rhizomes. Above ground stems erect; leaves ovate to triangular-ovate, ba- 
sally cordate, cordate-sagittate, or sagittate, typically peltate, often glaucous to glaucescent 
abaxially; spadix pedunculate, arising axillary and subtended by 1 or 2 acute to acumi- 
nate bracts, staminate and pistillate flowers separated by a section of synandrodes (ster- 
ile staminate flowers), appendix present; spathe present, 2-6 times longer than spadix, 
constricted basally to form a tube which encloses the spadix until anthesis; the basal 
portion remaining tightly closed; staminate flowers comprised of truncate synandria of 
3-6 stamens; pistillate flowers 1-locular, stigma discoid-capitate or weakly lobed, ovules 
numerous, placentation parietal; fruit a 1-numerous seeded berry with a persistent stigma; 
ovules numerous, hemiorthotropous, funicles long; seed ovoid to ellipsoid, testa costate, 
endosperm copious. 

Seven or eight species primarily of tropical Asia, some in Africa. Two species estab- 
lished in the US but only certain varieties of Colocasia esculenta widely naturalized. 

Etymology.—Greek: kolokasia, term used for the root of Nelumbo nucifera Gaertn. 
(water lotus) (Nel eae), thus applied to Colocasia because of its edible tubers. 


KEY TO SPECIES OF COLOCASIA 


1. Leaves and foliage predominately to entirely green, often somewhat glaucous on 
the lower surface, petioles and lamina sometimes with a reddish or purplish tinge; 
plants well-established in southeastern United States. 

2. Rhizomes up to 25 dm long; tubers usually poorly developed or nearly absent; 
widely naturalized (some material of C. esculenta var. antiquorum will key here). 


SIDA 19(1) 
3. Rhizomes originating at apex or middle of tuber body 1.C, esculenta var. aquatilis 
iginating only at very base of tubers 8.C. esculenta var. nymphaeifolia 
2, Rhizomes efien reduced in length or absent; tubers normally large and well- 
develop 


ed. 
4, eae large, few; terminal appendage of spadix short or oe abse 
rhizomes usually short or absent 3.C.e ans var. esculenta 
4. Tubers small, few to numerous, often with short or long rhizomes, occasion- 
ally rhizomes up to 25 dm long; terminal appendage of spadix elongated. 
5. Plants of aquatic areas; relatively wide-spread 2.C. esculenta var. antiquorum 
5. Plants of upland areas with well-drained soils, often occurring and sae sist- 
ing along field edges where cultivated but rarely occurring outside th 
areas 


eculent var. 1 lobules 

. Leaves with purplish or blackish sara or white- coreoune on ae under s 

face; plants not truly naturalized in the tates. 

6. Leaves white-glaucous on ieee and lacking purpl 
usually whitish; spathe white to cream 

6, Leaves not 


coloration,veins 

9.C. gigantea 
white-glaucous on lower surface, may be glaucous-green, blue or 
purple, and at least with some purple coloration on veins and ¢ or blade, veins 
never with white coloration; spathe yellow- green to orange, Oo 

7. 


rac ar n ( 


k, 

per surface (sometimes tinged with pt a petioles and 
inet surface veins purple or aque black, Gah margins usally pups 
8. Lamina dark green, almost appea 


ngt ‘ 
veins on the lower surface, nie or without purple margins; plas large, 

often 1.5-2.0 m tall culenta var. fontanesii 
8. Lamina bright green; leaves usually with veins purple, sometimes green 


on the lower surface, margins purple; plants small, ae 1m or less tall 


4.C. esculenta var. euchlora 
7. All or at least part of the upper surface of leaves dark purple to purple-black 
9, Leaves completely purple at maturity, almost black, glaucous-purple on 


the lower surface, primary veins dark purple to purple-black 5.C. esculenta var. 


fontanesii 
9. Leaves green with spots or broad bands of purple between the veins on 


the upper surface, glaucous-green on the lower surface, primary vein 


seculeae var. illustris 
Another variety of Colocasia esculenta—C. esculenta var.acris (R. Br.) A.F.Hill, Bot.Mus. Leafl. 7:7. 1939 
Caladium acre R. Br., Prodr. Fl. Nov. Holl. 336. 1810. Colocasia acris Schott in Schott & Endl., Melet. Bot 
18. 1832. Colocasia antiquorum Schott var. acris Schott, Syn. Aroid. 42. 1856; Prodr. Syst. Aroid 
1860; Engl. in A. DC. Monogr. Phan. 2:492. 1879; Engler & Krause in Engl., Pflanzenr. 4. 23E. 71:68 
1920—differs from the above varieties by the absence of an appendix. 


Colocasia esculenta (L.) Schott in Schott & Endl., Melet. Bot. 18.1832; Kunth, Enum. 3:57. 
841.Arum esculentum L., Sp. Pl. 1753. Caladium esculentum (L.) Vent., Mag. Enc. 4:471. 1801; 

Vent, Descr.Pl. Nouv. Hort.Cels t.30.1801;Vent.J.Bot.(Schrader) 4:319.1801.Colocasia antiquorum 

(L.) Schott in Schott & Endl., Melet. Bot. 18.1832; Kunth, Enum. 3:37. 1841; Hook. f.in Bot. Mag 

t. 7364, 1894; Engler & Krause in Pflanzenr., 4. 23E.65. 1920. 


This species is divided into numerous varieties and over 300 forms or land races prima- 


rily on the basis of vegetative differences (Engler & Krause 1920; Hill 1939; Neal 1991).The 


primary criteria for varietal separation are: size, shape, color, and number of tubers pro- 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 161 


duced, texture and chemical composition of tubers, degree of rhizome formation, and 
foliage morphology and coloration. Flower and fruit characters are also used. Varieties 
are used as ornamentals or as food crops for the edible, high carbohydrate tuberous 
rhizomes; some varieties are also invasive weeds. Spadix morphology and color can vary 
depending on variety. Flesh of tubers ranges in color from white, yellow, lilac-purple, or 
pink to reddish. Varieties of Colocasia esculenta are separated into two groups based on 
differential affinities for available soil or substrate moisture. Aquatic or wetland varieties 
prefer aquatic or semi-aquatic conditions of continual soil saturation, while upland forms 
prefer well-drained but continually moist soils. Both groups will survive under either set 
of conditions. Vegetative propagation occurs by division of the caudex, new secondary 
or daughter tubers, rhizome fragments, and buli (top one-third of tuber and 12-20 cm of 
stem; Neal 1991). Propagation by seed is also effective. 

In 1939, A.F. Hill published: The nomenclature of the taro and its varieties, showing 
Colocasia esculenta (L.) Schott, 1832 to be an earlier name than Colocasia antiquorum 
Schott, 1832. The species was originally described by Linnaeus as Arum esculentum, Sp. Pl. 
965.1753.Our varieties of Colocasia esculenta along with synonymy and specific varietal 
attributes are discussed below. 

Varieties aquatilis, antiquorum, and nymphaeifolia differ only slightly in their mor- 
phology. Variety antiquorum is most easily separated, differing by having petioles often 
nearly green in color, usually greater tuber development, and less developed rhizomes, 
whereas both aquatilis and nymphaeifolia have petioles usually with pronounced red- 
dish or purple coloration, often little to nearly no tuber development, and long, well- 
developed rhizomes. Varieties aquatilis and nymphaeifolia essentially differ from each 
other only in area of origination of rhizomes from the tuber or caudex (see key to spe- 
cies of Colocasia). These three varieties intergrade appreciably in form and ecology and 
probably encompass only a single, highly variable taxon. 


1, Colocasia esculenta (L.) Schott. var.aquatilis Hassk., Pl. Jav.Rar. 150. 1848. (Fig. 3). Cala- 
dium aquatile Rumph., Herb. Amboin. 5:313.t.110.f.1.1747. Colocasia antiq Schott var. 
aquatilis Hassk. in Engl., Pflanzenr. 4. 236. 71:68. 1 920. 


Colocasia anti hott var. stolonifera (Haines) H.B., Nathani, Flou. Pl.India, Nepal and Bhutan. 


Colocasia esculenta (L.) Schott var. stolonifera (Haines) H.B., Nathani, Flou. Pl.India, Nepal and Bhutan. 
454.1990 


Essentially acaulescent herbs to 1.5 m in height from a slightly tuberous caudex, tuber- 
ous Caudex usually poorly developed to nearly absent, well-developed rhizomes present 
to 2.5 min length, elongate, brittle, and covered with tan scale-like cataphylls, distal por- 
tion greenish-white and usually lacking the cataphyll covering; leaves 7-40 cm long and 
5-30.5 cm wide, ovate, ovate-triangular, or ovate-elliptic, apically rounded to acute, ba- 
sally cordate, cordate-sagittate, or sagittate, peltate, margins weakly undulate to entire; 
primarily lateral veins 3-8 per side, ranging in color from green to yellow, or purple, midvein 
and lateral veins even or minutely raised above the blade on adaxial side of leaf, blade 


SIDA 19(1) 


ANN 
< 
\ 


\\ q 
As 


Fic. 3. Colocasia esculenta (L.) Schott var. aquatilis Hassk. General habit. 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 163 


glaucous to glaucescent abaxially, petiole 40-180 cm in length, green to red or purple, 
often green and splotched with purple, sometimes pink basally; inflorescence of 1-nu- 
merous spadices, peduncles 15-50 cm long, spathe 10-40 cm long, chartaceous, broadly 
acuminate, yellow-green, yellow, yellow-orange, or orange; spadix 6-14 cm long, yellow- 
cream and often tinged with orange or pink, appendix 0.5-3 cm long, region of fertile 
staminate flowers 2-4 cm long, region of sterile staminate flowers 1.5-5 cm long, region 
of fertile pistillate flowers 1-2.5 cm long, pistillate flowers essentially without odor; ber- 
ries red, ellipsoid to oblong and 3-7 mm in diameter. 

Originally from southeast Asia, this variety along with varieties antiquorum and 
nymphaeifolia have become pantropical weeds of aquatic habitats, particularly rice fields, 
fresh water marshes, lake and waterway margins, and drainage sloughs. It is truly aquatic 
and cannot tolerate prolonged conditions of drought without the onset of dormancy. 
Extended drought over several months usually induces plant fatality. It was introduced 
into the United States as an ornamental and is completely naturalized as a member of 
our flora. Colocasia esculenta var. aquatilis has become an invasive weed in the south- 
eastern U S.It ranges from eastern Texas, to Louisiana, Mississippi,and Alabama, through- 
out Florida and north to Georgia. Variety aquatilis is the most established of the varieties 
of Colocasia esculenta in the United States. Though potentially invasive under any condi- 
tions with adequate moisture, the habitat preferences of variety aquatilis in the south- 
east US are continually wet or flooded roadside ditches; freshwater marshes; open areas 
of floodplains; fresh water beaches, and shallow water along the edges of ponds, lakes, 
streams, and slow moving waterways. Any environment with shallow standing water or 
continually wet soil is a potential habitat. Variety aquatilis requires little to no distur- 
bance for establishment.It has the potential to aggressively invade and quickly establish 
vegetative colonies regardless of the floristic component or degree of disturbance. Small 
colonies and isolated juvenile plants seem to indicate that spread is because of vegeta- 
tive propagule (rhizome fragments) fragmentation, translocation,and subsequent plant 
regeneration. This scenario is further supported because of the lack of fruit production 
in the US by Colocasia esculenta. 

Although roughly similar in form to Alocasia macrorrhizos and Xanthosoma 
sagittifolium, variety aquatilis is smaller and generally has more vigsieus rhizome produc- 
tion. These stoloniferous rhizomes are capable of producing | | 
(up to one per node). Typically only one or two plantlets are iodiveas at the rhizome 
terminus. Each plant can produce 2-10 or more rhizomes at any one time. Rhizomes are 
brittle and sever with minimal disturbance. Separation of rhizomes stimulates additional 
rhizome production.Rhizomes severed before leaves or roots have ged from the nodes 
can root and produce new plants on moist soil or in standing water. Extensive colonies of 
variety aquatilis occur throughout Florida, but are otherwise currently restricted to a zone 
approximately 100 miles inland from the Gulf of Mexico.Some isolated and localized popu- 
lations occur slightly farther inland and northward. Tuber development is usually reduced 
or almost absent, the subterranean portion usually only consisting of many fleshy fibrous 


g their length 


164 SIDA 19(1) 


roots and rhizomes. Large specimens sometimes have more pronounced tuber develop- 


ment at the crown region.Plants with larger more developed tubers (see sek esculenta 


e, the y re 
=) 


var.antiquorum) are not uncommon.Once rhizomes begin to elongat 
from the soiland become stoloniferous, rooting along the nodes and producing new plant- 
lets. At various lengths, the rhizomes sometimes repenetrate the given substrate. New colo- 
nies are nearly exclusively founded by translocation and subsequent establishment of nodal 
rhizome fragments or small plantlets. 

Variety aquatilis only occasionally produces flowers in the United States. Flowering usu- 
ally occurs in mid to late summer (late June through September or October), during which 
time conditions are dry throughout much of the southeastern US. If the site occupied be- 
comes lerately dry, flowers are often produced. Usually 1-3 inflorescences are produced. 
Plants will sometimes also flower in wet conditions. Fruit is not normally produced in the 
continental US and dispersal and colony establishment by seed is rare or absent. 

Colocasia esculenta var.aquatilis is propagated readily by plantlet division from rhi- 
zomes. Plantlets can be removed immediately after emergence from the rhizome so 
long as the node from which the plantlet originated is left attached, even leafy stem 
sections with a functioning apical meristem will usually root in water or wet soil. It can 
be grown with or without soil as long as standing water is provided, but establishment 
and growth is enhanced with some type of rooting medium. Plants thrive on excess 
nitrogen and phosphorous. Mature plants transplant readily. 

A variegated form of variety aquatilis with irregular lutescent areas of yellow to white 
on leaves and petioles, occurs infrequently within wild-type populations. 

Common names.—aquatic elephant-ear, elephant's-ear, wild taro. 


2: Colocasia eee (L.) Schott var.antiquorum (Schott) Hubb. & Rehd.in Bot. Mus. 
Leafl.1:1.5.1932. Colocasia Schott in Schott & Endl, Melet. Bot. 18.1832; Schott, 
Syn. Aroid, 41.1856; Schott, Prodr. Syst. Aroid. 138. 1860; Kunth, Enum PI. 3:37. 1841; Engl. in A. 
DC., Monogr. Phan. 2:491. 1879; Hook. fin Bot. Mag. 120. t. 7364. 1894; Engl. & Krause in Engl. 
Pflanzenr. 4. 23E. 71:65. 1920. 


Arum aegyptium Rumph., Herb. Amboin. 5:313.t. 109. 1747. 

Arum colocasia L., Sp. P|. 965. 1 ei 

Arum peltatum Lam., Encyc. 3:1 1786. 

Arum lividum Salisb., Prodr. ate ee Allerton 260. 1796. 
36. 


Colocasia peregrina Rafin. Fl. Tellur. 


Colocasia vulgaris Rafin., Fl Tellur. 3: a 1836. 
Caladium colocasia W.F.Wight in Contr. U.S. Natl. Herb. 9:208. 1905. 


Nearly identical in morphology and habit to varieties aquatilis and nymphaeifolia, variety 
antiquorum differs primarily on the basis of its nearly green petioles, larger tubers (espe- 
cially those at the crown) and usually shorter, less developed rhizomes, though these 
can approach the length and development of rhizome formation in varieties aquatilis 
and nymphaeifolia. 

Introduced for its edible tubers and as an ornamental from southeast Asia, Colocasia 
esculenta var. antiquorum is naturalized in the southeastern US but is less widespread 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 165 


than variety aquatilis. Variety antiquorum prefers margins of ponds and lakes, but will 
also rapidly colonize and invade lawns, flower beds, and other disturbed sites so long as 
adequate soil moisture is present.Establishment and complete invasion of various lawns 
and gardens was observed in Baton Rouge, Louisiana, in 1998 by both varieties antiquorum 
and nymphaeifolia. Plants prefer moist to saturated soil or shallow water and usually do 
not occur in water over 30 cm deep. 

The primary mode of reproduction is vegetative and occurs from secondary tu- 
bers, or plantlets produced from rhizomes. Variety antiquorum requires similar condi- 
tions for growth, colonization and expansion, and flowering as variety aquatilis. Flower 
morphology is similar to variety aquatilis, and as in variety aquatilis, fruit is not normally 
produced in the continental US. It seems to withstand somewhat drier conditions than 
variety aquatilis. lt occurs from east-central Texas, east to the Atlantic coast, and from 
Georgia southward throughout Florida. 

Common names.—Egyptian taro, culgas, golaas, black taro, eddoe, eddo, kalo, el- 
ephant-ear. 


3. Colocasia esculenta (L.) Schott var.esculenta Schott, Syn.Aroid.42.1856.Arum esculentum 
Sp. Pl.965. 1753. Caladium esculentum (L.) Vent., Descr. Pl. Nouv. Jard. Cels 30. t. 1800;Vent,, 
Mag. Enc. 4:471. 1801; Vent, J. Bot. (Schrader) 4:319. 1801. Colocasia antiquorum Schott var. 
esculenta Schott, Syn. Aroid.42. 1856; Schott, Prodr. Syst. Aroid. 140.1 860;Engl.in A.DC., Monogr. 

Phan. 2:492. 1879; Engl.& Krause in Engl., Pflanzenr. 4. 23E. 71:67. 1920. 


Arum maximum Bauhin in Verh. Batav. Gen. 4. 1790. 
Colocasia esculenta (L.) Schott var. typica Engl. Vers. Entwicklungsgesch. Extratrop. Florengeb. 8. 
1879. 


Variety esculenta is the most common ornamental form grown in the US and is charac- 
terized by its large tubers (up to 22 cm long and 18 cm in diameter), leaves, and size (to 
2m or more tall). Usually only a few tubers are produced per plant at any one time. 
Colocasia esculenta is the most important species of Araceae in terms of food produc- 
tion worldwide (Bailey 1997;Vaughan & Geissler 1997; Neal 1991).The tubers have a high 
starch content (approx. 25%), some protein,and 13 mg/100 g vitamin C.Small secondary 
tubers, also Known as cormels, are equally consumed (Vaughan & Geissler 1997). Sec- 
ondary tubers develop directly from the primary caudex or at the nodes or terminus of 
rhizomes. The foliage is also eaten and normally contains 7 mg/100 g carotene, and 52 
mg/100 g vitamin C; all parts of the plant are used as food in certain regions of Africa 
and Asia. Removal of calcium oxylate from the tubers is accomplished through boiling 
(Vaughan & Geissler 1997; Neal 1991). Chemical composition of the tubers produced by 
Colocasia is similar to those of Alocasia and Xanthosoma (Vaughan & Geissler 1997). 
Colocasia is grown as a food source in the United States and Mexico, throughout the 
neotropics, Africa, and Asia. It is a staple food crop in many areas of Africa where it is 
known as cocoyam (Vaughan & Geissler 1997; Neal 1991). The amount of tuber produc- 
tion varies with varietal type, soil texture, environmental conditions, and health of the 
plant (Young 1946). 


166 SIDA 19(1) 


Common names.—taro, elephant-ear, elephant-plant. 


4.Colocasia esculenta (.) Schott var.euchlora (C. Koch and Sello) A.F.Hill, Bot. Mus. Leafl. 

7:7.1939.Colocasia euchlora C.Koch and Sello in Ind. Sem. Hort. Bot.Berol.App.4. 1854.Colocasia 

antiquorum Schott var.euchlora Schott, Syn. Aroid.42. 1856; Schott, Prodr. Syst. Aroid. 140. 1860; 

Engl.in A.DC. Monogr. Phan. 2:491.1879; Engl. & Krause in Engl. Pflanzenr. 4. 23E. 71:67.1920. 
Plant to 92 cm tall, leaves with lamina bright green, margins usually purple, petioles and 
veins red to purple. Rhizomes similar to those of varieties aquatilis or nymphaeifolia but 
red or purple in color and usually shorter. Similar to variety fontanesii in coloration but 
smaller. Vegetative reproduction by rhizomes and secondary tubers. Grown in extreme 
southern US but no current record of establishment outside cultivation. 

Common name.—purple-stemmed taro. 


5. aaa esculenta (L.) Schott var. fontanesii (Schott) A.F.Hill, Bot. Mus Leaf 7:7.1939. 
asia fontanesii Schott in Oestr. Bot Wochenbl.4. 409. 1854. Colocasia Schott 
var. a Schott, Syn. Aroid. 42. 1856; Schott, Prodr. Syst. Aroid. 140. 1860; Engl in A.DC,, 
Monogr.Phan.2:491.1879;Hook.f.in Bot.Mag. 126.t.7732. 1900; Engl.& Krause in Engl. Pflanzenr. 

4, 23E. 71:67. 1920. 


Arum colocasioides Desf., Cat. Hort. Par. ed. 3. 7. 385.1829. 
. ladium violaceum Hort. ex Engl. in DC. Monogr. Phan. 2:492. 1879; Desf., Cat. Hort. Par. ed. 3.7. 
829. 


dm colocasioides Hort. Par.ex Brongn. in Nouv. Ann. Mus. Par. 3. 156. 1834; Kunth, Enum. Pl. 
ae 3.1841. 


Colocasia violacea Hort. ex Hook. f.in Bot. Mag. 126.t. 7732. 1900. 


Nearly identical in form to variety euchlora, but larger—to 2.14 m tall, and more robust 
with the petioles and veins of leaf usually black or dark purple;lamina dark green, often 
tinged with purple coloration. Rhizomes similar to varieties aquatilis and nymphaeifolia, 
but purple to purple-black and thicker. A form with completely purple to purple-black 
foliage is also cultivated in the US. Lamina, petioles, peduncles, and spathe all purple, 
undersurface of leaf glaucous-purple, roots often with pink or purple coloration. Leaves 
on plants grown under shade conditions and newly emergent leaves are blue-green 
with purple coloration along veins and petioles. Newly emergent leaves on juvenile plants 
sometimes green. This form actually appears to be an intervarietal cross between variet- 
ies fontanesii and illustris—being phenotypically intermediate between the two. Both 
forms are vegetatively aggressive similar to varieties aquatilis and nymphaeifolia. Culti- 
vated in extreme southern US and observed spreading vegetatively at these sites but 
no record of establishment outside cultivation. It is another potentially weedy variety. 
Common Names.—black-stemmed taro, black taro, purple taro, black magic taro. 


6. Colocasia esculenta (L.) Schott var. Saat uel & Krause) Young in U.S. Dept. 
Agric. Dept. Bull. 8:1247. 1924. Col im Schott var.globulifera Engl. & Krause 
in Engl. Pflanzenr. 4. 23E. 71:68. 1920. 


This upland variety is grown commercially in the US for its small edible tubers. Individual 
plants normally produce 20 or more 2-10 cm long secondary tubers terminating short 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 167 


rhizomes or originating from the primary caudex. Plants to 95 cm tall. Variety globulifera 
is not well-established in the US but does occur and persist near areas where it is in 
production. 

Common name.—dasheen. 


7.Colocasia esculenta (L.) Schott var illustris (Bull.) A.F Hill, Bot. Mus. Leafl. 7:7. 1939.Alocasia 

illustris Bull, Cat.4. 1873; Bull.in Flor. Mag. t. 107.1874. Colocasia antiquorum Schott var. illustris 

Engl.in A. DC. Monogr. Phan. 2:492. 1879; Engl. & Krause in Engl. Pflanzenr. 4.23E. 71:67. 1920. 
Plants to 1m tall and resembling varieties aquatilis and nymphaeifolia in form but less 
aggressive and usually with shorter, more tuberous rhizomes. Newly formed rhizomes 
are more slender than those of varieties aquatilis or nymphaeifolia. Leaves with lamina 
green and spotted to completely purple between primary veins, primary veins green, 
lamina glaucous abaxially, petioles cle with red or red-brown coloration or completely 
red.Variety i/lustris is cultivated in the extreme southern US. It is not currently naturalized 
but spreads locally where cultivated and could potentially be invasive under favorable 
conditions. 

Common names.—|mperial taro, black caladium. 


8. Colocasia sp pete .) Schott var.nymphaeifolia (Vent.) A.F. Hill, Bot. Mus. Leafl. 7:7. 
1939. (Fig. 4). Caladium nymphaeifolium Vent., Descr. Pl. Nouv. Jard. Cels sub t. 30. 1800;Willd., 
Sp.PI.4. ee 1805.Arum nymphaeifolium Roxb., Hort. Beng, 65. 1814; ROMP: Fl.Ind. aides rate 
Colocasia nymphaeifolia Kunth, Enum.PI.3:87.1840.Col 
Schott, Syn. Aroid. 42. 1856; Schott, Prodr. Syst. Aroid. 140. 1860: Engl.in A.DC., Monoar. Phan. 
2:492. 1879; Engl. & Krause in Engl, Pflanzenr. 4. 23E. 71:67.1920. 


Practically identical in form and habit to variety aquatilis (see varietal discussion follow- 

ing Colocasia esculenta) but differs by producing rhizomes exclusively from the base of 

tubers. Similar range and ecology as var. aquatilis but seemingly less common. 
Common names.—aquatic elephant-ear, taro. 


9. Colocasia gigantea (Blume) Hook.f, Fl. Britt. Ind. 6:524. 1893, 


Caulescent, erect, terrestrial or epilithic herbs to 2 m or more in height from a tuberous 
caudex. Leaves 20-120 cm long and 15-90 cm wide, ovate, apically rounded to acute, 
basally cordate, cordate-sagittate, or sagittate, peltate, apices short-acuminate, margins 
undulate; primarily lateral veins 4-8 per side, lamina green, petiole 20-150 cm in length, 
light green, white-pruinose; inflorescence of 1-numerous spadices, peduncles 20-55 cm 
long, spathe 25-50 cm long, chartaceous, short-acuminate, white; spadix 8-23 cm long, 
yellowish, appendix 3-6 mm long, region of fertile staminate flowers 2-9.5 cm long, re- 
gion of sterile staminate flowers 2—4.5 cm long, region of fertile pistillate flowers 4-8 cm 
long; berries oblong to 1 cm in diameter. 

Colocasia gigantea is similar to Colocasia esculenta and Alocasia macrorrhizos, and is 
included here to alleviate any confusion in identification of plants encountered. This 
species is rarely cultivated and not currently established outside cultivation in this coun- 
try. Native to Java. 


Fic. 4. Col 


r) 


Ley i 
i) IN 
! (ye 


GR WES 
(a 


(AS 


ta (L.) Schott var. ) : 


) A.F. Hill. General habit 


SIDA 19(1) 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 169 


Common names.—giant taro, elephant-ear. 


XANTHOSOMA 


Xanthosoma Schott in Scone Endl., Melet. Bot. 19. 1832; Engler Pflanzenr. 4. 23E. 71:41. 
1920. Lectotype species: Xanthosoma sagittifolium (L.) Schott, Arum sagittifolium L. 


Acontias Schott, Melet. Bot. 19.1 
Phyllotaenium André in Illust. Fiortie’ 19:3.t.88. 1872 


Perennial caulescent to acaulescent herbs, with or without a tuberous caudex. Tubers 
and or rhizomes typically present, asexual reproduction by means of secondary or daugh- 
ter tubers and or rhizomes. Above ground stems erect to decumbent; leaves ovate, trian- 
gular-ovate to ovate-elliptic, long petioled, basally hastate, three to several lobed or 
pedatisect, leaves and stems glaucous or not; spadix pedunculate, arising axillary and 
subtended by 1 or 2 acute to acuminate bracts, staminate and pistillate flowers sepa- 
rated by a section of synandrodes (sterile staminate flowers), sterile appendix reduced 
or absent; spathe present, 1-3 times longer than spadix, constricted basally to form a tube 
which encloses the spadix prior to anthesis, the basal portion remaining tightly closed; 
staminate flowers comprised of synandria of 4-6 stamens; pistillate flowers 1-carpellate, 
2-4-locular, stigma discoid, ovules numerous, placentation pseudoaxile or parietal; fruit 
a cylindrical, furrowed berry; ovules usually 12-20, but can be numerous, anatropous or 
hemianatropous, funicles long; seed ovoid, testa costate, endosperm copious. 

About 58 species originating in the neotropics. Currently only X. sagittifolium is natu- 
ralized in the United States. 

Etymology.—Greek:xantho, the color yellow and soma, a body; yellow body—prob- 
ably in reference to the yellow color of various stem tissues. 


KEY TO SPECIES OF XANTHOSOMA 


. Leaves entirely green to green-blue, young leaves green to green-blue; petioles 
1 


green to glaucous .X. sagittifolium 
1. Leaves green but margins and veins usually purple, young leaves blue-purple to 
primrose; petioles often purple but can be yellow-green 2.X.violaceum 


1. Xanthosoma sagittifolium (L.) Schott in Schott & Endl., Melet. Bot. 19. 1832; Engler, 
Pflanzenr. 4. 23E. 71:45. f. 9A. 1920. (Fig. 5). Arum sagittifolium L., Sp. Pl. 966. 1753. Arum 
sagittaefolium L., 1.c., ed. 2, 1369. 1763. Caladium sagittifolium (L.) Vent., Mag. Enc. 4(16):471. 
1801;Vent., Arch. Bot. (Leipzig) 2(3):351.1801;Vent., J. Bot. (Schrader) 4(6):319. 1801. Caladium 
sagittaefolium (L.) Willd. Sp.PI.4:489. 1805; Spreng. Syst.Veg.3:771.1826.Xanthosoma sagittaefolium 
(L.) Schott in Schott & Endl., Melet. Bot. 19. 1832. 


Nery 


Arum acaule Plum., Pl. Amer. t. 35.1756. 

Arum triangularis Plum.,Pl.Amer. t. 35.1756. 

Arum xanthorrhizon Jacq,, Pl. Hort. Schoenbr. 2:32. 1797. sea een Jacq.) Willd., Sp. 
Pl. 4:490. 1805. Xanthosoma xanthorrhizon Jacq.) C. Koch, Bonplan 1856. 

Caladium edule G.FW. Mey, Fl. Esseq.272. 1818. Xanthosoma en Be Schott in Schott & 
Engl., Melet. Bot. 19. 1832 


SIDA 19(1) 


Os? oa Fy 


(L.) Schott. General habit 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 171 


Xanthosoma atrovirens C.Koch and Bouché, Ind. Sem. Hort. Berol. 3. 1854. 
Xanthosoma utile C.Koch & Bouché, Ind. Sem. Hort. Berol. 2. 1854. 
Xanthosoma zanthorrhizon C. Koch, Bonplandia 4:4. 1856. 

Xanthosoma wendlandii Schott in Oestr. Bot. Zeitschr. 15:3. 1865. 
Caladium utile Hort. ex Engl.in A. DC. Monogr. Phan. 2:469. 1879. 


Caulescent herbs to 3.5 m in height from a tuberous caudex, tubers and tuberous cau- 
dex poorly developed or absent in juvenile plants; well-developed rhizomes present to 
1 mor more in length, elongate, slender, without scaly covering of cataphylls, often tu- 
berous at various places along the length, white and often tinged with light green; leaves 
20-120 cm long and 15-90 cm wide, ovate to triangular-ovate, apically acute to acumi- 
nate, basally hastate-each lobe distally acute, margins weakly but broadly undulate or 
entire, abaxial surface usually glaucous;primary lateral veins 3-9 per side, whitish or light 
green, lighter than the surrounding blade, midvein and lateral veins sunken below the 
blade on adaxial side of leaf; petioles green to glaucous; inflorescence of 1-numerous 
spadices, peduncles 20-30 cm long, glaucous or glaucescent; spathe 18-22 cm long 
and 4 cm wide, chartaceous, pale green to glaucescent, turning cream to peach with 
age; spadix 6-13 cm long, green to cream or yellowish, appendix 3-4 cm long, fertile 
staminate region 5-7 cm long, sterile staminate region 3-4 cm long, fertile pistillate re- 
gion 3.5-5 cm long, pistillate flowers with slight odor of ether; berries ovoid, yellow, 7-10 
mm long and 4-8 mm in diameter. 

Native to the neotropics and cultivated in Central and South America and the West 
Indies since pre-Columbian times (Vaughan & Geissler 1997).It was introduced into the 
United States as an ornamental for its large size, leaves, attractive flowers, and fruit (Bailey 
1997;Vaughan & Geissler 1997).Xanthosoma sagittifolium is naturalized throughout cen- 
tral and southern Florida and in southern Texas. It is highly rhizomatous and can be- 
come an invasive weed in areas of disturbance and adequate moisture. This species has 
the potential to increase as a weed under favorable conditions. 

Plants generally produce large, somewhat elongate, oblong, subterranean tubers 
along the length or at the end of slender rhizomes or from the base of the crown. Juve- 
nile plants have little or no tuber development; instead they possess a network of fleshy 
fibrous roots interspersed with elongate slender rhizomes. Large colonies are formed 
through vegetative propagation by rhizomes. Small to medium sized colonies of imma- 
ture plants at the rhizome stage are often observed. Such plants likely arise through 
vegetative fragmentation and subsequent relocation of the rhizome fragments. Dispersal 
and establishment from seed is also possible but less likely. Although fruit production is 
not often observed, plants are self compatible, with a single individual or colony produc- 
ing fruit and viable seed. It can be difficult to determine whether new populations origi- 
nated from seedlings or from vegetative plant propagules. Vegetative expansion seems 
a more plausible explanation. This species is probably not distinct from the similar x. 
atrovirens C.Koch and Bouché. 

Xanthosoma sagittifolium prefers moderate to high soil disturbance and soil water 
content. It is generally found in any disturbed area where water collects; along roadside 


172 SIDA 19(1) 


ditches, pond and lake margins, drainage areas, freshwater wetlands, marshy woods, sea- 
sonably wet sites, woodland edges, old fields, vacant city lots, roadsides, and home sites. 
Although X. sagittifolium prefers soil conditions that are moderately hydrated to highly 
saturated, it will grow on mesic slopes and other habitats with well-drained soils, as long 
as adequate moisture is present. It was observed in both wet and mesic sites in Ocala 
and Bellview, Florida, in 1996 and 1997. Xanthosoma sagittifolium has minimal cold toler- 
ance.|t withstands occasional freezing temperatures, but cannot sustain prolonged freez- 
ing. Juvenile, unestablished plants are most susceptible to freezing because older plants 
can regenerate from tubers. The tubers are used as a food source throughout the Ameri- 
can tropics and to a lesser extent in the South Pacific and Africa (Bailey 1997;Vaughan & 
Geissler 1997; Neal 1991). The tubers are similar in chemical composition to those of 
Colocasia or Alocasia, but the starch is more difficult to digest (Vaughan & Geissler 1997). 
In West Africa, the tubers are preferred over those of Colocasia for the production of the 
food substance known as fufu (Vaughan & Geissler 1997). 

Xanthosoma sagittifolium is easily cultivated. Vegetative propagation is accomplished 
through tuber division, rhizome tip development and fragmentation, plantlet removal, 
and crown division. Small plantlets can be removed from the parent plant at the 2-3 leaf 
stage so long as the entire stem is intact. Unlike Colocasia, rhizomes of Xanthosoma are 
primarily subterranean. Even mature individuals transplant readily. 

Common names.—arrowleaf-elephant-ear, elephant-ear, yautia, malanga, tannia, 
tanier, tanyah, tajer, ocumo, cocoyam. 


2. Xanthosoma violaceum Schott, Ind. Sem. Hort. Berol. 370. 1853; Schott in Oest. Bot. 
Wochenbl. 417.1854 


Arum nigrum Vell. Fl.Flumin.9.t.107. 1829. Xanthosoma nigrum (Vell.) Mansf, Verz. Pfl. Deut. Reich. 
1940. 


Arum variolatum Schott, Prod. Syst. Aroid. 81.1860. 

Xanthosoma ianthium C. Koch & Bouché, Ind.Sem Hort. Berol. 2. 1854. 
Caulescent herbs 30-150 cm in height, from a tuberous caudex, stems purple or yellow- 
green, tubers and tuberous caudex poorly developed or absent in juvenile plants; well- 
developed rhizomes present to 1 m or more in length, elongate, slender, without scaly 
covering of cataphylls, often tuberous at various places along the length, white and of- 
ten tinged with light green or pink coloration; leaves 30-100 cm long and 15-75 cm 
wide, ovate to triangular-ovate or ovate-oblong, apically acute to acuminate, basally sag- 
ittate to hastate—each lobe distally obtuse to acute and unequal, margins weakly but 
broadly undulate or entire, abaxial surface usually glaucous; primary lateral veins 4—9 per 
side, purple or rarely a pale yellow-green, midvein and lateral veins sunken below the 
blade on adaxial side of leaf, petioles usually purple, sometimes yellow-green, older leaves 
dark green, young leaves blue-purple or yellow-green with purple or yellowish primary 
or secondary veins; inflorescence of 1-numerous spadices, peduncles 30-50 cm long, 
glaucous or glaucescent; spathe 7-10 cm long and 3.5-4 cm wide, chartaceous, con- 
stricted portion purple to violet, green to yellow-green and violet-striate; spadix 15-25 


SERVISS ET AL., ARACEAE IN THE SOUTHEASTERN US 173 


cm long, initially purple but with age becoming grey or yellowish, appendix absent, fer- 
tile staminate region 10-15 cm long, sterile staminate region 3-5 cm long, fertile pistil- 
late region 3-5 cm long, pistillate flowers with slight odor of ether; berries ovoid, yellow, 
7-10 mm long and 4-8 mm in diameter. 

Cultivated in southern Florida but not strongly naturalized. Plants have been ob- 
served spreading vegetatively at sites of cultivation, and eventual establishment outside 
these conditions is probable. Xanthosoma violaceum is cultivated throughout the trop- 
ics for its edible tubers similar to X. sagittifolium. The exact area of origin (possibly the 
West Indies) for X. violaceum is uncertain, but it originated in the neotropics. Ecology 
similar to that of X. sagittifolium. 

Common names.—blue tannia, blue taro, blue ape, tanier, tajer. 


ACKNOWLEDGMENTS 


| would like to sincerely thank Thomas B. Croat at the Missouri Botanical Garden for his 
critical review and insightful suggestions on the content of this work. A high degree of 
gratitude is also in order for John R.MacDonald at Mississippi State University in Starkville, 
MS and Richard J. Carter at Valdosta State University in Valdosta, GA, for their editorial 
contributions and advisement in the preparation of this manuscript.| am also very thank- 
ful to Tricia K. Blake, Terri Ballinger, and Keri Denley. 


REFERENCES 
Baitey, L.H.,E.Z. Baitey, and Hortorium Starr. 1997.Hortus third. Cornell University. Barnes and 
Noble, New York, NY. 
Croat, T.B. 1998. Tropical aroids:T: ,diversity,and ecology In:Mathew and Sivadasan 
eds. Diversity and taxonomy of tropical flowering plants. Mentor Books, Calicut. Pp. 


. 1988 (1990). The ecology and life forms of Araceae. Aroideana 11(3-4):4—56. 

Enter, A. and K. Krause. 1920. Araceae In: A. Engler: Das Pflanzenreich, 4:23, A-F, Leipzig 
1905-1920. 

French,J.C.,,M.G.Chung, and Y.K. Hur. 1995. Chloroplast DNA phylogeny of the Ariflorae. In: 
P.J. Rudall, PJ. Cribb, D.F. Cutler, and C.J. Humphries, eds. Monocotyledons: Systematics 
and evolution. Royal Botanical Gardens, Kew. Pp. 277-286. 

Grayum, M.H. 1984. Palynology and phylogeny of the Araceae. Ph.D. dissertation. Univ. of 
Massachusetts, Amherst, MS. 

Hit, A.F. 1939. The nomenclature of the taro and its varieties. Bot. Mus. Leafl. 7(7). Cam- 
bridge. 

Mayo, S.J., J. Boaner, and P.C. Boyce. 1997. Genera of Araceae. Royal Botanical Gardens, Kew. 

1995. The Arales. In: PJ. Rudall, PJ. Cribb, D.F. Cutler, and C.J. Humphries, eds. 
Monocotyledons: Systematics and evolution. Royal Botanical Gardens, Kew. Pp. 277- 


Neat, M.C. 1991. In gardens of Hawaii. Bernice Pauahi Bishop Museum, Honolulu, HI. 


174 SIDA 19(1) 


VAUGHAN, J.G.and C.A. Geisster. 1997. The new Oxford book of food plants. Oxford University 
Press, Inc., New York, NY. 

Youne, R.A. 1946. The dasheen, a southern root crop for home use and market. USDA 
Farmers’ Bull. 1396, Rev. ed. 


GEOGRAPHIC DISTRIBUTION PATTERNS OF THE GENUS 
ARIOCARPUS (CACTACEAE) IN TAMAULIPAS, MEXICO 


José Guadalupe Martinez-Avalos Humberto Suzan- puelin 
loactityite do EF, / gia y Alimentos Ecrrjola do Rin}, g 
tInjivercidad Art ad Tamaulipas Universidad Aut6noma de Querétaro 
13 Blvd. Lopez Mateos 928 Cerro de las Campanas s/n Querétaro 
Cd. Victoria, Tamaulipas, MEXICO 87040 Querétaro, MEXICO 76010 
ABSTRACT 

The geographic distribution of the genus Ariocarpus in the State of Tamaulipas, Mexico was deter- 
mined based on literature, herbarium vouchers bamniel gardens and eens asia ups in 1991 
and 1992. Three speci bspecies were detected 
A. retusus subsp. retusus and A retusus subsp. Mee Ariocarpus agavoides is es restricted 
eee an area less than 2 km?; the main factor responsible for its distribution is 


oil type, which has clay texture, low organic matter content and a slightly alkaline pH. Ariocarpus 
retusus subsp. trigonus has the largest distribution (all the arid and semiarid portions of the state) 
probably because of its tolerance to different ecological factors, such as vegetation, soil types, and 
its ability to survive even if nurse plants were removed. 


KEY Ari , Geographic distribution. 


v 


RESUMEN 


Se determin6 la distribucién geografica del género Ariocarpus en el estado de Tamaulipas, México 
base en revisiones bibliograficas, consultas de herbarios, jardines botanicos, asf como en 
recorridos de campo realizados en 1991 y 1992. Se encontraron tres especies y dos subespecies: 
Ariocarpus agavoides, A. kotschoubeyanus, A. retusus subsp. retusus y A. retusus subsp. trigonus. 
Ariocarpus agavoides se encuentra euieinaciaieiie restringida geograficamente, ocupando un 
area no mayor a 2 km’;el factor de su distribucion es el tipo de suelo, el cual se caracteriza 
por su textura arcillosa, baj ntenido de materia organica, asi como un pH ligeramente alcalino. 
Ariocarpus retusus subsp. trigonus presenta el mayor rango de distribucién (ocupando las regiones 
dridas y semidridas del estado) debido a su tolerancia a diferentes factores ecoldgicos a como 
vegetacion, tipo de suelo, y a su sobrevivencia cuando las plantas nodrizas son removi 


INTRODUCTION 


The genus Ariocarpus described by Scheidweiler in 1838 ( Mitich & Bruhn 1977), is widely 
distributed in the Chihuahuan Desert (Anderson 1960).This desert is located in the Mexi- 
can highlands that includes portions of the states of Texas (U.S.A.) and Coahuila, Nuevo 
Leon, San Luis Potosi, Hidalgo, Querétaro, Zacatecas, and Tamaulipas in México (Shreve 
1951; Jaeger 1957; Rzedowski 1978; Bravo 1978).In Tamaulipas this region is known as 
the Tamaulipan Arid Zone (Bravo & Sanchez-Mejorada 1992) including the municipali- 
ties (municipios) of Tula, Palmillas, Miquinuana, Bustamante and Jaumave (Fig. 1). 

The genus Ariocarpus includes seven species: Ariocarpus agavoides (Castaneda) E.F. 
Anderson, A. bravoanus H. Hernandez & E.F. Anderson, A. kotschoubeyanus (Lemaire) K. 


SIDA 19(1): 175-183. 2000 


UNITE STATES OF AMERICA 


S ( 
‘ ee gy 
a, l < M : 
i A f a 
oe: 


24°30’ 


Golf 


Mexico 


SYMBOLS 


A. agavoides 

A. kotschoubeyanus 

A. retusus subsp. retusus 
A. retusus subsp. trigonus 


D0ece 


22° 40’ 


100°12' 


NUVO LEON 


Miquihuana 


SAN LUIS POTOSI 


arlos 
Oo 


Villa de Casas 


SLL 


98°40’ 


Fic 1 AA £4h a+ £Tamaulinas tth tha dictrihiiti £ Anz. 
oT. Map Or tit t 


(L)6L vals 


MARTINEZ AND SUZAN, DISTRIBUTION OF ARIOCARPUS IN TAMAULIPAS, MEXICO 177 


Shumann, A. fissuratus (Engelmann) K. Schumann, A. retusus Scheidweiler, Ariocarpus 
trigonus Weber, and A. scaphirostris Boedeker (Hunt 1992; Hernandez & Godinez 1994). 
Nevertheless in a recent revision of the genus Anderson and Fitz Maurice (1998) added 
two new subspecies, A. bravoanus H. Hernandez & E.F. Anderson subsp. hintonii E.F. Ander- 
son & Fitz Maurice,and A. retusus K.Schumann subsp.trigonus (Weber) Scheidweiler based 
on morphological traits,and hybridizing populations. All taxa are considered threatened 
or endangered (Anonymous 1991; CITES 1992; UICN 1985; Vovides 1988). The biggest 
threat to the survival of Ariocarpus species is habitat fragmentation due to the removal 
of natural vegetation in large areas, overgrazing, highway construction, development of 
urban areas and over-collection of wild populations by commercial and amateur collec- 
tors for sale as ornamental plants (Sanchez-Mejorada 1982, 1987; Hernadndez & Godinez 
1994) 

The literature published on the genus includes the works by Anderson (1958, 1960, 
1961, 1962, 1963, 1964): Bravo (1978), Bravo and Sanchez-Mejorada (1992) and Sanchez- 
Mejorada (1987). 

Only one species of the genus have been studied with an ecological perspective 
(Ariocarpus trigonus) by Suzan et al. (1989) and Martinez et al. (1993). 

In Tamaulipas little is known about the distribution and viability of these species 
because no floristic inventories and demographic studies necessaries for the definition 
of the real status of each species exists to date (Vazquez-Yanes 1979). The main objec- 
tive of the present study is to document the geographic distribution of the genus 
Ariocarpus in the state of Tamaulipas, México. 


METHODS AND MATERIALS 


The state of Tamaulipas is located at the northeastern portion of México, bordered by 
U.S.A. to the north, the States of San Luis Potosi and Veracruz to the south, Gulf of Mexico 
to the east, and State of Nuevo Ledn to the west. Three major climatic regions according 
to the Képpen classification modified by Garcia (1964) exist in the state: 1) the central- 
north region with semi-arid, semi-hot climates, with little annual rainfall (BS1 hw);2 ) the 
southeast region with hot sub-humid or humid climates with summer rainfall (Awo); 3) 
the south-west region located in the Sierra Madre Oriental with climates ranging from 
semi-hot sub-humid climates (A) C (W1) (W) to semi-hot humid climates (A) C (m) (w), in 
an altitude gradient on oe eastern slope, and the semi-hot subhumid (A)C(m)(w) to dry 
semi-hot Bsohw,in an alt gradient on the western slope (Secretaria de Programacion 
y Presupuesto (SPP) 1985). 

The vegetation is dominated by the thorn-forest (north central region), the xero- 
phytic shrubland (south-west region), and tropical deciduous forests (southeast region). 
In the highlands of the Sierra Madre Oriental, Pine-Oak and Cloud forests are the domi- 
nant vegetation types (Table 1) (Marroquin et al. 1964; Rzedowski 1978). 

Data w athered from voucher specimens at the following herbaria: DS, GH, IBUG, 
K, MEXU, MO, NY POM- RSA UAT, US (Holmgren et al. 1981). 


Se 


Tas.e 1.Climatic characteristics of the different municipalities in Tamaulipas where Ariocarpus species were detected. Data from Secretaria de Programacion 
y Presupuesto (1985). 


Species Municipality Vegetation Soils Climate Altitude Temperature Precipitation 
(m) °c) (mm) 
A.agavoides Tula Ms Xk BSohw 900-1250 21 436 
A. kotschoubeyanus Tula Ms Xk BSohw 1120-1200 2] 436 
A. retusus Tula Mr, Xk | Bsohw, BSIhw 1100-1300 21 434 
subsp. retusus Bustamente Mr,Ch |, Xk Cx, BSIkx 1700-1900 17 464 
Miquihuana Mr, Ch I, Xk Bsohx, BSIkx 1600-1900 17 464 
A. retusus Jaumave Mr, MeT Bk, |, Xk BSihw, Bsohw 600-900 23 469 
subsp. trigonus Palmillas MeT Bk, | (A)C(wo) 1100-1300 19 606 
. sas Sbc Bk BSl(h)hw, Awo 200-304 24 749 
(San Francisco) 

San Carlos MeT,M Bk, Zg (A)C(wo) 150-190 24 809 
Llera Sbc ZQ Awo 350--450 25 789 


Ms= submontane shrubland, MeT = tamaulipan shrubland; Mr = century plants shrubland; Ch = = chaparral; Sbc = tropical dry forest; M = mesquite forest; Xk 
= calcic alcic cambisol;| = Litosoil; Zg = Solonchak gleyic; Bsohw = semidry semi-warm;BS1hw = temperate semidry; (A)C(wo) = semi-warm 


sub- Mire with winter rains; BS1(h)hw = semidry warm; Awo = warm sub-humid with winter rains. 


BLL 


(L)6L Vals 


MARTINEZ AND SUZAN, DISTRIBUTION OF ARIOCARPUS IN TAMAULIPAS, MEXICO 179 


Living specimens of Ariocarpus were studied in the botanical gardens: Jardin 
Botanico de la Universidad Nacional Autonoma de México (UNAM): Coleccién de 
Cactaceas del Instituto de Ecologia y Alimentos (UAT); Jardin Botanico del Instituto 
Tecnoldégico de Ciudad Victoria, Tamaulipas (ITCV). Field work was carried out through 
the state, in order to visit reported sites of Ariocarpus occurrence. 


RESULTS 


In Tamaulipas only three species and two subspecies of the genus were found:Ariocarpus 
agavoides, A. kotschoubeyanus,A. retusus subsp. reutusus and A. retusus subsp. trigonus. Dis- 
tribution of the species is expressed in Fig. 1. The geographic distribution and character- 
istics of each species are: 


1. Ariocarpus agavoides (Castaneda) E.F. Anderson 
Common name.—"“Magueyito 
Geographic distribution.—San Luis Potosi and Tamaulipas. This species is endemic 

to the valley of Tula, municipality of Tula, Tamaulipas. The first population reported was 

located in the north slopes of the city of Tula. However, a new locality was discovered 
within the valley recently. Ariocarpus agavoides inhabit small hills with medium slopes 
and easily erodable rocky soils (xerosols), in altitudes from 900 m to 1200 m.The associ- 
ated vegetation in the population located north of Tula were dominated by Prosopis 
glandulosa, Koeberlinia spinosa and Acacia farnesiana, whereas for the new locality the 
dominant species were Hechtia glomerata, Agave lecheguilla and Agave striata. 

Field observations indicated strong dependence to nurse plants, low recruitment 
and pollination dependent on bees. The type locality is extremely perturbed by a waste 
disposal and overgrazing. Signs of plant extraction were detected several times. 


Cited specimens: MEXICO. Tamaulipas: Tula, 1200 m, Castafeda s.n.(DS);Tula, 1200 m, Cowper 1957 
(POM); Tula, 1200 m,Anderson 1616, (GH, K, MO, NY, POM). 

Specimens examined: MEXICO. Tamaulipas: Tula, 1200 m, Bravo s.n. (MEXU); Tula, 1170 m, 
Herndndez et al. 2054 (MEXU); Tula, 1240 m, Scheinvar & Sdnchez-Mejorada 750 (MEXU);Tula, 1200 m, 
Sdnchez-Mejorada 750 Jardin UNAM); Tula 1200 m, Herndndez Jardin ITCV); Tula, 1200 m, Martinez- 
Avalos & Jiménez 0441 (UAT); Tula, 914 m, Martinez-Avalos 0446 (UAT). 


2. Ariocarpus kotschoubeyanus (Lemaire) K. Schumann 

Common names.—"Pezuna de venado,”“Pata de venado" 

Geographic distribution.—Tamaulipas, Nuevo Leén, Zacatecas, San Luis Potosi and 
Querétaro. This species is reported for Tamaulipas in two sites in Tula, located northeast 
of Tula City (1200 m).The species grows in rocky soils, with poor organic material (xerosols), 
and inhabit medium slopes in a desert shrubland (Rzedowski 1978) dominated by Agave 
lecheguilla, A. striata, Hechtia glomerata and Dasylirion longissimum. 

We detected a well-preserved locality (location concealed for conservation pur- 
poses), with populations represented in different size classes. Sites close to Tula City ex- 
hibited heavy habitat disturbance by overgrazing and over-collection of plants. 


180 SIDA 19(1) 


Cited specimens (Anderson 1960, 1963): MEXICO. Tamaulipas: Tula 1200 m, Albert 1959 (POM. US); 
Tula, Anderson 161 alee US) 

Exa specimens: MEXICO. Tamaulipas: Tula, 1190 m, Anderson 1738 (MO); Tula, Arreola 
736 (IBUG); an Scheinvar et al. 4070 (MEXU); Tula, 1200 m, Martinez-Avalos 091 (UAT); Tula, 1127 m, 
Martinez-Avalos 0437 (UAT). 


3. Ariocarpus retusus K. Schumann subsp. retusus (Scheidweiler) E.F Anderson & Fitz 
Maurice 

Common names.—"Chaute,”"Chautle” 

Geographic distribution.—Tamaulipas, Coahuila, Nuevo Leon, San Luis Potosi y 
Zacatecas. In Tamaulipas it is common in the municipalities of Miquihuana, Bustamante, 
Tula and Palmillas. This species grows over hills and rocky slopes with clustered popula- 
tions at altitudes from 700 to 2100 m, in a desert shrubland (Rzedowski 1978), domi- 
nated by Agave lecheguilla, A. striata, Dasylirion longissimum, D. texanum and Yucca 
carnerosana. 

Populations of A. retusus subsp. retusus inhabit also perturbed areas dominated by 
Dodonea viscosa, in small canyons. Hybridizing specimens with A. resusus subsp. trigonus 
were found in eastern slopes of “El Cielo Biosphere Reserve.” Populations were also de- 
tected in pinyon pine forests dominated by Pinus nelsonii and Pinus cembroides in 
Miquihuana. 


a 


Cited specimens (Anderson 1964): MEXICO. ea a Miquihuana, 1900 m, Cowper 1889, 1958 
(POM, US); Tula, 1200 m, aAngae vom 

Spec B te, 1700 m, Herndndezet al. 2029 (MEXU); 
Bustamante, 1590 m, buat et al, 2033 “(MEXU): Bustamante, 2000 m, Jiménez 0018 (UAT); 
Miquihuana, 1524 m, Martinez-Avalos 054 (UAT); Miquihuana, 1487 m, Martinez-Avalos 0151 (UAT); 
Tula, 1250 m, Martinez-Avalos 0664 (UAT). 


4. Ari I t K.Sch nn subsp.trig (Weber 

Common name.—"Chaute,""Chautle” 

Geographic distribution.—Tamaulipas and Nuevo Leon. This species is distributed in 
the municipalities of Jaumave, Palmillas, San Carlos, Villa de Casas and Llera de Canales. 
The populations inhabit areas from 250 m to 1400 m, in different soil types. The centre of 
the distribution is the Jaumave Valley (Martinez-Avalos et al. 1993). 

Field observations indicate an ability to survive in exposed and eroded soils, A survey 
in the Jaumave Valley indicated 11 sub-populations with approximately 35,000 individuals. 
The plants exhibited a non-exclusive cactus-nurse plant relationship. The must impor- 
tant nurses were Prosopis laevigata and Cercidium praecox. Plants beneath healthy nurses 
exhibited better conditions than those with damaged nurses (and with smaller cano- 
pies).A. trigonus plants are able to thermo-regulate in open sun but individuals beneath 
the shade of nurses exhibited better thermo-regulation (Suzan unpublished data). 


E.F. Anderson & Fitz Maurice 


oF 


Cited specimens (Anderson 1964): MEXICO. Tamaulipas: Jaumave Valley, 760 m, Anderson 1580 
(POM, US, NY, MO, GH, K); Jaumave Valley, 760 m, Meyer & Rogers 3115 (MO,US); E of San Vicente, 
Jaumave Valley, 760 m,Anderson 1089 (POM, US, NY, MO, GH);W of Cd.Victoria, 700 m, Anderson 1153, 
1708 (POM, US). 


MARTINEZ AND SUZAN, DISTRIBUTION OF ARIOCARPUS IN TAMAULIPAS, MEXICO 181 


Specimens examined: MEXICO. Tamaulipas: W of San Antonio, 676 m, Arreola 275 (IBUG); NE 
of Tula, Sdnchez-Mejorada 2070 (MEXU); Jaumave 750 m, Herndndez et al. 2038 (MEXU); Jaumave, 750 
m, Herndndez 2047 (MEXU): Valle de Jaumave, 610 m, Martinez-Avalos 068 (UAT); Villa de Casas, 304 
m, Martinez-Avalos 0383 (UAT); San Carlos, 189 m, Martinez-Avalos 541 (UAT); San Carlos, 400 m, 
Martinez-Avalos 0745 (UAT). 


DISCUSSION 


It’s difficult to determine the ecological requirements that affects the geographic distri- 
bution of the genus Ariocarpus with the analysis of herbarium specimens, and even with 
field studies in the Chihuahuan Desert in México (Anderson 1958, 1961). In Tamaulipas 
this genus is restricted to the Chihuahuan Desert region, the Tamaulipan arid zone 
(Tamaulipas biotic province) and some specific tropical neighboring areas with similar 
climatic parameters (i.e. LLera Valley) (Fig. 1). The area in Tamaulipas where Ariocarpus 
grows is similar habitats in the neighboring states of Nuevo Leon and San Luis Potosi. 

The genus is distributed in altitudinal gradients from 200 to 2100 m (Table 1). Of 
the three species in the state, A. retusus subsp. trigonus shows the widest range in distri- 
bution, inhabiting different types of vegetation, soils and climate in the southwestern, 
central and southern parts of the state. Ariocarpus agavoides has the most restricted dis- 
tribution in an area smaller than 2 km2, being the species most highly disturbed due to 
direct and indirect factors such as habitat destruction for garbage incineration, marked 
hydric soil erosion, overgrazing, and excessive over-collection of individuals (Hernandez 
1992; Sanchez-Mejorada 1987). Ariocarpus kotschoubeyanus is a difficult species to find 
in the field due to its size and form; therefore, the true size of wild populations is rela- 
tively unknown.Ariocarpus retusus subsp. retusus inhabits the southwestern arid zones of 
the state and is well-adapted to the Chinuahuan Desert region. 


ACKNOWLEDGMENTS 


Héctor M. Hérnandez from the Universidad Nacional Autonoma de México provided 
permits to use and consult materials from this University. Donald J.Pinkava and James C. 
Solomon provided excellent comments and revisions to the paper. This study was sup- 
ported partially by the CONABIO (Comisién Nacional para la Conservacion y Uso de la 
Biodiversidad) projects P120, H14 


REFERENCES 


Anonimous, 1991. Diario oficial De La Federacién, (17 de Mayo). 1991. Listado de especies 
raras, amenazadas en péligro de extincidn o sujetas a proteccién especial y sus 
endemismos en la Republica Mexicana. SEDESOL. México. 

Anberson, E.F. 1958. En busca de Ariocarpus.. Cact. Suc. Méx. 3:79-84. 

. 1960.A revision of Ariocarpus (Cactaceae). |. The status of the proposed genus 
Roseo-cactus. Amer. J. Bot. 47:582-587 
.1961.A taxonomic revision of Ariocarpus, Lophophora, Pelecifora and Obregonia. 


Doctoral dissertation, Claremont Graduate School, Claremont, California. 


SIDA 19(1) 


-1962.A revision of Ariocarpus (Cactaceae).||.The status of the proposed genus 
Neogomesia. Amer. J.Bot. 49:615-622. 
_ 1963. A revision of Ariocarpus (Cactaceae). Ill. The status of the proposed ge- 
nus Roseocactus. Amer. J. Bot. 50:724—731. 
_1964.A revision of Ariocarpus (Cactaceae). IV. Formal taxonomy of the subge- 
nus Ariocarpus. Amer. J. Bot. 51:144-150. 
ANpERSON, E.F. and W.A. Fitz Maurice. 1998. Ariocarpus revisited. Haseltonia.5:1—20. 
Bravo,H. 1978. Las Cactdceas de México. Universidad Nacional Autonoma de México. México. 
1: 103-104. 
Bravo, H and H. SANcHEz-Mesoraba. 1992. Las Cataceas de México. (Universida Nacional 
Autonoma de México. México. 2:252-263. 
Cites. 1992. Convention on International Trade in Endangered Species of Wild Fauna and 
Flora. (11 of June of 1992), CITES. 
Garcia, E. 1964. Modificaciones al sistema de clasificacién climatica de KOppen (para 
adaptarlo a las condiciones de la Republica mexicana). Cartes, México. 
HernAnoez, H. and H. Gooinez. 1994. Contribucidn al conocimiento de las cactaceas 
mexicanas amenazadas. Acta Bot. Mex. 26:33-52. 
HerNnaAnpez, T. 1992. Aspectos bioldgicos de Ariocarpus agavoides (Castaneda) E.F. Ander- 
son. Cact. Suc. Mex. 37:40-45. 
Houmaren, P.,W. Keuken, and E.K. Scueriet. Index herbariorum. Part 1 vol 7.Regnum Veg. Vol. 
106 
Hunt, D.R. 1992. CITES. Cactaceae checklist. Royal Botanic Garden Kew. 
Jaccer, E.C. 1957. The North American deserts. Standford University Press. Standford, Cali- 


fornia. 
Marroauin, J.S.,G. Boris, R. Vazquez, and J.A.De La Cruz. 1964. Estudio ecoldgico - dasondmico 


de las zonas aridas del norte de México. Instituto Naciot val de Ir westigacior eS Forestales. 


Publicacidn especial. No. 2 México. 

Maartinez-Avatos, J.G., H.SuzAn, and C. Sacazar. 1993.Aspectos ecoldgicos y demograficos de 
Ariocarpus trigonus (Weber) Schumann. Cact. Suc. Mex. 38:30-38. 

Miticu, L.and J. Brum. 1877. The genus Ariocarpus. Cac. Succ. J.49:122-127. 

Rzepowski, J. 1978. La vegetacion de México. Ed. Limusa, México 

SANCHEZ-MEJORADA, H. 1982. Informa sobre la Reunion de Tucson para analizar el comercio 
de Cactaceas. Cact. Suc. Mex. 27:90-95 

. 1987. Observacion sobre el Estado de conservacion de doce especies 

amenazadas del noreste de México. Cact. Suc. Méx. 32:61-71. 

SECRETARIA DE PROGRAMACION Y PresupuEsTO (SPP). 1985. Sintesis geografica del estado de 
Tamaulipas. Instituto Nacional de Estadistica, Geografia e Informatica. 

Sureve, F. 1951. Vegetation of the Sonoran Desert. Carnegie Inst. of Washinton, publ. No. 
591. 

SuzAn, H., G. MALDA, J. Jimenez, L. HERNANDEZ, M. Martinez, and G. NasHan. 1989. Evaluacién de 
plantas amenazadas y en peligro de extinci6n en Tamaulipas. Biotam 1:20-27. 


— 


MARTINEZ AND SUZAN, DISTRIBUTION OF ARIOCARPUS IN TAMAULIPAS, MEXICO 183 

ULILC.N. (UNION INTERNACIONAL PARA LA CONSERVACION DE LA NATURALEZA Y DE LOS Recursos NATURALES). 
1985. Rare and threatened plants of Central America. January 85. 

VAzauez-Yanes, C.P. Pérez, and J. Rezeoowski 1979. La biosfera: extincién de las plantas. 
Naturaleza. 10:15-22. 

Vovines, A. 1988. Relacién de plantas Mexicanas raras o en péligro de extincion. In: 
Conservacion en México: Sintesis sobre Vertebrados Terrestres, Vegetacion y Uso del 
suelo, O.Flores Villela y P. Gerez, eds. Instituto Nacional de Investigaciones sobre Recursos 
Bioticos, Conservacion Internacional. México. 


184 SIDA 19(1) 


BOOK REVIEW 


Kevin Apams and Marty CassTevens. 1996. Wildfl of the Southern Appalachians: How 
to Photograph and Identify Them. (ISBN 0-89587-143-2, pbk.). John F. Blair Pub- 
lisher, 1406 Plaza Drive, Winston-salem, NC 27103 (910-768-1374; 910-768-9194 fax). 
$26.95.257 pp. Color ae 


You can find field guides that are less expensive, more compact, and include more cies, but this 
book has several features that mole it well a the money. First, it is the most pani Sere 
guide to herbaceous wildflowers of this specific geographic region. The southern Appalachians— 
sometimes called the botanical jewel of the eastern United States, with over 2500 species of flow- 
ering plants—includes sections of Virginia, West Virginia, Kentucky, Tennessee, North Carolina, South 
Carolina, Alabama, and Georgia. However, since many of the 180 species in this book have wide 
aes distributions, the book has application far beyond these states 
A second important feature of the book is the high quality of the sntoaniehe along with 
hints on on to photograph wildflowers. The photos are striking because they have exceptional 
focus contrast to separate the flowers from their backgrounds, a feat the authors usually accom- 
plished with a 75-300 mm zoom lens with an attached close-up lens. The 28-page chapter on 
“Photographing Wildflowers" has 23 photos of flowers (copies taken from the main section of the 
book) that demonstrate the excellent results that can be obtained with the equipment and tech- 
niques described. This chapter would be much improved if it also included photographs of the 
equipment items themselves (or the authors using the equipment), since many amateur photog- 
raphers may not be familiar with such items as a Bogen super clamp, Bogen accessory side arm, 
ball head, focusing rail, or wind barrier. The ten-page chapter on “Format” includes an additional 
even pages of “Photo Tips” and “Wildflower a Categories,” which correspond to the 
useful sia tips adjacent to each flower phot 
third important feature is the emphasis on conservation. The authors strive to educate the 
s ee wildflowers, in the belief that “education is the best environmental policy.”The book 
promotes environmental stewardship in the introductory chapter, and in the chapters on “The 
Natural Setting,”“Botanical Exploration of the Southern Appalachians,” and “Environmental Con- 
gee 


The authors hotograph rare flowers, rather than pick them. Their pho- 
tographic tips include Ways . minimize ae ee to flowers and their habitats, and warn that 
sometimes flowers should not even be photographed if intrusion into the habitat is too damag- 
ing.—Jerry Carpenter, Northern Kentucky University. 


SIDA 19(1): 184. 2000 


PLATEILEMA (ASTERACEAE: HELENIEAE) 
A NEW GENERIC REPORT FOR THE UNITED STATES 


Billie L. Turner 
Department of Integrative Biology 
niversity of Texas 
Austin, TX 78713, U.S.A. 
The monotypic genus Plateilema was first proposed by Cockerell (1904) in his treatment 
of the genus Hymenoxys for North America, basing this upon Actinella palmeri A. Gray, 
the sole member of section Plateilema. The latter was maintained as a genus by subse- 
quent workers (eg. Rydberg 1914), and | intend to accept its generic status in my forth- 
coming treatment of the Helenieae for Mexico (Turner 1999, in prep,). 

The single species concerned, Plateilema palmeri (A. Gray) Cockerell, acaulescent 
perennial herb superficially resembling Gaillardia comosa A. Gray, has been largely ig- 
nored by most phyletic workers (eg. Bierner and Jansen 1998), presumably because it is 
represented in herbaria by relatively few collections, mostly type material, the latter ob- 
tained from near Saltillo, Coahuila, Mexico. Indeed, prior to the present report, in addition 
to the type, the species was known by only a few skimpy collections from the Ocampo 
region of central Coahuila. 

The present article calls attention to two collections of the species from the United 
States, 200 kilometers or more from the previously known collections (Fig. 1), as follows: 


TEXAS. Brewster Co.: “Infrequent at Schuler mail box 44 mi S of Alpine; Terlingua Road; altitude 
3,600 ft," 10 Apr 1929, Henry T. Fletcher 219 (SRSC); “infrequent annual along Chalk Draw; Schuler 
Ranch; limestone soil; altitude 3,700 ft,"10 Apr 1929, Henry T. Fletcher 884 (SRSC). 
Henry T. Fletcher (1884-1955) was a remarkable man in the Brewster County region, 
serving his community in many ways:Vice President of the State National bank in Alpine, 
Rotary Club Governor, owner and manager of the Fletcher Cattle Company, to name but 
a few of his enterprises (Casey 1976). He was also an excellent collector and keen ob- 
server of range land plants, as attested to by his rarely cited but excellent publication for 
the times, Vegetation of the Green Valley Region (Fletcher 1928),a 40 km? of rolling plains 
with bordering mountains some 40-50 kms south of Alpine. Indeed, in a brief survey of 
the collections on file at SRSC, | calculate that he assembled and placed on deposit at 
that institution five thousand or more plant specimens. And his are not fragmentary 
collections as often collected by non botanists, rather they are neatly pressed, solidly 
mounted and in the best possible flowering and fruiting condition. In short, Fletcher 
collections served as the founding core of the SRSC herbarium, this subsequently built 
upon by the late Barton H. Warnock (Turner 1998). 

Both of the above specimens were said to have been collected on the same date, 
but possess very different collection numbers (279 and 884), suggesting that the collec- 


SIDA 19(1): 185-187. 2000 


186 SIDA 19(1) 


Fic. 1. Known distribution of Plateilema palmeri. 
tor arbitrarily assigned his collection numbers and/or dates, or else he collected over 
600 different specimens on the same date, an unlikely feat considering the time (April is 
usually a very dry month in the region concerned and even heavy early rains are unlikely 
to bring out a profusion of collectable species of this magnitude.) 

No collection book or records survive to sort out the peculiar numbers assigned 
Fletcher's specimens, but most likely he assembled his plants over several months and 
then assigned his plants numbers irrespective of their order of collection; similar anoma- 
lies as that found with the numbers alluded to in the above are also encountered with 
other assemblages of this collector. 

Since the above two sheets were only recently sorted out of a pile of unidentified 
specimens at SRSC little opportunity has been available to search the area concerned to 
see if the species might still exist. My hunch is that they are still represented by small 
populations along Chalk Draw, a poorly collected area of the Big Bend region of Texas. 
Unfortunately, most ranchers in the trans-Pecos at the present time are reluctant to al- 
low botanists on their ranchlands, fearful that an endangered taxon might be observed 
and as a result their use of their rangelands placed in some sort of jeopardy. 

Finally, it should be noted that two recent collections of Plateilema palmeri have 
been made in the state of Nuevo Leon,Mpio. Galeana, Mexico, by the Hinton family (Hinton 
et al.27195.TEX) some 100 kilometers south of the type locality.What was once thought 
to be a very localized species is now known to be quite widespread, albeit relatively rare, 
to judge from the collections at hand. 


TURNER, PLATEILEMA, A NEW GENERIC REPORT FOR THE U.S. 187 


REFERENCES 


Bierner, M.and R.K. JANSEN. 1998. Systematic implications of DNA restriction site variation in 
Hymenoxys and Tetraneuris. Lundellia 1:17-26. 


Casey, C.B.1976.Sul Ross State University, the cultural center of Trans-Pecos Texas. Pioneer 
Book Publishers, Seagraves, Texas. 

Cockerett, T.D.A. 1904. The North American species of Hynenoxys. Bull. Torrey Bot. Club 
34:461. 1904. 

FietcHer, H.T. 1928. Vegetation of the Green Valley region. West Texas Hist. Sci. Soc. Publ. 
2:6-50. 

Rypger, PA. 1914. Plateilema,in N. Amer. Fl. 34:101.1915. 

Turner, B.L. 1998. Barton H.Warnock (1911-1998). PI. Sci. Bull. 44:78-80. 

. 1999. Helenieae, in Comps of Mexico.Vol.5.Phytologia Memoirs (in prep.) 


188 SIDA 19(1) 


BOOK REVIEW 


Susyn Anorews, ALAN Lestic, and Criman ALexanoer, (eds). 1999. Taxonomy of Cultivated Plants. 
(ISBN 1 900347 89, hbk.). Royal Botanic Gardens, Kew. £ 27,553 Pp. 


This volume contains the results of the 1998 “Third International Symposium on the Taxonomy of 
Cultivated Plants,” sponsored jointly by the Royal Botanic Garden Edinburgh, the Royal Botanic 
Gardens, Kew, and the oe Horticultural a a unges the auspices of the International soley 
for Horticultural Science. T y HORTAX (Horticultural Taxonomy Group) 
and held at the Pollock Halls of the University of Edinburgh from 20-26 July 1998. 

The book is organized into introductory material, papers and shorter papers, but | found short 
papers or abstracts in each section. In the papers division, 43 works are distributed among ten 
conceptual sections, including: 1) Plant and germplasm collections, 2) Nomenclature in the orna- 
mental seed trade, 3) Intellectual property rights and plants, 4) registration of plant names, 5) prin- 
ciples of cultivated plant eee 6) the coat: Gelnihion ae cn 7) databases for 
collections, nomenclature and taxon my eyan dern technique: ing and taxonomy, 9) case 
studies in the ta ny Iti plant introductions. The next division covers short 
papers one were delivered as s posters curing the Sones and includes 50 abstracts divided 
among the r techniques, 2) databases and reg 
isters, 3) systematics 4) caeeicailon and nomenclature 5) conse! and collections, and finally 

6) publications. 

This volume truly sums up the latest in horticultural taxonomy from a plethora of points of 
view, and gives the reader a precis of current concerns and vacuums in our understanding of cul- 
tivated plant systematics. It is impossible here to thoroughly review each subject presented, but 
such subjects as living collections’ importance, germplasm collection, ex and in-situ conservation, 
role of herbaria, etc. are written to address the concerns of the practicing horticulturalist in the 
commercial world as much as ine academic. The chapter on “the cultivar-definition and recog 
tion” was particularly noteworthy and significal tly enhanced my understanding of the ge 
horticultural texonamy faces. che thoroughly convinced that th papers on fingerprinting of culti- 

ens Itant 


presented | Diy oats tic | indigens and their culti- 


gens, will be extremely valuable as lessons for us all, to help us tackle what has largely been a 
neglected area. Several of the molecular papers highlighted the fact that many cultigens may have 
their origin traced to but a few individuals, rather than representing an adequate sample of the 
infraspecific genetic ake present in wild populations of the species. These papers can surel 
guide those who are attempting endangered a rescues, or introducing wild species into 
cultivation and have poe conservation consequences. 

| believe this book can open the ‘Adiionslhy ee systematist’s eyes to the many prob- 


{ =l/ | 


lems facing horticultural taxonon ny and hould foster a ee appreciation for it.While the papers 


are brief summaries, their extensive permit further study of each area, aes the boo 
an exueinely useful tool. ae poo physical alas are a a of me other fine Kew public 
tions, and the volume of worth the price. igre lel 


recommend it to anyone working in 1 botanical gardens or universities, those who train future sys- 
tematists, plant conservationists or epee ace —John J. Pipoly Ill; Head of Research, Botanical 
Research Institute of Texas, 509 Pecan St., Ft. Worth, TX 76126-3019, USA. 


SIDA 19(1): 188. 2000 


WHICH NON-NATIVE PLANTS ARE INCLUDED 
IN FLORISTIC ACCOUNTS? 


Guy L.Nesom 
North Carolina Botanical Garden 
Coker Hall, CB 3280 

University of North Carolin 
Chapel Hill, NC 27599-3280, on 


ABSTRACT 


It is suggested that four terms provide an ad t al description of the major categories of 
non-native plants in our flora: cultivated, persisting, ver ae naturalized. Comparative definitions 
are given for seven related terms that Gssciing: the non-native origin and floristic integration of 
vascular plant species in North A alien, escaped, established, exotic, introduced, 


and non-native. 


RESUMEN 


Se sugiere sas cuatro términos dan una descripciédn general adecuada de las grandes categorias 
d nuestra flora: cultivada, persistente, abandonada, y naturalizada. Se dan 


las planta 
elas p d 


definiciones dompareines de siete términos relacionados que describen el origen no nativo y la 
integracion floristica de plantas vasculares en Norteamérica: adventicia, extrana, escapada, 
establecida, exotica, introducida, no-nativa. 


Plants are usually included in floristic accounts of the North American flora on the basis 
of their status either as native or as non-native but integrated into the flora to some 
degree. For non-native plants, an assessment of their integration is based primarily on 
reproductive status and dispersive success. The more summary floristic accounts com- 
monly do not provide information necessary for a clear assessment of the degree of 
floristic integration of non-native plants. 

A selection of general statements of intent for major floristic studies (from numer- 
ous examined) indicates that a broad range of non-native taxa is treated. Naturalized 
plants are invariably treated, but the definition of “naturalized” is not consistent, and plants 
recognized as“waifs" and as merely “persistent” may or may not be included. 


Radford et al.(1968) for the Carolinas: "species of vascular plant known to grow with- 
out cultivation.” 

Correll and Johnston (1970) for Texas: all native and naturalized flowering plants and 
ferns.” 

Voss (1972) for Michigan: “all species ... known to grow (or to have grown) outside of 
cultivation in Michigan, whether originally native or not.” See Voss (Preface, 1996) for 
related comments. 

Great Plains Flora Association (1986) for the Great Plains: “all vascular plants known 


SIDA 19(1): 189-193. 2000 


190 SIDA 19(1) 


to occur spontaneously”—"the native members of the flora and those introductions 
that appear to be permanently established outside of intentional cultivation." 

Welsh et al. (1993) for Utah: “All indigenous plant species known to occur in Utah are 
included in the flora. Introduced plants are also covered, but not so intensively as are 
the native ones.“ This flora attempts to present coverage of all established species, all 
common adventive taxa,and many of the commonly grown cultivated species." 

Gleason and Cronquist (1993) for northeastern United Stat ladj 
plants “growing wild” (including—from the text—“often escaped from cultivation,’ 
“adventive’and “rarely adventive,"“occasionally introduced,”and"sparingly established” 

Hickman (1993) for California: “plants growing wild in California.”“All aliens that have 
become an integral part of the Californian flora are included. The general policy was 
not to include (or to note only in passing) waifs or non-reproducing but long-persist- 
ing individuals or clones.” For non-native species the area of origin is specifically named 
(eg.,"native to Europe’) and a brief description of its habitat, geographic range, and 
relative abundance is provided, without the use of any further terminology. 

Cooperrider (1995) for Ohio: “the native members of these families, the alien plants of 
these families that have become established or that appear occasionally in the state's 
flora, and to a lesser extent the families’ major cultivated plants.” 

Flora of North America Editorial Committee (1996) for North America north of 
Mexico: to be included in full are“all native plants”and “introduced taxa that are natu- 
ralized or found frequently outside cultivation;” to be mentioned in the discussion are 
“naturalized plants now known only as historic records” and “non-native, economically 
important or extensively cultivated plants that tend not to escape cultivation (e.g., 
alfalfa).” 

Diggs et al. (1999) for north central Texas: all known native and naturalized vascular 
plant species ....A few long-persistent (e.g., Ficus carica—the common fig), but appar- 
ently non-reproductive taxa have been included because of the likelihood of them 
being encountered.” 

Kartesz (1999) for North America north of Mexico: “all Known native, naturalized or 
waif occurrences of vascular plant taxa within the region.” Also included, but not ex- 
plicitly, are non-native taxa recognized as “persisting” (as described below). 


anada: 


— 


Despite the demonstration in the recent Jepson Manual (Hickman 1993) that the floris- 
tic integration of non-native plants can be precisely described without specialized ter- 
minology, a minimal set of descriptive terms is useful if consistently applied. The four 
terms below provide an adequate general description of the major categories of non- 
native plants in our flora. 


1. Cultivated 

Plants of native and non-native species, across various categories of duration and habit, 
that are deliberately planted, actively maintained, and grown for ornament, interest, con- 
sumption, or other use. These are rarely included in floristic studies (but see Welsh et al. 


NESOM, NON-NATIVE PLANTS IN FLORISTIC ACCOUNTS 191 


1993). Native species are sometimes encountered only as cultivated plantings, and these 
should be distinguished from naturally occurring plants. Another distinction can be drawn 
by recognizing a cultigen (a cultivated plant of unknown or obscure origin) as different 
from a cultivar (a cultivated plant of known origin). 


2. Persisting 

Perennial plants (woody, suffrutescent, and herbaceous) cultivated for ornament or in- 
terest and remaining in place at old home sites, roadsides, etc., without human assis- 
tance after the site has returned to a more natural state, sometimes after evidence of 
associated human presence is decayed or completely gone. Such plants are not repro- 
ducing or at least not spreading beyond the original planting. The difference between 
“cultivated” and “persisting” is primarily a temporal one, referring to the duration of exist- 
ence, and persisting plants usually are not included in floristic accounts with the same 
status of native and naturalized species. Because, however, in a sense they grow outside 
of cultivation, they may appear in unannotated summaries or they may be explicitly 
described in floristic studies as “persisting.” 


3. Waif 

Plants of non-native species growing outside of cultivation but not maintaining a viable 
population for more than one or a few seasons. A number of waifs in the North Ameri- 
can flora are known only as single historic records; other essentially non-reproductive 
plants as waifs apparently may be recurrent. Some species reported with new distribu- 
tion records may become known more precisely as waifs as the site of their reported 
occurrence is reinvestigated. More than a single season is required to observe the repro- 
ductive status of such a plant or population, and some of those initially suspected of 
being waifs may become naturalized. Waifs may originate from various kinds of natural 
and human-mediated long-distance dispersal. They may originate from seeds washed 
out from garden plantings and commonly are found at sites where seeds are mixed 
with other exotic materials (e.g., ballast dumps, wool mills). Tomato seeds, which pass 
unharmed through digestive tracts and sewage treatment, are spread in sludge-based 
fertilizer and may spawn waif plants in repeated cycles. The usage here for “waif” is 
a good match for its ancient European etymology as “ownerless property, stray, or 
homeless.” 


4. Naturalized 

Plants of non-native species accidentally or deliberately introduced into the flora, now 
reproducing and maintaining viable populations from year to year (more than just one 
or a few seasons), and dispersing without deliberate h assistance beyond the popu- 
lation or populations of original establishment. The degree of naturalization may vary 
from widespread and abundant occurrence to local establishment (from recent intro- 
duction or sluggish reproduction and dispersal). Perhaps the most difficult in interpreta- 
tion are cultivated plants spreading slightly beyond their original planting—these might 
be best characterized as waifs, if their reproductive ability is slight (e.g., /mpatiens spp), 


192 SIDA 19(1) 


or they may be“incipiently naturalized’ if reproduction and spread appear to be vigor- 
ous, even though restricted in area. Species native to North America may become natu- 
ralized in areas of the continent other than where recorded as native in the past or present. 
Some species apparently have a mixture of native and non-native North American popu- 
lations (e.g., Achillea millefolium, Galium aparine, and Prunella vulgaris). 


Associated Terms 

In addition to the four terms suggested above to cover the major categories of integra- 
tion of species and other taxa in the North American flora, various other related terms 
have commonly been used in floristic works to characterize the origin and integration 
of such plants. The usage of these terms varies considerably, and without other refer- 
ences that provide comparative definitions in a floristic context, it is hoped that those 
provided here will contribute toward more consistent application."Native species,"alien 
species,” and “introduction” are formally defined in the recent Executive Order (U.S. Ex- 
ecutive Order 13112).Because of the difficulty in ordering this set of terms in the context 
of floristic criteria, they are listed here alphabetically. 


Adventive—non-native, becoming naturalized either from deliberate plantings or acci- 
dental introduction. 

Alien—non-native, commonly considered to have originated from a different continent, 
but not necessarily exclusive of an origin from an ecologically different region of the 
same continent. 

Escaped—non-native, becoming naturalized from deliberate plantings, usually only in 
a local area or represented by relatively few individuals over the range of naturalization. 
Established—non-native, similar to ‘adventive” and “introduced” but with the implica- 
tion of being more securely naturalized. 

Exotic—non-native, essentially similar to the term “alien,” but with stronger implication 
of an extra-continental origin.|n the recently issued "Synonymized Checklist” for the North 
American flora (Kartesz 1999), exotic species in the continental flora are considered to 
originate from other continents or from Mexico or Central America. 
Introduced—non-native, released accidentally or deliberately into the flora and grow- 
ing without cultivation, at least for one generation, Similar to“adventive” but with slightly 
stronger implication of having originated through plants used for horticultural or prac- 
tical purposes. 

Non-native—originating from outside the geographic region of concern, either from a 
different continent or from an ecologically different area of the same continent. Among 
a group of terms often used interchangeably (adventive, alien, exotic, introduced, non- 
native),"non-native’ is the most general, at least in the definitions suggested here. 


ACKNOWLEDGMENTS 
Helpful comments from John Kartesz, Robert Kiger, Rogers McVaugh, Rudolph Schmid, 
and John Strother are appreciated.| am grateful for help with publications costs provided 
by the Mouzon and Mouzon Fund of the University of North Carolina Herbarium (NCU). 


NESOM, NON-NATIVE PLANTS IN FLORISTIC ACCOUNTS 193 


REFERENCES 


U.S. Executive Orver 13112. 1999 [3 Peon: eee Order on invasive alien species. 
W.J. Clinton, The White House. [http:/, | itet gov/urires/I2R?urn:pdi:// 
oma.eop.gov.us/ 1999/2/3/14.text.2] 

Cooperniner, I.S. 1995. The Dicotyledoneae of Ohio. Part 2: Linaceae through Campanulaceae. 
Ohio State Univ. Press, Columbus. 

Corrett, D.S. and M.C. JoHnston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner 

Diccs, G.M., Jr., B.L. Liescome, and R.J. O’Kennon. 1999. Shinners & Mahler's illustrated flora of 
North Central Texas. Sida, Bot. Misc. No. 16. 

Fora oF NortH America Epitoriat COMMITTEE, EDS. 1996. Flora of North America North of Mexico: 
Guide for contributors (August 1996 version). FNANM Organizational Center, Missouri 
Botanical Garden, St. Louis. 

Gteason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada (ed. 2). The New York Botanical Garden, Bronx. 

GREAT PLAINS Flora Association. 1986. Flora of the Great Plains. Univ. Press of Kansas, Lawrence 

Hickman, J.C. (ed.). 1993. The Jepson manual: Higher plants of California. Univ. of California 
Press, Berkeley. 

Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vas- 
cular flora of the United States, Canada, and Greenland. First Edition.In:Kartesz,J.T.,,and 
CA. Meacham. 1999. Synthesis of the North American flora, Ver. 1.0. North Carolina 
Botanical Garden, Chapel Hill. 

Raprorb, A.E., H.E. AHies, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill. 

Voss, E.G. 1972. Michigan flora. Part |, Gymnosperms and Monocots. Cranbrook Inst. Sci- 
ence Bull.55 and Univ. of Michigan Herbarium, Ann Arbor. 

Voss, E.G. 1996. Michigan flora. Part Ill, Dicots (Pyrolaceae—Compositae). Cranbrook Inst. 
Science Bull.61 and Univ. of Michigan Herbarium, Ann Arbor. 

We sh, S.L., N.D. Atwooo, S. GoopricH, and L.C. Hiccins. 1993. A Utah flora. Brigham Young 
Univ., Provo, UT. 


SIDA 19(1) 


BOOKS RECEIVED 


N.W.Simmonos and J. Smaart (chapter 9 by S.Millam and W.Spoor).1999.Principles of Crop 
Improvement. (ISBN 0-632-04191-9, hbk). Blackwell Science Ltd, 350 Main Street, 
Malden, MA 02148-5018, U.S.A. (800-759-6102; 781-388-8255 fax; www.blackwell- 
science.com). $104.95, hbk. 412 pp. Illustrated. 

From the cover: "The second edition of this highly regarded text provides a broad introduction to 

the principles of plant breeding. It presents a synoptic view of crop history, linking original studies 

of crop origin and early evolution with recent and possible future trends in breeding, The text has 
been revised and expanded to include recent developments relevant to crop improvement. There 
is a new chapter on biotechnology and new information on the use and conservation of genetic 
resources. Throughout the book examples are drawn from a wide range of crops and locations, and 
flow diagrams are used to supplement and expand nia given in the text. This book is 
aimed primarily at senior undergraduate and graduate students of agriculture, horticulture and 
applied biology.” Contents: 1) The evolution of crops; 2) Basic features of plant breeding; 3) Objec 
tives of plant breeding; 4) Genetic aspects: Populations and selection; 5) Breeding plans; 6) Trials 

and Multiplication; 7) Disease resistance; 8) Special techniques; 9) Biotechnology and crop im- 

provement; 10) New crop and genetic conservation; 11) The social context; Bibliography, Index of 

plant names, and General index. 


Epward A.Weiss (Foreword by Sir Charles Pereira, FRS). 2000. Oilseed Crops, Second Edi- 
ion. (ISBN 0-632-05259-7, hbk.). Blackwell Science Ltd, 350 Main Street, Malden, MA 
02148-5018,U.S.A. (800-759-6102; 781-388-8255 fax; www.blackwell-science.com). 
$156.95, hbk. 364 pp. Illustrated. 
From the cover:"Oilseeds are a very important component of semi-tropical and tropical agriculture, 
Hal easily available and highly nutritious human and animal food. Many also have industrial 
s and are relatively easy to incorporate into coy sania products. This book gathers 
to oe scientific, economic and field pract t ee Aaa tas ole oe 
After a discussion on world production and trade in oilseeds there are separate chap 
groundnut, safflower, sesame, oe and sunflower, crambe, niger and jo fone: A final chapter ae in 
detail at oilseed p ns and the great variety of products and uses for oilseeds. This 
new edition has been somplciely ued to inclige the cea eg Teta! hawsiak the ne 
ence of biotechnology on crops " 
‘The book is a valuable reference for all involved directly or indirectly with oilseed proedeton 
research and processing.” 


SIDA 19(1): 194. 2000 


CINNA AND LIMNODEA (POACEAE): NOT CONGENERIC 


David M. Brandenburg John W.Thieret 
The Dawes Arboretum ee sclenices 
Newark, OH 43056 U.S.A. Northern K iV 


Highland Hei ights, kVA 1099 uU SA 


A decade after Clayton and Renvoize (1986) suggested that Limnodea LH. Dewey is “a 
prairie version of Cinna,"Tucker (1996) reduced this long-recognized grass taxon to syn- 
onymy under Cinna L. As the only justification for the change he noted that Limnodea 
arkansana (Nuttall) Dewey “has been treated as constituting a monotypic genus ... but 
there seem to be no compelling differences other than length of lemma awn, a minor 
distinction."We regard this nomenclatural aberration as being unsupportable. A similar 
reduction of Limnodea to synonymy occurred in 1841 when Trinius in Steudel (Steudel 
1841) transferred it to Limnas, a Siberian genus of two species from which Limnodea 
differs in several features (Bentham 1881). (On the basis of the lemma awn of Limnodea 
alone, the genus could just as well be allied with Stipa.) 

Our purpose in this note is to point out that, in addition to features of the awn, 
there are indeed other compelling differences between Limnodea and Cinna and that 
considering these two taxa to be congeneric is untenable. Unfortunately the new com- 
bination has already been adopted in two recent floras (Wunderlin 1998; Yatskievych 
1999). 

Nuttall (1835-1836) recognized that a grass he had in hand constituted a distinct 
genus, which he called Greenia Nutt. Unfortunately, this was a preoccupied name, as 
were two other generic names later applied to the genus, Sclerachne Torrey in Trinius 
(Trinius 1841) and Thurberia Bentham (Bentham 1881). Finally, Dewey (1894) published 
the generic name Limnodea, which has hitherto remained unchallenged. Whatever the 
generic name, for over a century and a half Limnodea has been recognized as distinct by 
agrostologists and other botanists (except Tucker and the two authors of floras cited 
above) who have dealt with it; we list some of these here (e.g., Allen 1980; Beal 1896; 
Beetle 1977;Bentham 1881;Bentham & Hooker 1883;Bews 1929;Campbell 1985;Clayton 
& Renvoize 1986; Correll & Johnston 1970; Dewey 1894; Diggs et al. 1999; Dore 1956; 
Featherly 1946; Gould 1968, 1975, 1979; Gould & Box 1965; Gould & Shaw 1983; Great 
Plains Flora Association 1977, 1986; Hackel 1887, 1890; Hatch et al. 1999: Hitchcock 1920, 
1935, 1937,1951;Johnston 1990; Kartesz 1994; Kartesz & Meacham 1999;Lamson-Scribner 
1900a, 1900b; Lonard 1993; Powell 1994; Shinners 1972; Silveus 1933: E.B. Smith 1994; J.P. 
Smith 1981;Stebbins & Crampton 1961; Terrell 1971;Vasey 1883, 1885;Watson & Dallwitz 
1992). The genus Limnodea is also recognized in the Missouri Botanical Garden's New 
World Grass Checklist (<ww bot.org/mobot/tropicos/Poa/Limnodea.html> Ac- 
cessed 1999 September 30) and in the Texas A&M University Grass Images (<http:// 
www.csdl.tamu.edu/FLORA/image/k4606700.htm>Accessed 1999 September 30). 


— 


SIDA 19(1): 195-200. 2000 


196 


Taste 1.Comparison of various characters of Cinna and Limnodea. 


SIDA 19(1) 


Character Cinna Limnodea 

Flowering/ Late estival/autumnal Late hibernal/vernal 
fruiting time 

Duration Perennial Short-lived annual 

Spikelet Laterally compressed Subterete 
compression 

Texture of Membranous Chartaceous™ 
glumes 

Abaxial surface Sharply keeled Rounded (or broadly 
of glumes keeled in distal portion) 

Margin of glumes Flat, hyaline Inrolled, not hyaline 


Vestiture of 
glumes 


Texture of 
lemma 


Abaxial surface 
of lemma 


Awn 
Palea 


Veins of palea 


Number of 
stamens 


Endosperm 
Chromosome number 


Articulation 


Habitat 


Sparingly to moderately 


Membranous 


Sharply keeled 


Much shorter than spikelet, 


Straight, not twisted, not 
hygroscopic, or lacking 
Well developed, 
prominent, relatively long 
compared to olem 


1 or if 2, these closely 


nearly to tip of palea 


lor2 


Semi-liquid (“so 
-fleshy’ ; oo io 
Dore 1956) 


2n = 28 (Bowden 1960; 
Davidse & Pohl 1978; 
Pohl & Davidse 1971) 
Spikelets not readily 
disarticulating 


Woods 


Sparingly to densely scabrous 
or pilose especially on veins 
(var. pilosa (Trin.) Scribn.) 


Chartaceous 
Rounded 


Much longer than spikelet, 
geniculate, twiste 
proximally, hygroscopic 


Poorly developed, 
dian short compared 


2-veined only at base 


Liquid (Brown 1955; 
Terrell 1971; Dore 1956) 


2n = 14 (Watson & Dallwitz 
1992 onward 


Spikelets readily 
disarticulating 


Prairies and disturbed areas 


*The describer of Limnodea, Nuttall (1835-1836), wrote of the glumes as a coriaceus” and, in 
English translation, as“indurated."His adjectives formed the basis for Torrey’s new name, Sclerachne 


NOT CONCENERI 197 


Cinna was recently revised by Brandenburg et al. (1991). Differences between its 
four species and Limnodea are summarized in Table 1. 

True it is, of course, that Liminodea and Cinna share certain features. Among these 
are 1-flowered spikelets, articulation below the glumes, extended rachilla, and awned 
spikelets (sometimes awnless in Cinna). Each of these features is found in other genera 
of Agrostideae sensu Hitchcock (1951). The combination of the four may be unique to 
Cinna and Limnodea—it is among U.S. grasses—and may indicate a possible relation- 
ship between the two genera. Bentham and Hooker (1883) suggested a resemblance 
between Limnodea and Polypogon; Nuttall (1835-1836) considered Limnodea to be “al- 
lied to Oryzopsis." However, resemblance and possible relationship are not necessarily 
the same. Combining the two genera because of a feature of the awn is, we suggest, 
most emphatically negated by the many other features separating them. 

Distinguishing among the spikelets of the species of Cinna takes some experience, 
but distinguishing between spikelets of Cinna and those of Limnodea takes hardly more 
than a glance. As an experiment we removed the awns of several spikelets of Limnodea 
and then mixed them with a number of Cinna spikelets representing all four species of 
that genus. Next we invited several colleagues to examine the mixture under a dissec- 
tion scope to determine which spikelets “did not belong.”No one had any trouble point- 
ing out the Limnodea spikelets—even without their conspicuous awns—as the “differ- 
ent” ones. 

The genera Cinna and Limnodea are easily separable by the following synoptic key. 
1. Spikelets | II rel membranous, sharply keeled; lemmas awn- 

less or with a raig at: awn finch shorter than the spikelet; palea well developed, 

relatively long compared to lemma, the veins 1 or 2, ca {0 or See te pales 

tip, when veins 2, the veins closely approximate; stamens 1 or tival/autum 

nal perennials Cinna 
. Spikelets subterete; glumes chartaceous, rounded on back;lemmas with a genicu- 

late, twisted, hygroscopic awn much longer than the spikelet; palea poorly devel- 

oped, relatively shart compared to lemma, 2-veined only at base; stamens 3; 

short-lived, late hibernal/vernal annual Limnodea 


paar 


CONCLUSION 


The inclusion of Limnodea in Cinna introduces a markedly discordant element into the 
latter small and well-circumscribed genus. The four currently recognized species of Cinna 
are quite similar to each other and all differ consistently from the monotypic Limnodea 
in several significant features, especially in the strikingly different spikelet morphology, 
These genera are not congeneric. 


REFERENCES 
ALLEN, C.M. 1980. Grasses of Louisiana. Univ. Southwestern Louisiana, Lafayette. 
Beat, W.J. 1896. Grasses of North America. Henry Holt and Company, New York. 
Beetle, A.A. 1977. Noteworthy grasses from Mexico V. Phytologia 37:317-407. 


198 SIDA 19(1) 


BentHam, G. 1881. Notes on Gramineae. J. Linn. Soc. Bot. 19:14-134. 

BentHaM, G.and J.D. Hooker. 1883. Gen. Plant. III(2). 

Bews, J.W. 1929. The world’s grasses. Longmans, Green and Company, London. 

Bowoen, W.M. 1960.Chromosome numbers and taxonomic notes on northern grasses. Ill. 
Twenty-five genera. Canad. J. Bot. 38:541-557. 

Branpensurc, D.M., W.H. BLackweLt, and J.W. THieret. 1991. Revision of the genus Cinna 
(Poaceae). Sida 14:581—586. 

Brown, W.V. 1955. A species of grass with liquid endosperm. Bull. Torrey Bot. Club 82: 
284-285. 

Campsett, C.S. 1985. The subfamilies and tribes of Gramineae (Poaceae) in the southeast- 
ern United States. J. Arnold Arbor. 66:123-199. 

Cayton, W.D.and S.A. Renvoize. 1986.Genera graminum. Grasses of the world. Her Majesty’s 
Stationery Office, London. 

Corrett, D.S. and M.C. Jonnston, 1970. Manual of the vascular plants of Texas. Texas 
Research Foundation, Renner 


Daviose, G. and R.W. Port. 1978. Chromosome numbers of tropical American grasses 
(Gramineae). Ann. Missouri Bot. Gard. 65:637-649. 

Dewey, L.H. 1894. Gramineae. In: J.M. Coulter. 1891-1894. Botany of western Texas. Contr. 
U.S. Natl. Herb. 2:484-551 

Dices, G.M., Jr, B.L. Liescome, and R.J. O’Kennon. 1999. Shinners and Mahler's illustrated flora 
of north central Texas. Botanical Research Institute of Texas, Fort Worth. 

Dore,W.G. 1956.Some grass genera with liquid endosperm. Bull. Torrey Bot. Club 83:335— 
S37. 

FeatHerty, H.I. 1946. Manual of the grasses of Oklahoma. Bull. Oklahoma Agric. Mechan. 
College 43(21). 

Foster, R.C. 1944. The publication data of Nuttall’s “Arkansas flora.” Rnodora 46:156-157. 

Goutp, FW. 1968. Grass systematics. McGraw-Hill Book Company, New York. 

Goutp, FW. 1975.The grasses of Texas. Texas A&M Univ. Press, College Station. 

Gouto, FW. 1979. A key to the genera of Mexican grasses. Texas Agric. Exper. Station, Col- 
lege Station. 

Goutb, EW. and T.W. Box. 1965. Grasses of the Texas Coastal Bend. Texas A&M Univ. Press, 
College Station. 

Gouto, FW. and R.B. SHaw. 1983. Grass systematics. 2nd ed. Texas A&M Univ. Press, College 
Station. 

Great PLains Fora Association. 1977. Atlas of the flora of the Great Plains. lowa State Univ. 
Press, Ames. 

Great PLAINS FLora AssociATION. 1986. Flora of the Great Plains. Univ. Press of Kansas, Lawrence. 

Hacket, E. 1887. Gramineae (echte Grdser). In A. Engler and K. Prantl, Die naturlichen 
Pflanzenfamilien II(2). 

Hackel, E. 1890. The true grasses. Translated from Die naturlichen Pflanzenfamilien by F. 
Lamson-Scribner and E.A. Southworth. Henry Holt and Company, New York. 


BRANDENBURG AND THIERET, C/NNA AND LIMNODEA: NOT CONGENERIC 199 


HATCH, S.L., J.L. ScHuster, and D.L. Drawe. 1999. Grasses of the Texas gulf prairies and marshes. 
Texas A&M Univ. Press, College Station. 

HitcHcock, A.S. 1920. The genera of grasses of the United States. U.S.D.A. Bull. 772. 

HitcHcock, A.S. 1935. Manual of the grasses of the United States. U.S.D.A. Misc. Publ. 200. 

HitcHcock, A.S. 1937. Limnodea. N. Am. Flora 17(7):538-539. 

HitcHcock, A.S. 1951. Manual of the grasses of the United States. 2nd ed., revised by Agnes 
Chase. U.S.D.A. Misc. Publ. 200. 

Joxnston, M.C. 1990. The vascular plants of Texas. A list, up-dating the Manual of the vascu- 
lar plants of Texas. Published by the author, Austin?, Texas. 

Karresz, J.T.1994.A synonymized checklist of the vascular flora of the United States, Canada, 
and Greenland. Timber Press, Portland, Oregon. 

Karresz, J.T., and CA. MeAcHam. 1999. Synthesis of the North American flora. Version 1.0. 
CD-ROM. ISBN 1-889065-05-6. North Carolina Botanical Garden, University of North 
Carolina, Chapel Hill. 

Lamson-Scrianer, F.1900a. American grasses—I.U.S.D.A. Div. Agrostol. Bull. 7 (3rd ed). 

Lamson-Scriener, F. 1900b. American grasses—lll.U.S.D.A. Div. Agrostol. Bull. 20 (revised). 

Lonaro, R.|. 1993. Guide to grasses of the lower Rio Grande Valley, Texas. Univ. Texas-Pan 
American Press, Edinburg, Texas. 

Nurtatt, T. 1835-1836. Collections towards a flora of the Territory of Arkansas. Trans. Am. 
Philos. Soc. n.s.5:139-203. [Greenia, 142; published 1835 (Foster 1944)] 

Pout, R.W. and G. Davidse. 1971. Chromosome numbers of Costa Rican grasses. Brittonia 
23:293-324. 

Powel, A.M. 1994. Grasses of the Trans-Pecos and adjacent areas. Univ. Texas Press, Austin. 

SHINNERS, LH. 1972. Shinners’ spring flora of the Dallas-Fort Worth region[] Texas. 2nd ed. 
Prestige Press, Fort Worth. 

Siwveus, W.A. 1933. Texas grasses. Published by the author, San Antonio, Texas. 

SMITH, E.B. 1994. Keys to the flora of Arkansas. Univ. Arkansas Press, Fayetteville. 

Smith, J.P. Jr.1981.A key to the genera of grasses of the conterminous United States. Mad 
River Press, Eureka, California. 

STEBBINS, G.L. and B. Crampton. 1961. A suggested revision of the grass genera of North 
America. Recent Advances Bot. 1:133-145. 

STEUDEL,E.T. 1841. Nomenclator botanicus.Editio secunda. Pars Il. Lit. L—Z.J.G. Cotta, Stuttgart. 

Terrett, E.E. 1971. Survey of occurrences of liquid or soft endosperm in grass genera. Bull. 
Torrey Bot. Club 98:264-268. 

Trinius, C.B. 1841. Gramina agrostidea, Il. Callus rotundus. (Agrostea). Mém. Acad. Imp. Sci. 
St-Pétersbourg. Sér. 6. Sci. Math., Phys. Nat. Seconde Partie: Sci. Nat. 6:247-390. 

Tucker, G.C. 1996. The genera of Pooideae (Gramineae) in the southeastern United States. 
Harvard Papers Bot. 9:11-90. 

Vasey, G. 1883. The grasses of the United States. U.S.D.A. Spec. Rep. 63. 

Vasey, G. 1885.A descriptive catalogue of the grasses of the United States. Gibson Broth- 
ers, Washington, D.C 


aa 


200 SIDA 19(1) 


Warson, L. and MJ. Datuwitz. 1992. The grass genera of the wohl ca inteina tion 
Wallingford, Oxon, United Kingdom. (See also <http://biodi delta/grass/ 
>Accessed 1999 October 5.) 

WuNDERLIN, R.P 1998. Guide to the vascular plants of Florida.Univ.Press of Florida, Gainesville 

YarskievycH, G. 1999. Steyermark’s flora of Missouri. Vol. 1. Missouri Department of Conser- 
vation, Jefferson City, in cooperation with the Missouri Botanical Garden Press, St.Louis. 


THEMEDA QUADRIVALVIS (POACEAE: ANDROPOGONEAE) 
IN KANSAS: AN EXOTIC PLANT INTRODUCED 


FROM BIRDSEED 
E.Gene Towne lralee Barnard 
Division of Biology Herbarium, Division of Biology 
Kansas State University Kansas State University 
Manhattan, KS 66506-4901, U.S.A. Manhattan, KS 66506-4901, U.S.A. 
egtowne@ksu.edu iralee@ksu.edu 


In November 1998, a peculiar grass growing in a residential yard in Wamego 
(Pottawatomie County), Kansas was submitted to the Herbarium at Kansas State Univer- 
sity (KSC) for identification. The grass was identified as Themeda guadrivalvis (L.) Kuntze, a 
plant native to southeastern Asia. In the United States, this grass has been collected in 
Louisiana (Brown 1945) and Florida (Wunderlin 1998), presumably as an escapee from 
cultivation. A search of herbaria records revealed that Themeda also has been collected 
in California (RSA). 

Themeda quadrivalvis can be recognized by the following description: Annual, 20- 
50 cm tall; blades papillose-ciliate on the lower margins; ligule membranous, lacerate, < 
1 mm long; inflorescence a lax flabellate cluster of racemes, each subtended by a strongly 
keeled spathe; spikelet clusters with a fertile sessile spikelet surrounded by four infertile 
sessile spikelets and two infertile pedicellate spikelets, all partially enclosed by a spatheole; 
fertile spikelet terete, bearded, 4.7-7.2 mm long with a 3-4 cm long geniculate awn, 
disarticulating obliquely to form a pointed callus; infertile spikelets 4-5 mm long;flower- 
ing Sep-Oct. The specimens may be assigned to variety helferi based on the presence of 
tuberculate hairs on the involucral spikelets and the plant stature (Bor 1960). The inflo- 
rescence of Themeda is a complex arrangement of spikelet clusters that is described in 
greater detail by Baird and Thieret (1985). 

In a follow-up examination of the collection site, eight additional Themeda plants 
were found growing under a bird feeder and in an unmowed area near the porch. The 
yard was semi-shaded, and the lawn was predominately crabgrass (Digitaria sanguinalis) 
and bluegrass (Poa pratensis). Microscopic examination of birdseed from the feeder found 
numerous Themeda spikelets among the thistle (Guizotia abyssinica: Asteraceae) seeds, 
indicating that it was the source of the introduction. 

To determine if Themeda seeds were prevalent in birdseed, nine sacks of commer- 
cial thistle seed with different lot numbers were purchased and inspected thoroughly 
(30.1 kg of seed). All of the sacks contained Themeda and other contaminant seeds, which 
were removed and compiled. Three hundred of the Themeda seeds and numerous other 
unidentified seeds were planted in vermiculite in a 22° C greenhouse and monitored for 
five months. In the spring, an additional 900 Themeda seeds were broadcast sown in a 


SIDA 19(1): 201-203. 2000 


202 SIDA 19(1) 


lawn dominated by crabgrass. The area was not fertilized or mowed during the growing 
season but did receive occasional supplemental water. 

Themeda seeds removed from the birdseed were apparently inert. However, some 
Vigna radiata var. radiata (Fabaceae) and Arthraxon hispidus (Poaceae) seeds germinated 
in the greenhouse. Three unidentified dicotyledons also germinated but died before 
reaching maturity. Both Vigna radiata and Arthraxon hispidus are weedy annuals from 
southeastern Asia that have been introduced into the United States (Kartesz 1999). 

One year after the initial collection, examination of the yard where Themeda was 
discovered did not find any recurrence of the grass. A floristic survey of numerous resi- 
dential yards where thistle seed was fed to birds also did not detect any exotic plants. 
Themeda may be unable to persist in Kansas, but in Louisiana it has flourished for more 
than 40 years in disturbed sites near cultivated fields (Reese & Landry 1985). 

India and Ethiopia are major producers of thistle for birdseed (Sharma 1982; Vin- 
cent & Cusick 1998), and Themeda seeds, as well as numerous other weedy species, are 
present in the harvested material. Although imported seeds are treated to prevent ger- 
mination, the process is fallible. Contamination or inconsistent sterilization procedures 
apparently were responsible for introduction of Themeda into Kansas. The incidence of 
viable seeds in thistle may be small, but birdseed represents a potential source for ad- 
ventive plants. 


ACKNOWLEDGMENTS 


Thanks to Larry and Sarah Heinken who initially observed the grass in their backyard, to 
Gail Wilson and Emily Benson for overseeing the greenhouse operations, to Gordon 
Shumway for purchasing the thistle seed, and to the Audubon Society members who 
consented to a floristic survey around their bird feeders. Support for this project was 
provided by the Kansas Agricultural Experiment Station and the National Science Foun- 
dation Long-Term Ecological Research Program. This paper is contribution no. 00-139-J 
from the Kansas Agricultural Experiment Station. 


REFERENCES 


Bairo, J.R. and J.W. THierer. 1985. Notes on Themeda quadrivalvis (Poaceae) in Louisiana. 
Iselya 2:129-137. 

Bor, N.L. 1960. Grasses of Burma, Ceylon, India and Pakistan (excluding Bambuseae). 
Pergamon Press, New York. 

Brown, C.A. 1945. Notes on additions to the flora of Louisiana. Proc. Louisiana Acad. Sci. 
9:4-13. 

Kartesz, J.T.1999.A synonymized checklist and atlas with biological attributes for the vas- 
cular flora of the United States, Canada, and Greenland. First Edition. |n:Kartesz, J.T.and 
C.A. Meacham. Synthesis of the North American Flora, Version 1.0. North Carolina 
Botanical Garden, Chapel Hill, NC. 


T 203 


Reese, W.D. and G.P. Lanory. 1985. Themeda quadrivalvis (L.) Kuntze (Poaceae) in Louisiana. 
Sida 11:99-102. 

SHARMA, S.M. 1982. Niger cultivation in India. Indian Farming 31:27-34. 

VinceNT, M.A. and A.W. Cusick. 1998, New records of alien species in the Ohio vascular flora. 
Ohio J. Sci. 98:10-17. 

WUNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. Univ. Press of Florida, Gainesville. 


204 SIDA 19(1) 


BOOKS RECEIVED 


Pup Precitt and Nancy Voukman. 1999. Landscapes in History: Design and Planning in 
the Eastern and Western Traditions, Second Edition. (ISBN 0-471-29328-8, hbk.). 
John Wiley & Sons, Inc.,605 Third Avenue, New York, NY 10158-0012.$89.95.844 pp. 
Numerous b&w photos. 


From the Cover:"This revised edition of Landscapes in History features for the first time new informa- 
tion—rarely available elsew! in the literature—on landscape architecture in India, China, South- 
east Asia, and Japan. It also expands the discussion of the modern period, including current North 
American Dieta, and design practices. 
Sania Highly oe ppok oa me a of landscape architecture and 
d e, the Middle East, Asia,and North 
America. It covers the manly cultural, political, technological, and philosophical issues influencing 
land use throughout history, focusing not only on design topics but also on the environmental 
impact of human activity. Landscape architects, urban planners, and students of these disciplines 
will find her 
The most comprehensive, in-depth, and up-to-date overview of the subject 
Hundreds of stunning photographs and design illustrations 
A scholarly yet accessible treatment, drawing on the latest research in archaeology, geogra- 
phy, and other disciplines 
The authors’ own firsthand observations and travel experiences 
on into the evolution of landscape architecture as a discipline 
Useful chapter summaries and bibliographies” 


C. Wayne Smit and J. Tom Coturen (eds.). 1999. Cotton: Origin, History, Technology, and 
Production. (ISBN 0-471-18045-9, hbk.). John Wiley & Sons, Inc.,605 Third Ave., New 
York, NY 10158-0012. $250.00.850 pp. Numerous b&w photos 


From the preface: The genus COSTE contains 49 species distributed throughout most tropical 
and subtropical regions of the world. The genu tains an amazing amount of diversity, ranging 
from herbaceous perennials to small trees. Seed coverings, or fibers, range from ie nonexist- 
ent to short, stiff, Gree brown hairs that aid in wide dispersal to long, fine white fibers that charac- 
terize highly improved cultivars. This monograph is truly about a miracle plant. It provides fibers for 
an array of textile suede including the clothes we wear, oil for human consumption, feed for 
livestock, and base chemicals for a plethora of industrial products.” Contents: 1) Origin and history, 
2) The cotton plant; 3) Production and production hazards; and 4) Processing and products. 


SIDA 19(1): 204. 2000 


NODULATING LEGUMES FROM THE 
TAHOE BASIN, CALIFORNIA 


Mohammad Athar James Harding 
Dept. of Environmental a Dept. of Environmental Horticulture 
University of California-Davi University of California-Davis 
Davis, CA 95616, U.S.A. Davis, CA 9 
atariq@cdfa.ca.gov jaharding@ucdavis.edu 
ABSTRACT 


A survey was conducted to determine the nodulating ability of 44 legume species from the Tahoe 
Basin. All were found to nodulate under natural soil eondiiaes Nodulation is s reported for the first 
time in nine species in five genera of Papilionoideae. These SI 

Genisteae, Thermopsideae, Trifolieae and Vicieae. Nodule ai shape and Poa are also de- 
scribed. The results pot eSing the morphology of nodular structures to classify legumes. The 
soils Ohne Tahoe Gas nerally eed eae Quins: survive and grow well there. 
Healthy plants h foli in nutri ts, suggest that they 
were nurtured by nitrogen-fixing aealiles Balcitaion of tegune Rhizobium symbiosis may help 
to improve nutrient poor soils and may provide refuge and quality browse for wildlife in the Tahoe 
Basin 


RESUMEN 


Los condicionantes g y bioclimaticos han permitido la existencia en el drea de Tahoe de 
un entorno unico. Nuestra investigacion se centrd en la determinacion de la capacidad de 
nodulacion de algunas especies de leguminosas en el area del lago Tahoe donde predominan los 
suelos pobres en nutrientes. Todas las especies estudiadas resultaron nodulantes en condiciones 
edaficas naturales.En nueve de pak penericcle tes a cinco géneros de Papilionoideae e incluidas 
en las tribus Gal ,Genistea Trifolieae y Vicieae, se describe por primera vez el 
proceso de nodulacion ee la coloracion, forma y frecuencia de las formaciones nodulares. 
Los resultados apoyan la estrecha relacion existente entre los diferentes grupos taxondémicos y la 
morfologia estructural de los nédulos. El desarrollo de poblaciones sanas con follaje exuberante 
colonizando estaciones con suelos pobres en nutrientes Pe ser er eae como bioindicador 
del proceso nodular de fijacion del nitrégeno. La utilizacidn de leguminosa-Rhizobium 
quizas ayude a mejorar la pobreza en nutrientes de los suelos del area de Tahoe asf como, en 
general, a proveer de mejores habitats para el incremento y la supervivencia de la vida silvestre en 
la cuenca del lago Tahoe. 


INTRODUCTION 


The California flora is rich in Fabaceae members or legumes; there are 69 genera and 491 
species found in California (Hickman 1993). Most legumes are indigenous while others 
are naturalized, often widely so (Witham 1994). Bacteria associated with legumes fix 
atmospheric nitrogen and are helpful in improving and maintaining soil fertility both in 
agriculture and natural ecosystems. This novel legume-Rhizobium association provides 
an ideal system for the study of plant-microbe interactions. Athar (1996a) studied nodu- 


SIDA 19(1): 205-210. 2000 


206 SIDA 19(1) 


lation in 66 Papilionoid species from Sacramento Valley, California and found that the 
majority of the species were abundantly nodulated under natural soil conditions. How- 
ever, a majority of the legumes found in California have not been examined for their 
nodulating ability under natural conditions. A survey was conducted to determine the 
nodulating ability of some legume species from the Tahoe Basin of California's Sierra 
Nevada Range (39° N, 120° W). Plants were collected from accessible areas within the 
Basin, ranging from Desolation Wilderness in the west to the Carson Ridge in the east, 
Alpine Meadows in the north,and Hope Valley and Carson and Luther passes in the south. 

Geologic and climatic forces have combined to form Tahoe's unique and varied 
environment (Blackwell 1997; Graf 1999). Tahoe's floral diversity can be attributed in part 
to its central location between plant communities of the Sierra Nevada western foot- 
hills, the western Great Basin desert, and the northern and southern Sierra Nevada. The 
variety of plant species can further be explained by the area's rapidly changing and var- 
ied habitats, which have contributed to the large number of Tahoe plants with limited 
geographic ranges (Graf 1999; Strong 1999). Although vegetation in the basin is mixed 
due to variations in temperature, precipitation and soil, coniferous forests dominate 
(Blackwell 1997; Strong 1999). 


MATERIALS AND METHODS 


Legume species growing under natural conditions were surveyed for their nodulating 
ability. Periodic field trios were made from late spring to early fall in various parts of 
Tahoe Basin. Observations were made as described previously (Athar 1996a). Legumes 
examined included indigenous and introduced herbs, shrubs and vines. At least five plants 
of each species were examined to minimize error. Legumes were identified by speci- 
mens of mature plants. Nodules were distinguished from other kinds of morphological 
modifications or pathogenic root malformation, and nodulation data were recorded. In 
some cases, nodule smears and nodule slices were prepared and examined under the 
microscope (Somasegaran & Hoben1994). 


RESULTS AND DISCUSSION 


Nodulation status was examined in 44 Papilionoid species growing under natural con- 
ditions in the Tahoe Basin. All the species observed were nodulated to various extents 
(Table 1). These results were compared with the available reports on nodulation (Aguilar 
et al, 1994: Allen & Allen 1981; Athar 1996a,b, 1997a,b; Athar & Mahmood 1990; Corby 
1988; de Faria et al. 1994; Nasim et al. 1998; Roggy & Prevost 1999). Nodulation is reported 
for the first time in nine species within five genera of Papilionoideae. These species are 
distributed in tribes Galegeae, Genisteae, Thermopsideae, Trifolieae and Vicieae. Joe 
Kirkbride, USDA, Beltsville, Maryland has been scanning the literature to create a database 
that records the presence or absence of root nodules. The database currently contains 
about 3,000 taxa reported with nodules and about 400 taxa reported without nodules. 
The results of the present investigation were also sent to him for comparison. Kirkbride 


CALIFORNIA 207 
Taste 1.Nodulation characteristics of some legume species from the Tahoe Basin, California. 
Nodulating Species! Sieur Frequency? Color Shape 
Repor 
Galegeae 
Astragalus bolanderi A. Gray B ++ Brown Elongated 
Astragalus gambelianus E. Sheld. A ++ Brow Elongated 
Astragalus tener A.Gra A + Reddish Elongated 
brown 
Astragalus whitneyii A. Gray B 44 Brown Elongated 
Genisteae 
“Lupinus affinis J. Agardh A ++ Pink Globose 
“Lupin nus albifrons Benth. A + Brown Globose 
_Lupinus ais © A + Brown Globose 
tiff astw B ++ Pink Globose 
Lupinus arbustus poteks ex ‘Lindl B ++ Pink Globose 
Lupinu us argenteus Pursh A + Brow Globose 
“Lupinus arizonicus (S. Watson) A + Reddish Globose 
S.Watson brown 
Lupinus brewri A. Gray B +++ Pink Globose 
Lupinus fulcratus Greene B +++ Pink Globose 
Lupinus grayi (S.Watson) S.Watson A +4 Pink Globose 
Lupinus lepidus Douglas ex Lindl. A ++ Pink Globose 
“Lupinus mi pus Sin A + Reddish Globose 
brown 
Lupinus polyphyllus Lindl. A + Brown Globose 
“Lupinus succulentus Douglas A + Brown Globose 
ex K. Koch 
Loteae 
“Lotus corniculatus L. A +++ Pink Elongated 
Lotus nevadensis (S.Watson) Greene A + Pink Elongated 
“Lotus unifoliatus Benth A ++ Pink Elongated 
Millettiea 
“Wisteria chinensis DC. A + Brown Globose 
Thermopsidea 
“Thermopsis acai Nutt. B +4 Brown Semi-globose 
deitelicass, 
lina L. A +++ Pink Elongated 
“Medicago pee L. A th Pink Elongated 
“Medicago sativa L A +44 Pink Elongated 
“Melilotus alba Medi A +++ Pink Elongated 
*“Melilotus indicus (L.) All A b++ Pink Elongated 
ee ee MnO GrOiger ay Torr. A +++ Pink Elongated 
Schreb. A ++ Pink Elongated 
"Trifolium dubs “i A + Pink Elongated 
"Trifolium h A ++ Pink Elongated 
Trifolium kin gil eee A +4++ Pink Elongated 
Trifolium onge Nutt B ++ Pink Elongated 
"Trifolium ns L. A +4 Pink Elongated 
"Trifolium ee L. A $44 Pink Elongated 


208 SIDA 19(1) 


Tasce 1. continued 


Nodulating Species’ Previous Frequency? — Color Shape 
Report? 

"Trifolium subterraneum L. A +++ Pink Elongated 
Trifolium variegatum Nutt. A ha Pink Elongated 
eiieae 

us nevadensis S.Watson B ++ Pink Elongated 
ee odoratus L. A +++ Pink Elongated 
"Vicia fa A ++ Pink Elongated 

Vicia hirsuta (L .) Gray A +++ Pink Elongated 
"Vicia sativa L. A 44 Pink Elongated 

"Vicia villosa Roth A ++ Pink Elongated 


‘Species are arranged alphabetically within genera, The nomenclature and tribal classification are 
as described by Polhill and Raven (1981). Author citations are quoted following instructions of 
Brummitt and Powell (1992). Introduced species are marked with an asterisk (*). 

*Nodulating status 

A= Ne orrae previously observ 

B = Nodu tae oe for the aang 

coe frequency 

ou icates ae nodulation (1 to 5 nodules per plant) 
ie ee moderate nodulation (6 to 10 nodules per plant) 
= Indicates abundant nodulation (more than 10 nodules per plant) 


compared the results with his global listing and has confirmed the new reports. The 
nodules observed in other species corroborate earlier studies (Allen & Allen 1981; Athar 
1996a, b; Athar & Shabbir 1997; Corby 1988). 

Nodules in most of these species were generally distributed on the main, as well as 
lateral roots, and were found in the 10 cm layer of the soil. Nodules of some legumes, 
particularly Lupinus spp., sometimes grew on the surface of the soil and were covered 
by a layer of damp litter. Similar observations were made by Athar (1996a) for some 
legumes from the Sacramento Valley. The size, number and color of the nodules varied 
for various species, as well as with the growth stage of the plants. Nodules varied from 
semi-globose to globose with streaked or smooth surfaces, to elongated and branched 
forms (Table 1). They occurred singly or as lobed structures. Nodule morphology strongly 
coincided with the descriptions of earlier workers (Allen & Allen 1981;Athar 1996a;Corby 
1988; Pueppke & Broughton 1999; Roggy & Prevost 1999; Somasegaran & Hoben 1994). 
Nodules were mostly pink or brown with reddish interiors. Nodule morphologies in le- 
gume species described by Corby (1988), showed that they were uniform at the tribal 
level and linked to their position in the evolution of the family. It is becoming clear that 
nodulation is a robust taxonomic character, both at the presence/absence level and at 
the structural/physiological level (Sprent 1999). The pioneer work of Corby (1988), link- 
ing the morphology of nodular structures to the taxonomy of the legumes, is still rel- 
evant, provided that it is coupled to anatomical analyses taking into account the modes 


CALIFORNIA 209 


of establishment of the symbiosis between the two partners (Pueppke & Broughton 
1999; Roggy & Prevost 1999; Sprent 1999). 

Attempts to isolate rhizobia or to test nitrogenase activity were not made for these 
nodulated legumes. However, healthy plants with lush green foliage growing in a nutri- 
ent-limited environment imply that they were nurtured by nitrogen-fixing nodules 
(Hartwig 1998;Somasegaran & Hoben 1994). This is supported by the generally accepted 
view that nodulated legumes have an internal regulatory system to allow them to ad- 
just nitrogen fixation to environmental conditions (Hartwig 1998; Sprent 1999). 

Lupinus (Lupines) were the most prevalent nodulated legumes in Tahoe Basin with 
14 species followed by Trifolium (clovers), Vicia (vetch) and Astragalus (locoweeds) with 
10,4 and 4 species each respectively (Table 1). The soils of the Tahoe Basin generally are 
nutrient poor, especially near the surface where drainage greatly exceeds the rate of 
chemical erosion from individual soil particles. All the legumes grew well in the nutrient 
poor environment of the Tahoe Basin. Physiological processes that plants undergo to 
survive in the Basin help in understanding Tahoe's vegetation ecology. Nutrient limita- 
tion plays an important role in plant distribution. Plants cope with nutrient deficiency 
through mutualisms with mycorrhizal fungi or through associations with bacteria capable 
of nitrogen fixation. Nitrogen fixation is a crucial component of many plant communi- 
ties in the Sierra Nevada Range, where nitrogen is ways in short supply. Exploitation of 
the legume-Rhizobium symbiosis in nutrient poor environment of the Tahoe Basin may 
help improve natural ecosystem, and provide refuge and quality browse for wildlife. 


ACKNOWLEDGMENTS 


Special gratitude is expressed to Joseph H. Kirkbride, USDA Beltsville, Maryland for his 
cooperation in checking the legume nomenclature and taxonomy, and comparing the 
results against his global database of legume nodulation. Thanks are due to Steve Shoenig, 
California Department of Food and Agriculture, Sacramento, California for helpful dis- 
cussion and suggestions; Ellen Dean and Grady Webster and the staff at the J.M. Tucker 
Herbarium, Davis, California for providing access to their herbarium facilities. 


REFERENCES 


AGuiLar, N.O,, F.C. Pitarcue, and M.O. Casano. 1994. Nodulation of legumes in Philippines. In: 
Jl. Sprent and D. McKey, eds. Advances in legume systematics. 5. The nitrogen factor. 
Kew, Royal Botanic Gardens. Pp. 25-31. 

Auten, O.N. and E.K. AtLen. 1981. The Leguminosae. A source book of characteristics, uses 
and nodulation. University of Winconsin Press, Madison. 

ATHAR, M. 1996a. Observations on nodulation status of some Papilionoid species of po- 
tential agricultural and forestry value from Sacramento Valley, California. Taiwania 
41:301-308 

ATHAR, M. 1996b. New nodulating legume species from natural ecosystem of Pakistan. 
Phytologia 80:385-388. 


210 SIDA 19(1) 


AtHar, M.1997a.A qualitative study of the nodulating ability of legumes of Pakistan. List 5. 
Acta Bot. Gallica 144:67-72. 

AtHar, M.1997b. Nodulation status of some legume species from Cache Valley and north- 
ern Utah. Phytologia 81:145-150. 

AtHar, M.and A.Mahmood.1990.A qualitative study of the nodulating ability of legumes 
of Pakistan. List 4. Trop. Agric. (Trinidad). 67:53-56. 

AtHar, M. and S.M. Shabbir. 1997. Nodulation characteristics of some of the forage and 
browse legumes. Phytologia 82:12-19. 

Biackwett, L. 1997. Wildflowers of the Tahoe Sierra. Lone Pine Publishing, Redmond, 
Washington. 

Brummitt,R.K.and C.E. Powett (eds.). 1992. Authors of plant names. Royal Botanical Gardens, 
Kew, UK. 

Corey, H.D.L. 1988. Types of rhizobial nodules and their distribution among the 
Leguminosae. Kirkia 13:53-124. 

Faria, S.M. de, H.C.de Loma, A.M. CarvatHo, V.F. Concatves, and J.I. Sprent. 1994. Occurrence of 
nodulation in legume species from Bahia, Minas Gerais and Espirito Santo states in 
Brazil. |n:J.1. Sorent and D.McKey, eds. Advances in legume systematics. 5. The nitrogen 
factor. Kew, Royal Botanic Gardens. Pp. 17-23. 

Grar, M. 1999. Plants of the Tahoe Basin: Flowering plants, trees, and ferns. California Na- 
tive Plant Society, Sacramento 

Hartwic, U.A.1998.The regulation of symbiotic N; fixation:a conceptual model of N feed- 
back from the ecosystem to the gene expression level. Perspect. Plant Ecol. Evol. Sys- 
tem. 1:92-120. 

Hickman, J.C. (ed). 1993. The Jepson manual: Higher plants of California. University of Cali- 
fornia Press, Berkeley. 

Nasi, M.,M.ATHaR, and S.M. SHaseir. 1998. Observations on some new nodulating legume 
species from Azad Kashmir. Phytologia 85:1 10-114. 

PotHitt, RM. and PH. Raven (eds.). 1981. Advances in legume systematics. Royal Botanic 
Gardens, Kew, UK. 

Pueppxe, S.G. and W.J. BRouGHTON. 1999. Rhizobium sp. Strain NGR234 and R. fredii USDA257 
share exceptionally broad, nested host ranges. Mole. Plant Microbe Interactions 12: 
293-318 

Roaey, J.C.and M.F. Prevost. 1999. Nitrogen-fixing legumes and silvigenesis in rain forest in 
French Guiana:a taxonomic and ecological approach. New Phytol. 144:283-294. 

SOMASEGARAN, P.and H. Hosen. 1994. Handbook for rhizobia: Methods in legume-Rhizobium 
technology. Springer-Verlag, New York. 

SprENT, J. 1999. Nitrogen fixation and growth of non-crop legume species in diverse en- 
vironments. Perspect. Plant Ecol. Evol. Syst. 2:149-162. 

StrONG, D.H. 1999. Tahoe: From timber barons to ecologists. University of Nebraska Press, 
Lincoln. 

Witham, C.W. 1994. Jepson prairie preserve plant check list. Jepson Prairie Docent Pro- 
gram, Davis, California. 


NOTEWORTHY PLANTS FROM NORTH FLORIDA. VII 


Loran C. Anderson 


Department Of Biological Science 
Florida State University 
Tallahassee, FL 32306-4370 U.S.A. 


ABSTRACT 


The following appear to be first reports for the state of Florida:Ajuga reptans,Aster la 
Cyperus retrofractus, Linum macrocarpum, and Rhynchospora thornei. Several saifonet to he dics 
of north Florida are also documented, and some species that were thought to be restricted in 
Florida to the western panhandle are noted for extreme northeastern Florida. 


RESUMEN 


Los siguientes taxa parecen ser las primeras citas para el estado de Florida: Ajuga reptans, Aster 
laevis var. concinnus, Cyperus retrofractus, Linum macrocarpum, y Rhynchospora thornei. Se 
documentan también varias adiciones a la flora del bole ae Honea: y algunas dusaan cis se 
pensaba que estaban restringidas en Florida ala franja 
nordeste de Florida. 


IN previous installments of this series (Anderson 1984, 1986, 1988, 1989, 1991, 1995),"north 
Florida” was defined as the Florida panhandle west of the Suwannee River. Recent bo- 
tanical surveys, particularly in Clay, Nassau, and Putnam counties of northeastern Florida, 
have prompted me to expand the area of coverage to include that portion of the state 
from Putnam County northward. Particularly interesting sites were found at Black Creek 
Ravines Conservation Area just E of Middleburg in Clay County and Ralph E. Simmons 
Memorial State Forest just E of Boulogne in Nassau County. Exotics that appear to be 
adventive or naturalized are included. Nomenclature generally follows Wunderlin (1998). 
Herbarium specimens are at FSU unless noted otherwise. These findings will update 
species distributions as mapped by Wunderlin et al. (1999). 


TAXA NEW TO THE AREA 


Ajuga reptans |.—Leon Co.: extensively naturalized in Tallahassee lawns, 7 May 1999, 
Anderson 18620; new to Florida. 

Aster laevis L. var. concinnus (Willd.) House.—Okaloosa Co.: N side of Karick Lake, 11 
Oct 1996, Anderson 17191;Santa Rosa Co.:shrubby thicket around lake at Camp Paquette, 
6 mi NNE of Munson, 27 Oct 1995,Anderson 16036, margin of Krull Lake, W of Sweetwater 
Creek, 11 Oct 1996, Anderson 17157; new to Florida. 

Duplicates identified by Almut G. Jones (pers. comm.), but | think they may repre- 
sent a new taxon. These Florida samples have disk corollas 3.7-4.8 mm long, whereas 
disk corollas from four samples of A. /aevis (fram neighboring states) ranged from 5.2-6.0 
mm. The Florida disk flowers have long lobes (1.3-1.6 mm long) and short throats, whereas 


SIDA 19(1): 211-216. 2000 


212 SIDA 19(1) 


the A. laevis samples | measured had short corolla lobes and long throats. A large lobe/ 
throat ratio is characteristic of A. lateriflorus (L.) Britton, but that species is cespitose or 
with short, stout rhizomes and a branching paniculate inflorescence, whereas the Florida 


plants have long, slender rhizomes and more racemiform inforescences. 
Chenopodium murale |.—Leon Co.: cracks in sidewalk along Sixth Avenue in Tallahas- 
see, 6 Mar 1999, Anderson 18268; new to Florida panhandle. 

Conobea multifida (Michx.) Benth—Leon Co.: railroad crossing at Capitol Circle NW in 
Tallahassee, 2 Oct 1992, Godfrey 84404, same locality, 6 Oct 1999, Anderson 19056; new to 
north Florida and second county of record for Florida. Many authors (e.g., Gleason & 
Cronquist 1991) list this taxon as Leucospora multifida (Michx.) Nutt. 

Cyperus retrofractus (L.) Torr. in A. Gray.—Jackson Co.: Florida Caverns State Park with 
Desmodium ochroleucum M. A. Curtis and D. rotundifolium DC., 10 Oct 1998, Anderson 
18188; new to Florida. Clewell (1985) listed C. retrofractus for the Florida panhandle, but 
the plants to which he was referring are now considered C. plukenetii Fernald; see Carter 
and Jarvis (1986) for clarification. 

Linum macrocarpum Rogers.—Bay Co.: pine flatwoods near Long Beach, 7 Jul 1963, 
Godfrey 62966; Franklin Co.: Apalachicola National Forest, E of Rte 65,5 Jul 99, Anderson 
18810;new to Florida. The Bay County specimen was annotated as L. floridanum (Planch.) 
Trel. by Rogers, but later Bruce Sorrie noted its capsules were 3.4—4.6 mm long and iden- 
tified it as L. macrocarpum. The Franklin County specimen was immature, with capsules 
3,3-3.5 mm long and shrunken seeds 2.5—2.6 mm long. This species was previously known 
only from the 1897 type collection (in swamps, Spring Hill) in Mobile Co., Alabama (Rogers 
1963).Anderson 16859 adds a second locality (east of Bayou LaBatre) in Mobile County, AL. 
Lipocarpha aristulata (Cov.) G. Tucker—Putnam Co.:wet depression beside railroad, 1 
mi N of Palatka, 18 Sep 1997, Anderson 18119,same locality, 22 Oct 1998, Anderson 18500; 
new to north Florida. Kral (pers. comm.) prefers Hemicarpha aristulata (Cov.) Smyth. 
Mecardonia procumbens (Mill.) Small—Putnam Co.: frequent in border of lawn, S of 
rte 100, W edge of Palatka, 16 May 1997, Anderson 17444, same locality, 19 Sep 1997, 
Anderson 18152;new to north Florida. 

Rhynchospora thornei Kral —Jackson Co.:marshy area bordering lake 1 mi £ of Marianna, 
27 Sep 1957, Kral & Godfrey 5996, edge of small pond 2 mi NW of Grand Ridge, 10 Oct 
1997, Anderson 18203; Putnam Co.: wet roadside depression 1 mi N of Palatka, 18 Sep 
1997, Anderson 18133; new to Florida. The species was previously known only from the 
type locality in Baker County, Georgia (Kral 1977). The Kral and Godfrey specimen (origi- 
nally labeled “Rhynchospora divergens Chapm.?") was annotated as R. thornei by Kral in 


Taraxacum laevigatum (Willd.) DC —Putnam Co.: lawn weed at St. Johns River Com- 
munity College in Palatka, 15 Jan 1999,Anderson 18550;new to north Florida and second 
county of record for Florida. 

Thunbergia alata Bojer ex Sims.—Leon Co. escaped from cultivation and naturalized 
in Tallahassee, 22 Jun 1998, Anderson 18388; new to north Florida. 

Thymophylla tenuiloba (DC.) Small—Leon Co.: escaped from cultivation and natural- 


ANDERSON, NOTEWORTHY PLANTS FROM NORTH FLORIDA 213 


ized in Tallahassee, 7 Jul 1999, Anderson 18817;new to north Florida. 

Xanthosoma sagittifolium (L.) Schott—Leon Co. naturalized (vegetative) in swampy 
woodland by Ochlockonee Rd, NW of Tallahassee, 1 Nov 1996, Anderson 17247, same 
locality (flowering), 20 Sep 1999, Anderson 19053; new to north Florida. 


ADDITIONAL RANGE EXTENSIONS 


Abutilon theophrasti Medik—Leon Co.: edge of flood-control pond on Call Street in 
Tallahassee, 7 Jul 1999, Anderson 18816; new county of record and second collection for 
north Florida. 

Aureolaria virginica (L.) Pennell—Clay Co.: steephead ravine at Black Creek Ravines 
Conservation Area, 10 Jun 1999, Anderson 18691; new to northeast Florida and first col- 
lection in Florida east of Ochlockonee River. 

Boltonia asteroides (L.) L'Her—Nassau Co.: bordering St. Marys River at Hwy 17 bridge, 
9 air mi NW of Yulee, 5 Oct 1985, Anderson 8911: new to northeast Florida. 

Callisia repens (Jacq.) L—Leon Co.:escaped from cultivation and naturalized along road- 
side ditch in Tallahassee, 20 Oct 1998, Anderson 18491;second county of record for north 
Florida. 

Carex crebriflora Wiegand.—Clay Co.:deep shade of swampy floodplain of Black Creek, 
Black Creek Ravines Conservation Area, 10 Jun 1999, Anderson 18674; new to northeast 
Florida. 

Carex styloflexa Buckley —Clay Co.:deep shade of floodplain of Black Creek near junc- 
tion of the two forks, Black Creek Ravines Conservation Area, 10 Jun 1999, Anderson 18687: 
new to northeast Florida. 

Chasmanthium latifolium (Michx.) Yates —Nassau Co.:shaded banks of St.Marys River 
at Simmons State Forest, 25 Jun 1999, Anderson 18788, 8 Jul 1999, same locality, Anderson 
18842; new to northeast Florida (first collection in Florida east of Ochlockonee River). 
Croton willdenowii G.L.Webster—Bay Co.:abundant in white sand of depression just E 
of Enfinger Rd, S of rte 20 (ca. 2 mi W of Econfina Creek), 27 Oct 1999, Lisa & Ed Keppner; 
second county of record for the state of Florida. This species was previously known as 
Crotonopsis elliptica Willd. Pennell (1918) stated that, in the south part of its range, it oc- 
curred “mostly inland,on granitic rocks.” Here it occurs in loose white sand, and the other 
known occurrence in Florida is on citronelle sandstone outcrops at the summit of Rock 
Hill in Washington County. 

Cyperus pseudovegetus Steud.—Nassau Co.:seepage slope, Simmons State Forest, 25 
Jun 1999, Anderson 18764; second county of record in Florida east of Leon County. 
Desmodium rotundifolium DC.—Jackson Co.: Florida Caverns State Park with D. 
ochroleucum and Cyperus retrofractus, 10 Oct 1997,Anderson 18185;third county of record 
for Florida. 

Fimbristylis decipiens Kral—Leon Co.:lawn and open weedy slopes above pond E of 
Mary Ellen Drive in Tallahassee, 8 Oct 1992, Anderson 14062, same locality, 25 Oct 1997, 
Anderson 18241. Listed by Clewell (1985) for Florida panhandle, but no specimens seen; 
these collections confirm its presence. 


214 SIDA 19(1) 


Hyperi gy tk Englem & A.Gray.—Nassau Co.:seepage slope, Simmons State 
Forest, 25 Jun 1999, Anderson 18777,new to northeast Florida and first collection in Florida 
east of Ochlockonee River. 

Kyllinga squamulata Thonn.ex Vahl.—Leon Co.: locally common at Tallahassee South- 
east Farm, S of Tram Rd, 25 Sep 1995, Horne 773, same locality, 26 Sep 1995, Anderson 
15899; new to Florida panhandle. 

Leersia lenticularis Michx.—Nassau Co.:along stream in Plummer Swamp,4 air mi WSW 
of Yulee, 24 Jul 1999, Anderson 18926; new to northeast Florida and first collection in 
Florida east of the Ochlockonee River. 

Lithospermum incisum Lehm.—Putnam Co.:dry sand of open oak woodland in Palatka, 
27 Mar 1998, Anderson 18926; second county of record for northeast Florida. 
Murdannia keisak (Hassk.) Handel-Mazz—Leon Co.: under Meridian Rd bridge W of 
Lake lamonia, 18 Oct 1993, Anderson 14590, 27 Oct 1999, Anderson 19082, shaded streamlet 
at Elinor Klapp-Phipps Park, N edge of Tallahassee, 23 Sep 1996, Anderson 17068; second 
county of record for Florida panhandle for this recently introduced, aggressive weed. At 
the Lake lamonia site, it appears to be displacing Polygonum meisnerianum Cham. & 
Schlecht., which is an endangered species in Florida (Coile 1998). 

Oxycaryum cubense (Poeppig & Kunth) Lye.—Bay Co.: frequent along edge of drain- 
age ditch,W of Rte 389 at Girl Scout Camp, 5 Jun 1999, Keppner 1120.Reported (as Scirpus 
cubensis Poeppig & Kunth) by Clewell (1985), but no vouchers from the Florida pan- 
handle were found in major Florida herbaria (Wunderlin et al. 1999). This collection docu- 
ments its presence; the species has also been found recently in Alabama and Georgia 
(Bryson et al. 1996). 

Paederia foetida L.—Duval Co.: alley behind Clemson Rd near University Blvd in Jack- 
sonville, 13 Nov 1998, Podris s.n.; Franklin Co.: St. George Island, 13 Apr 1995, Jubinsky s.n.; 
Leon Co.:;woodland along Crestdale Ln, NE of Tallahassee, 30 Sep 1997, Jubinsky s.n.;new 
counties of record for this Class | weed (FLEPPC 1999.) 

Paspalum conjugatum Berg.—Clay Co.: mesic woodland bordering St. Johns River, 
Bayard Point State Preserve, 17 Aug 1999, Anderson 18955; new to northeast Florida. 
Plantago major L.—Nassau Co.: beside rte 2 at St. Marys River bridge, W of Callahan, 24 
Jul 1999, Anderson 18931;second county of record for northeast Florida. 

Plantago rugelii Decne —Jackson Co.: Pine Shelter service road, Florida Caverns State 
Park, 10 Oct 1997, Anderson 18190; second county of record. Additional sites in Leon Co.: 
Anderson 18078, 18389; first reported for this county (and the state) by Anderson (1986). 
Platycladus orientalis (L.) Franco—Marion Co.: naturalized, several trees of differing 
ages in pine-oak woodland $ of Ferry Rd,E of rte 19 in Ocala National Forest, 23 Oct 1998, 
Anderson 18507;second record for state of Florida (not collected since 1905,see Wunderlin 
1998). 

Polygonum lapathifolium |.—Nassau Co.: edge of cultivated field, ca. 4 air mi NE of 
Boulogne, 8 Jul 1999, Anderson 18852; new to northeast Florida. 

Pycnanthemum flexuosum (Walt.) B.S.P—Nassau Co.: semishade of seepage slope, 


ANDERSON, NOTEWORTHY PLANTS FROM NORTH FLORIDA 215 


Simmons State Forest, 25 Jun 1999, Anderson 18764,same general area in full sun, 18 Aug 
1999, Anderson 19009; new to northeast Florida and first collection in Florida east of 
Suwannee River. 

Rhynchospora macrostachya Torr.—Nassau Co.: shallow pond on seepage slope, 
Simmons State Forest, 25 Jun 1999, Anderson 18762. Reported for the state in 1933 by 
Small (but not by Godfrey and Wooten 1979 or Wunderlin 1998), but this is apparently 
the first specimen documenting its presence (G. Moore, pers. comm,). Another horned 
beaksedge from two sites at Simmons State Forest (Anderson 19007, 19014) is puzzling in 
having achenes with short bristles typical for R. corniculata (Lam.) A.Gray but with stout 
rhizomes as in R. careyana Fernald. 

Schisandra glabra (Brickell) Render—Leon Co.:mesic woodland along Trillium Court in 
Tallahassee, 1 May 1999, Anderson 18617;new county of record for this endangered spe- 
cies (Coile 1998), 

Scirpus koilolepis (Steud.) Gleason.—Nassau Co.:;campground at Cary State Forest, just 
NE of Bryceville, 1 May 2000, L.C. Anderson 19196; first report for Florida east of the 
Apalachicola River. 

Spermacoce glabra Michx.—Madison Co: steep limestone slopes of Withlacoochee 
River at Blue Spring, 16 Jul 1999, Anderson 18876; first collection in Florida east of 
Ochlockonee River. 

Spermacoce verticillata | —Putnam Co.: roadside 1 miN of Palatka, 18 Sep 1997, Ander- 
son 18122, same general area, 5 Dec 1997, Anderson 18263; third county of record for 
northeast Florida. 

Vaccinium tenellum Aiton.—Clay Co.: overgrown seepage slope, Bayard Conservation 
Area, SE of Green Cove Springs, 10 Dec 1998, Slaughter 10717, sandhill seepage at Black 
Creek Ravines Conservation Area, 10 Jun 1999, Anderson 18665; St. Johns Co.: powerline 
corridor, Moses Creek Conservation Area, 2 mi WNW of Crescent Beach,9 Apr 1998, Ware 
s.n. These collections confirm its occurrence in Florida. Vander Kloet (1988) noted a single 
1943 collection from Clay Co.and implied the species may no longer be found in Florida; 
the same situation was noted in 1996 (Luteyn et al.). Surprisingly, Wunderlin et al. (1999) 
listed the species for over 20 counties in north Florida. The discrepancy between this 
wide distribution and all previous reports may arise from confusion of this species with 
other elements of the genus. 


ACKNOWLEDGMENTS 


My field work in northeastern Florida was sponsored by a contract from the St. John’s 
River Water Management District. Ed and Lisa Keppner are thanked for their intensive 
field work in the Panama City area. 


REFERENCES 


ANDERSON, L.C. 1984. Noteworthy plants from north Florida. Sida 10:295-297. 
. 1986. Noteworthy plants from north Florida. ll. Sida 11:379-384. 


216 SIDA 19(1) 


1988. Noteworthy plants from north Florida. lll. Sida 13:93-100. 
_1989.Noteworthy plants from north Florida. IV. Sida 13:497-504. 
_1991.Noteworthy plants from north Florida. V. Sida 14:467-474. 

_ 1995. Noteworthy plants from north Florida. VI. Sida 16:581-587. 

Bryson, C.T., J.R. MacDonato, R. Carter, and S.D. Jones. 1996, Noteworthy Carex, Cyperus, 
Eleocharis, Kyllinga, and Oxycaryum (Cyperaceae) from Alabama, Arkansas, Georgia, 
Louisiana, Mississippi, North Carolina, Tennessee, and Texas. Sida 17:501-518. 

Carter, R., and C.E. Jarvis. 1986. Re-evaluation and leptotypification of Scirpus retrofractus L. 
Rhodora 88:451—-455. 

Ciewett, AF. 1985. Guide to the vascular plants of the Florida panhandle. Florida State 
University Press/University Presses of Florida, Tallahassee. 

Cote, N.C. 1998. Notes on Florida's endangered and threatened plants. Florida Depart- 
ment of Agriculture & Consumer Serv., Division of Plant Industry—Botany Section 
Contrib. No. 38, Gainesville. 

FLEPPC. 1999. Florida's most invasive plants list. Florida Exotic Pest Plant Council. http.// 
www/fleppc.org. 

Gueason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada, 204 ed. New York Botanical Garden, Bronx. 

Goprrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United 
States. Monocotyledons. University Georgia Press, Athens. 

Krat, R. 1977. A new species of Rhynchospora (Cyperaceae) from southwestern Georgia. 
Sida 7:42-50. 

Luteyn, J.L., W.S. Jupp, S.P. VANDER KtoeT, LJ. Dorr, G.D. Wattace, K.A. Kron, PF. Stevens, and S.E. 
Ciemants. 1996. Ericaceae of the southeastern United States. Castanea 61:101-144. 
PENNELL, FW. 1918. Notes on plants of the southern United States. lV. Bull. Torrey Bot. Club 

45:477-482. 

Rocers, C.M. 1963. Yellow flowered species of Linum in eastern North America. Brittonia 
15:97-1 

Smatt, J.K. 1933. Manual of the southeastern flora (1972 facsimile reprint). Hafner, New 
York. 

Vanoer Ktoet, S.P. 1988.The genus Vaccinium in North America. Research Branch Publ. 1828, 
Agriculture Canada. 

WuNberLN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, 
Gainesville. 

_B.F. Hansen and E.L Brioces. 1999. se of Florida vascular plants. http:// 


+| + 


www. Weedumebiome ts/atlas/atlas.Atm 


WHO NAMED EUTHAMIA (COMPOSITAE: ASTEREAE) 
AND WHEN? 


John L. Strother 


/ fi act ‘ 
niversity PTELOCEIUT 


1001 Valley Life Sci fences Bui ai ing #2465 
University of California 
Berkeley, a See U.S.A. 


Nesom (1999) and Gandhi (1999) reviewed author attributions and dates for the generic 
name Euthamia and for binomials in Euthamia. They expressed slightly different inter- 
pretations; here is a third interpretation: 

The first known use of Euthamia as a botanical name was by Nuttall (1818). Nuttall 
wrote of his Euthamia,"A subgenus, or rather genus, reciprocally allied to Solidago and 
Chrysocoma' (Fig. 1).Only a taxon of generic rank can be said to be “reciprocally allied’to 


oO o ~ 
162 SYNGENESIA, SUPERPLUA. SYNGENESIA. SUPERFLWA. 163 
44. Virgaurea, 45. vimine * puberula, ein sim- * 
ple and ae somewhat eee leaves Junceolate, 561. *BRACHYR 
entry De co each side minutely pUBSsCeN ‘stented. - Calix elinde. ic-ov mate cosy sae nae scales 
ithe emity emes 
st ity, Wa a nee ate d. Pee ore ut 5; discal 
axillary §§ 
of the oe linear. peas acute; AN gated, about orets 05. plc naked Cpa short 
10. Ha os the sandy nei of New Jersey, near Am- and eet leaflet 8, persistent, 
Boy, ol: Saeent brownish, 1 ta tof eS Beh, ainpleyand Vegetation almost ae ae to that of Eushania 
de gree; racemes shorter than the low er leaves, collected sa 
into a leafy spike, rays conspicu- . Euthamiz. Solidago Sarothre. Pu. 2. 
ous and of The whole aupect of au Perennial. Stems nu tely 
the pla ant is that of SS veer, the HaHa gh nd up- und scabrous, 6 to 12 inches oaEh ‘faiticiately een 
snl carpi sce: Leaves id esl ee Narrow linear, entire, 
a. multiradiata. 48. elata. 49. riyida. The largest punctate, and vant iator on the argin; br: anchlets PIE o- 
North Am tomous. Flower: j mall 
and yellow. | Scales of iy ealix subearinate, a ie 
*Eurnamia. t Calix cylindric-ovate, closely im- lutinous. Rays 
iieuted, scales ch emt Radial florets 10 to 5, entire, Sas pa slong as the eal: *Discal florets the 
20, very small, yel Receptacte setose. Papi- sam ah number, becca thed. oe ee ae 
. : ersely conic, pubescent; pappus us, about the 
jus simple, Seed villous length of the see » 5 to 8-parted, leaflets unequal, linear 
ound tery sl nu y bral ached, teavea nar- and subacute. Has. On the ~ hills of the Missouri, 
e, longitudinally nerved; flowers ter- from the Arikarees to the Mountains?) The whole plant 
minal, een ‘glomeruli fastigiate, “corymbose. Ca- possesses a strong balsamic but Niiacieeisle scent, and 
lix resinosely viscid. A subgenus, or a genus, reci- ig used oe by the aborigines, operating power 
procally a allied to saan a and Cirysoce fully as a diuretic. 
50. zraminifolia ifolia, Lin. Solidu- 
go lanceolata Aiton. Ss ib 1 "E an 562, DONIA. 
gular, under side of the Icavea Calix hem faherical: imbricated, squarrose 


minutely hispi id; leaves eee) linear, entire, J to 5. 7 1S] 
eee natn serious rays 15 to 20, minute, scarcely and glutinous. Radial flore esta 
Ha rginia 


30 to 
35, yellow). Jteceptacle naked, scrobiculate. 


m Canada to ue yello 
ce: cof 8. “emudfaia, Pu. 2. p. 540. Stem low Pupjus setaceous, sete 3 ur 4, deciduous, some- 
and branched, angular, a as well as | 
the leaves: smoot tea aves narrow vy linea ary nume By mar- hat paleaceous. 

Suffruticose, or biennial; leaves serrated, resinosely 
resinous atoms, ar ee a "reds. rays about 10, dis- pene flowers terminal, fustigiate. Seeds obovate, 
tinetly. exse red. w Jer rsey to Florida. oth. 

AX a of u ee 2 1 De sge lage si Px. Oss. Biennial or suf- 
vlidago is exclusively 2 a North Amcrican genus, with ieitcees "Bian 3 or 4 fee high, erect or oe 
o “exception o of 5 5 es 6 species in Europe, and 2 near vie or Purp lists branches fasta few-A nai 
Canton in China arborescent species of St. Helena muli owered b- 
and New Zealand ‘i ak a be excluded from this tus, an leaves oblon ng, acute, serrate, amp asic 
genus, if ever carefully examine oath and reticulately veined, glandularly punctate. 


+ In allusion to the crowding of the flowers, + From Bg%xX 5; short, and #X¢4l, a chaffy acale; the pap- 
pus being short and palcaceous. 


Fic. 1. Pages 162 and 163 f | 2 of Nuttall’s The G f North American Plant 


SIDA 19(1): 217-218. 2000 


218 SIDA 19(1) 


two other genera and a taxon cannot be both a subdivision of Solidago and, at the same 
time,"reciprocally allied to Solidago and Chrysocoma." Nuttall (Fig. 1) associated the epi- 
thets graminifolia and tenuifolia with the generic name Euthamia and, in his protologue 
of Brachyris (Fig. 1), Nuttall used the binomial Euthamia tenuifolia. 

Elliott (1823), Cassini (1825), and de Candolle (1836) treated Nuttall’s Euthamia as a 
generic name published in 1818 and Elliott and de Candolle treated Nuttall’s E.graminifolia 
and E. tenuifolia as published in 1818. 

Regardless of typographic lapses or errors or inconsistencies of numbering and/or 
positioning in production of Nuttall’s Gen. N.Amer.pl. (see Gandhi 1999; Nesom 1999; and 
papers cited by them), citations for Nuttall’s Euthamia names should be: 

Euthamia Nutt.,Gen.N. Amer. pl. 2:162. 1818. 
Euthamia graminifolia (L.) Nutt.,Gen.N.Amer.pl.2:162.1818.Basiony:Cl minifolia 
Sp. pl. 841.1753. 
Euthamia tenuifolia (Pursh) Nutt. Gen.N. Amer. pl. 2:162. 1818. Basionym: Solidago tenuifolia 
Pursh, Fl. Amer. sept. 540. 1813 [1814]. 


ACKNOWLEDGMENTS 
| thank K.N. Gandhi, G.L. Nesom, P.C. Silva, and A.R. Smith for discussions on Euthamia. 


REFERENCES 


Canpotte, A.-P. be. 1836. Solidago. In: Prodr. 5:330-342. Treuttel and Wurtz, Paris. 

Cassini, H. 1825. Euthamia. In: F. Cuvier, Dict. sci. nat., ed. 2.37:471. Paris. 

E.uort, S. 1823. Solidago. In: Sketch bot. S. Carolina 2:368-392. J. R. Schenck, Charleston. 
Reprint: 1971. Hafner Publishing Co., New York. 

GANoHI, K.N. 1999.Nomenclatural novelties for the Western Hemisphere plants-—ll. Harvard 
Pap. Bot. 4:295-299. 

Nesom, G.L. 1999. Review of early nomenclature in Euthamia (Asteraceae: Astereae). Sida 
18:1009-1018. 

Nuttatt, T. 1818. Euthamia. In:Gen.N. Amer. pl. 2:162. D. Heartt, Philadelphia. Reprint: 1971. 
Hafner Publishing Co., New York. 


JOSEPH ANDORFER EWAN 
OCTOBER 24, 1909 - DECEMBER 5, 1999 
A MEMOIR 
Anne S. Bradburn 


Tish. aren roy an OS L a 
PUICATIC UEMVETOICTY PICIRUGTIUIET 
Dept. of Ecology and Evolutionary Biology 
Tulane University 
New Orleans, LA 70118-5698, U.S.A. 


The lights were back on, projection equipment was being put away, when aman with a 
neat white goatee, twinkling eyes, and the most active eyebrows one can imagine said 
“That's such a good show, you need to know all the plant names. | can help youl” After 
dinner the following week, Professor Ewan, who would become my mentor and friend, 
did just that; adding names to flowers in the slide show used in a seven-year, finally- 
successful effort to have two small Mississippi barrier islands added to the National Wil- 
derness System. By the time the campaign was over, | had taken all the botany courses 
offered at Tulane, wound up with a master’s degree, and found a new career. 

For over thirty years, Professor Ewan touched the lives and minds of Tulane stu- 
dents. None of the hundreds of students who enrolled in biology 431 (Plant Systematics 
in the catalogue but really a course in “Plant Appreciation”) could ever walk across the 
campus in the same way. Their eyes had been opened to a world of wonder. Music, art, 
and his beloved history were woven into plant biology. You never missed a Ewan lecture, 
nor forgot one! 

For one whose focus would be the history of science, Joseph Ewan left a remarkably 
thin personal paper trail. The facts of his life are scant and not easy to find. Born in Philadel- 
phia, educated in Los Angeles, he received an A.B. degree in botany from the University of 
California, Berkeley in 1934. In 1935, he married Ada Nesta Dunn, a fellow student who 
shared many of his interests and his life for the next 64 years. They had three daughters: 
Kathleen, Dorothy, and Marjorie and five grandchildren. He was an instructor at the Univer- 
sity of Colorado (1937-1944), botanist with the Foreign Economic Administration (1944— 
1945), Assistant Curator, Smithsonian Institution (1945-1946) and Associate Botanist, 
Bureau of Plant Industry, U.S. department of Agriculture (1946-47). In 1947, he came to 
Tulane as an Assistant Professor, moved steadily up the academic ladder, and in 1972 was 
named the Ida Richardson Professor of Botany, a chair he held until 1977 when he was 
appointed Emeritus Professor. For nearly forty years, Tulane students, faculty,and staff mem- 
bers were exposed to his mind, his vast knowledge, and his enigmatic wit. 

Along the way, Ewan was a Guggenheim Fellow (1954), a National Science Founda- 
tion Fellow (1959-61),a Smithsonian Regents’ Fellow (1984), and held visiting professor- 
ships at the Universities of Hawaii and Oregon, and at the Ohio State University. 


os: 


SIDA 19(1): 219-222. 2000 


220 SIDA 19(1) 


He received the Founders Medal from the Society for the History of Natural History, 
London in 1977, the Elosie Payne Luquer Medal from the Garden Club of America in 
1978, and with his wife Nesta Dunn Ewan, the Henry Shaw Medal from the Missouri 
Botanical Garden in 1994. Ewan was an elected Fellow of the Linnean Society of London. 
The Botanical Society of America awarded him a Certificate of Merit in 1989. The College 
of William and Mary and Tulane University awarded him honorary doctorates. 

“A bite of immortality” is the Ewanian concept of publication. His own first “bite’ at 
age 19, was A Report on the California Black Railin Los Angeles County published in Condor. 
His early botanical interests focused on ferns, and he was President of the American Fern 
Society in 1951. Taxonomic studies on Delphinium and other Gentianaceae followed 
but his interest had always been captivated by biogeography, bibliography, and history 
of biology and by the time | became his student in the early 1970's, history had com- 
pletely replaced taxonomy in his research. Questions to Ewan were most frequently an- 
swered by questions and one left his office burdened with books. He believed books 
were to be used, and was always generous with his personal library. As a new graduate 
student, | remember being awe struck when he handed down from above his rickety 
desk, a beautiful vellum copy of Bauhin’s Prodromus with the disclaimer that it was only 
the second edition from 1671. Back in my cubicle, | was afraid to open it! 

Other samples from his “bites” include such tempting titles as:"From Calcutta and 
New Orleans, or tales from Barton's greenhouse’Roots of the California Botanical Soci- 
ety””Who Conquered the World? or four centuries of exploration in an indehiscent cap- 
sule” and “The Botany of Cook's Voyages; or around the world on six shillings a day.” 

Then there are his many contributions to The Dictionary of Scientific Biography 
where ones finds, among others, sketches of George Engelmen, Albert Spear Hitchcock, 
Elmer Drew Merrill, Frederick Pursh, and the irascible Constantine Samuel Rafinesque. 
His introductions to the Classica Botanica Americana series are classics themselves. As a 
book reviewer, Ewan was in constant demand. His writings into his venue were always 
tantalizing. As in"A good beginning is the true story of nine pigs driven 71 miles by a peon 
carrying a pine torch’ which certainly tempts one to dip into Archie Carr's “High Jungles 
and Low." Or"How can you know the gardens of the deep south without this book close 
to the hammock?" and the fact that its author is “not frightened by information” makes 
one want to dash out to acquire Charlotte Seidenberg's The New Orleans Garden." 

Perhaps more meaty “bites” are his books. Rocky Mountain Naturalists (1950), its fol- 
low-up, Biographical Dictionary of Rocky Mountain Naturalists (1981), and John Banister 
and His Natural History of Virginia (1970) the later two co-authored by his wife, Nesta Dunn 
Ewan.In 1969 Ewan edited A Short History of Botany in the United States and contributed 
the Calendar of Events and chapters on Early History and Plant Geography. Here he graphi- 
cally illustrated the “Role of the Bartrams in natural History” for Ewan believed William 
Bartram’s contributions central to the development of botany in the United States. His 
William Bartram and His Botanical and Zoological Drawings was published by the Ameri- 
can Philosophical Society in 1968 and the Ewans' monumental biography of Philadel- 


BRADBURN, JOSEPH ANDORFER EWAN 221 


rt 


Joseph Andorfer Ewan 
October 24, 1909 — December 5, 1999 


222 SIDA 19(1) 


phia botanist Benjamin Smith Barton will be published posthumously by the Missouri 
Botanical Garden. 

In 1986, the Ewans moved from Tulane. The Missouri Botanical Garden had bought 
his prodigious library and given them a new home. There they were housed in the old 
museum building and for the first time the Ewans had spacious working conditions 
with large double desks for each of them and his 4,500 books carefully catalogued and 
shelved within easy reach. In announcing the move, Peter Raven, Director of the Garden, 
said “The Joseph Ewan collection is an extremely valuable acquisition for the Garden. 
The historical significance of the collection, its relevance to the work we do, and the 
respect Joseph Ewan commands in the scientific community make this announcement 
one of the most exciting I've made while at the Garden.” 

We a Tulane are left with his name on the door, lots of happy memories, and his 
herbarium now over 110,000 specimens. Just last week we received a request from Swit- 
zerland for Tulane’s holdings of Macrocarpaea, a genus monographed by Ewan in 1948. 
As | compose on the Pentium, in my mind's ear, | can still hear the clicking of his manual 
typewriter which sounded in the same room for so many years. 

The 1989 Ewania:The Writings of Joe and Nesta Ewan lists 358 "bites of immortality.” 
The following ten years produced many other “bites.” Surely a veritable feast for genera- 
tions of botanists, natural historians, and biogeographers in the generations to come. 


BOOK REVIEWS 223 


BOOK REVIEWS 


Joe Liccio and Ann O. Liacio (David H. Riskind, Scientific Advisor). 1999.Wild Orchids of Texas. 
(ISBN 0-292-7471 2-8, hbk). Univ. of Texas Press, PO. Box 7819, Austin, TX 78713-7819, 
U.S.A. $29.95, hbk. i-xii, 1-228 pp., color photos, distribution maps 


The Wild Orchids of Texas is an outstanding treatment of the fifty-two species of orchids known to 
occur in Texas and the authors deserve high praise for such an accomplishment. It begins with a 
chapter giving an overview of the state’s orchids emphasizing their diversity and conservation. 
Subsequent chapters give a clear and interesting discussions of such topics as pollina- 
tion, saprophytic species, and orchid-fungal partnerships. These are followed by a detailed discus- 
sion of orchids by the main habitat in which they occur, incl bogs 
habitats, forests and woodlands, and mountains and canyon 

The main part of the book, taking up about three- eee of the text, is a wonderfully com- 
plete trip through all of Texas’ orchids by genus and species. For example, the treatment of the 
genus Calopogon (grass pink orchids) begins with a discussion of the genus, giving such informa- 
tion as the number of species (five) and the range (eastern North America from Newfoundland to 
Cuba and westward to Minnesota and Texas). The derivation of the genus name is said to be de- 
rived from two classical Greek words meaning iaeue: peal: in PSIEISnce to the up tufted with 
colorful hairs. An insightful discussion on pollinatio Pog w the pollinating bees 
are victims of these deceptive orchids which give no reward iene as eee in return for the 
insects’services. Both of the Texas species of Calopagon are then discussed with such information 

n name, range,a detailed description of the species characteristics, ta~onomic and no- 
menclatural history, habitat and other ecological data including blooming time, and a county dis- 
tribution map showing precise locations in Texas. Such thorough accounts are given for each of 
the state's genera and species. While my basic reaction is that only praise is due this work, | could 
make a few of the most minor criticisms. For example, a few recently rey | distributional records 
were missed. 

The authors have clearly given extensive thought to making the book as useful as possible to 
a varied audience. For example, for amateurs wanting to identify an orchid with which they are not 
familiar, there is a list of orchids by flower color. However, for both professional and interested lay 
readers, the most exceptional and enjoyable part of the book is the quality of the remarkable color 
photography. Joe Liggio is a gifted photographer and the approximately 90 photos are a testa- 
ment not only to his artistic and technical skills but aS to his intimate aes - ann 
Some of the species photographed rarely and irregularly t mely 
peat places. The complete coverage of the state's species is thus ey the eed . hard 

work covering many years. The beauty of the photographs is complimented by the pleasing layout 
and the obvious thought given to design. There are, for example, numerous appealing details such 
as interesting quotes in the spacious margins and special text boxes pointing out particularly fas- 
cinating bits of information (e.g.,"ice Age Legacy" describes a rare yellow Lay's slipper discovered in 
the Texas Panhandle, apparently a relict of a colder and wetter climate regime during the last ice 
age). 

The scientific accuracy and attention to detail evident in this volume also make it valuable to 
professional botanists. The Liggios and their scientific advisor, David Riskind, have used the most 
up-to-date information on scientific nomenclature, have visited herbaria to obtain information for 
the detailed distribution maps, and have carefully documented their sources in an extensive Lit- 
erature Cited section. As a taxonomist currently working on a treatment of the orchids of East Texas 
(as part of the /lustrated Flora of East Texas project), | found the book an extremely valuable and 
helpful resource. For example, the genus Spiranthes (ladies’-tresses orchids) is known for its diffi- 


,open sunny 


SIDA 19(1): 223. 2000 


224 SIDA 19(1) 


ue in estate ae adn eee There is a oe clear and ee section Clscubsing the 

S.C orata,t ifficult species. 
Finally, the authors’ intimate en. of the ae in ‘the field is evident throughout the book 
and is a welcome relief in a scientific climate where work in the laboratory is too often valued more 
than knowledge of organisms in the wild. 

In summary,among state and regional treatments of orchids or other plant families, this is the 
most detailed, aesthetically pleasing, and just plain enjoyable book I've seen. From the beautiful 
dust jacket to the superb photography and details about orchid biology, it was clearly a labor of 
love for the authors. This is a book that anyone interested in Texas plants definitely should not 
miss.—George M. Diggs, Jr, Dept. of Biology, Austin College, Sherman, TX 75090, and Botanical Research 
Institute of Texas, Fort Worth, TX 76102. 


Joun H.Wiersema and Bianca Leon. 1999.World Economic Plants: A Standard Reference. 
(ISBN 0-8493-2199-0, hbk.). CRC Press LLC, 2000 Corporate Blvd. NW, Boca Raton, FL 
33431, U.S.A. (1-800-272-7737; 1-800-374-3401 fax).$125.00, hbk. 749 pp. 


While discussing some of my research on the Kallawaya herbalists of Andean South America with 
one of the researchers here at BRIT, it was recommended | consult World Economic Plants to help 
validate some of my work. Covering almost 10,000 vascular plants of economic importance, this 
reference volume is sure to be on the shelves of any botanist or anthropologist concerned with 
economic plants. Data were compiled for the Germplasm Resources Information Network (GRIN), 
which is the USDA/ARS's genetic resource management database. While much of this database is 
open to the public via internet access, this volume makes a strong argument that i a book 
off the shelf,as decadent as that may Scum) sonia still bea ce way to get an answ 

Wiersema and Leon divide th J s, both of which are organize. oa 
betically, Part one is the catalog shee economic caine offer if ie scientific name, syno 
mon names, economic impacts, and geographical distribution for a given plant. Part two ee an 
index of common names that includes the list of common names from part one with correspond- 
ing scientific names. All of the above information was validated by 150 specialists, something few 
reference volumes can boast about. Following a brief introduction that describes the methods of 

ili mes a discussion of content where the authors discuss the rationale for part one 
and its fee While serving as an excellent reference to confirm plant names, scientific and com 
mon, this book first and foremost is concerned with ie ic salience Sika plants world- 
wide. Under the heading of economic importance th reference to the plant's role as 
a food or food additive, animal food, bee plant, be eae food and poison nes fuel, social 
use, medicine, environmental use, gene source, w and harmful oe host. For plants that 
are subject to the regulation of the Convention on pee 1 Species (CITES) 
the authors have made the appropriate reference. Completing ve mission of the book, the « au- 
thors cite the geographical distribution of each plant, following the standard World G hical 
Scheme for Recording Plant Distributions (Hollis & Brummitt 1991), that divides the ierectial were 
into nine areas: Africa, Antartica, Asia-Temperate, Asia- Tropical, Australasia, Europe, Northern America, 
Pacific, and Southern America. However, the actual breakdown does not quite gel with the divi- 
sions. For instance, for North America the authors include Canada, U.S.A,and most of Mexico.While 
_ America includes Mesoamerica, the West Indies and South America. 
current research is concerned with the plant origins of the Kallawaya pharmacopoeia, 

meet exotic botanicals or non-native plants. Using my own data as well as “Pharmacopeia of 
Qollahuaya Andeans” by Joseph W. Bastien (1983), Journal of Ethnopharmacology, 8:97—111,| looked 
for information on a number of plants, using both their common and scientific names. | looked in 


SIDA 19(1): 224. 2000 


BOOK REVIEWS 225 


the common name index ot altansa, for which the authors give the scientific name Tanacetum 
parthnium, but Basti entific name Ambrosia peruviana.| looked for A. peruviana in the 
catalog of economic plants and vale many species of the Ambrosia genera are listed, there is no 
citation for A. peruviana. When | looked for berro in the common name index for its scientific name, 
both World Economic Plants and Bastien cite the name Nastrium officinale. But, when | looked in the 
talog of economic plants for N. officinale the authors cite its only economic in perenice asa 
pee eae Wee Bastien cites a medicinal use (a Mate for Hepatitis and Kidney infections) the 
authors leave out.! decided to supplement my investigation with hlamoduced Plants in the Indig- 
enous Pharmacopoeia of Northern South America” by Brad Bennett and Ghillean Prance (2000) 
Economic Botany, 54(1):90-102, my rationale being a more recent article might limit confusion. | 
consulted the catalog of economic plants for Plantago major, a plant that Bennett and Prance cite 
as used to treat 12 body systems with 15 pharmacological properties. World Economic Plants lists 
its only economic importance as a Weed with possible seed contamination. Coincidentally, while 
investigating P major, | noticed that one of the common names listed was Manten. Bastien lists 
Hlanten as Plantago tomentosa, which until now,| had had a great deal of trouble locating at all. 
Using another plant from Bennett and Prance, Rosmarinus officinalis, | consulted the catalog of 
economic plants again. The authors cite it as a medicinal and its geographic distribution, not a 
South American plant, corroborates with Bennett and Prance. 
nsummary, while this book is not definitive, it still exists as th one of its kind. And as the 
forward states,"this handy reference will address the ongoing needs os broad spectrum of users 
including sOsietitules eo mnmodty traders, farmers eerily for poral crops or seeking to eradi- 


cate harmful veterinary workers concerned with poisonous plants; Scent Ie 
mation specialists; and scientists in a variety of disciplines that involve Kevin D. 
Janni. 


Ernest SMALL and Paut M. CaTuna. 1999. Canadian Medicinal Crops. (ISBN 0-660-17534-7, 
pbk). National Research Council of Canada. Monograph Orders, NRC Research Press, 
M-55, National Research Council Canada, Ottawa, Ontario Kl 1A OR6, CANADA. (613- 
990-2254:613-952-7656 fax) <htt ca/cisti hs.| I>.$29.95. 

240 pp., 25 color plates, b&w line drawings, distribution maps. 


Responding to the demands of the agricultural community for more information on native me- 
dicinal plants, the National Research Council of Canada has produced a reference book that sets 
the ne ey other countries to follow. Canada is no stranger to the medicinal plant jie 
They gros million annually from the cultivation of Ginseng, as well as producing other crops, 
like pie Pacific Yew, and goldenseal. With the current surge of interest in herbal ene 
to costly prescription drugs, medicinal plants are in vogue to a currently sluggish agricultural in- 
dustry. Canadian Medicinal Crops is targeted to the agricultural community, offering not only an 
encyclopedic reference to 25 potential medicinal crops, but also a section devoted to the business 
of medicinal plants. 
ening with an appropriate introduction for the uninitiated, the authors discuss medicinal 
plants from ancient times to present, medicinal chemicals, herbals vs.pbharmaceuticals, nutraceuticals, 
and economic opportunities. This sets the stage for the core of the book and the 25 plants of 
economic potential and their detailed species accounts. For each plant the authors provide the 
scientific, English and French names, morphology, classification and geology, ecology, medicinal 
uses, toxicity, chemistry, non-medicinal uses, agricultural and commercial aspects,and a section on 
ths, legends, tales, folklore, and interesting facts. For each plant there is also a page-sized color 
esacee of the plant,a geographical distribution map and a list of selected references and World 


SIDA 19(1): 225. 2000 


226 SIDA 19(1) 


Wide Web links. | found the references to web links a nice bonus, the authors cite web sites in the 
text, so—for example—while reading about the toxicity of Arnica Montana, the authors provide 
the web address to a 1995 Health Canada document on herbs used as non-medicinal ingredients 
in nonprescription drugs. The web links listed after the selected references section are more plant 
specific. For example, in the chapter on echinacea the authors cite http://res.agr.ca/lond/pmrc 
study/newcrops/echinacea.html, plus nine other sites. The addition of web links makes informa- 
tion readily accessible, and along with the text itself, comprehensive 

Following the information on plants comes chapters eee to the business of growing 
medicinal plants, the regulatory and legal framework in Canada for producing and marketing me- 
dicinal plants, various hazards, and a general overview of medicinal plant research in Canada. This 
section is quite interesting- the auIOls cna principal determinants of the commercial impor 
tance of medicinal plants, marketplace timing, a list of nati tic, insecticidal, 
crops, and a list of non-native medicinal crops a could be grown in Canada. While this section 
does pay some lip service to conservation and sustainability, it would be nice to see in future edi- 
tions a full-blown chapter on the subject, with references and web links. There is also a section on 
medicinal cautions that discusses health fraud and gives recommendations to legislators, regula- 
tors, product manufacturers, physicians, and consumers, and discusses herbal remedies in Cana- 
dian and American laws. The last 30 pages consist of a list of ee resources, such as con- 
tacts to Canadian experts, organizations, and publications, an ff a regional review 
of medicinal plant research in Canada, a list of general selefences a list of wee sites on medicinal 

herbs, and mally a aa of pharmacological and medical terms relevant to medicinal plants. 
| was sligt ws eenpol ited pe there was no general index and that the authors did not fully 
but, otherwise Canadian Medicinal Crops is a model text for 

other countries to follow. The sections discussing the business side of the medicinal crop industry 
were particularly interesting and I'm sure, invaluable to anyone interested in starting a medicinal 
crop. Even though this book has a target audience (the agricultural community),! wouldn't expect 
readership to be exclusive. This book would also serve well as part of a medical botany class or as a 
general reference to anyone interested in Canadian medicinal crops.—Kevin D. Janni. 


AA 


JOHN T, tals and CuaistorHer A. Meacham. 1999. Synthesis of the North American Flora, 
Version 1.0. (ISBN 1-889065-05-6). North Carolina Botanical Garden and the Uni- 
nee of North Carolina at Chapel Hill, CB #3375, Totten Center, Chapel Hill, NC 27599- 
3375, A 

Minimum hardware configuration: Pentium 90 MHz-class processor, 32 megabytes RAM, 25 mega- 

bytes free hard disk space, SVGA display (800 X 600 pixel resolution) with 16 colors, Microsoft Win- 

dows 3.1,NT, 95,98, or 2000 operating system, CD-ROM drive for installation,a mouse, and, of course, 

a keyboard. 

This CD-ROM includes an updated version of Kartesz’s A Synonymized Checklist of the Vascular 

Flora of the United States, Canada, and Greenland, but it is so much more. Besides being able to 

switch between thesaurus and checklist format at the click of a mouse, you alt have th tion of 

omitting authors ie the listing, omitting synonyms alla or new with the Synthesis, dis- 
playing common names. And this is only the tip of the ic 

There are many ane practical functions packed into this program. They are so numerous 
and applicable in so many different ways that it's actually a little confusing at first.I'll start with the 
more straightforward functions. The Synthesis allows you to highlight any taxon from a family, ge- 
nus, or species list and displays its distribution on a map of the circumscribed area. It also color 
codes regions (the U.S. is divided into its states, and Canada into its provinces) as to status of the 


SIDA 19(1): 226. 2000 


BOOK REVIEWS 227 


taxon in terms of noxious, rare, extinct, eradicated, extirpated, or simply present. Another touch is 
discovered when you hover the cursor over a region and a flag pops up telling you the location of 
a voucher or in some cases a citation rel the vas on tet cr 
other basic function of th window. Ini is Siapiesel use 
it in. the biological attributes of the currently selected taxon. Some of the attr 
are general plant habit (forb, grass, shrub, tree etc.), major plant group (monocot, dicot, fern gym- 
nosperm etc.), duration (annual, perennial or biennial), nativity (native or exotic), habitat (alpine, 
aquatic, submerged, floating, wetland etc.), and the list goes on. 
hese functions alone are worth the investment in the CD. But it is only the beginning of the 
functions of the program. The rest revolves around three possible queries and four operations you 
can perform on the queries in numerous Boolean permutations. From here | found it somewhat 
convoluted if not in design at least in the explanation. The four operations (union, intersection, 
restricted, and “not") can be performed on any single, pair, or trio of queries (geographic, attributes 
or taxonomic). As an example you could select two geographic regions like Texas and North Da- 
kota and perform the union operation on the selection. The program would then highlight all of 
the a. ner occur in pESVIESe wp tales combined. Further you could add an attribute query 
tates. Finally you could add the third query wi th something 
ike Asteraceae and all annul dicots in the Asteraceae in both states would a highlighted. Besides 
the taxa in the list being highlighted a table with the number of taxa at each rank is displayed in 
the lower right hand corner. The intersection operation displays the taxa with the particular at- 
tributes or in a particular taxon, if selected, that are common to the chosen geographic regions. 
The restricted operation displays taxa that are restricted to the selected region or regions. Finally, 
the “not“ operation is the functional opposite of the restricted function and displays all of the spe- 
cies of the circumscribed area not found in the selected region.For example, there are about 28,000 
ane in the Synthesis (North America north of Mexico) and about 6,000 species in Texas. If the 
ot" operation were performed on Texas as the selected region the program would display 22,000 
species. This may be the least usable of the functions found in the program 
Besides this dizzying ile of fein eee combinations there are also several different 
colors of fonts used int q sts that carry different kinds of information. This add 
information an sometime to sink in ne a or may not be useful to some. 
lin all, once you spend some time with the program, it is quite usable. It will probably 
become the standard for its medium just as the Kartesz books have become a standard for the 
printed medium.—Robert J. George. 


W.S. Juno, C.S. Campbell, E.A. KeLLoce, and PF. Stevens, 1999, Plant Systematics: A Phyloge- 

netic Approach. (ISBN 0-87893-404-9, hbk.). Sinauer Associates, Inc. 23 Plumtree 

pocd Sunderland, MA 01375, U.S.A. (publis! ht l com/) 
$67.95, hbk. xvi + 464 pp., with CD- ROM. 


| shared the general enthusiasm of many colleagues in the summer of 1999 upon learning of the 
availability of anew textbook for plant oe . by Judd and colleagues, whose collective depth, 
breadth, and expertise in taxonomy is considerable. Promotional literature distributed by the pub- 
lishers indicates the text has been ene by a considerable number of institutions in the United 
States. However, this review is mixed because of eee reel interests in the text as a researcher 
and a teacher of introductory plant systematics. 

a detailed and modern source information for graduate students and thoroughly trained 
plant en the text is highly recommended. Stated simply, Plant Systematics is the only 


SIDA 19(1): 227. 2000 


228 SIDA 19(1) 


modern and relatively COMPIES: source of ntornation summarizing the numerous recent changes 
in sais Leo at the family level based on phylogenetic analysis. The text has an abundance 
of classic and more recent citations concerning the taxonomy of Reed groups. For these rea- 
Sons sre the text can be consulted profitably by all practicing taxonomists. The coverage of 
general topics by and large reflects what graduate students in taxonomy should master. The vol- 
ume probably should be consulted by all instructors of plant taxonomy, given emerging changes 
in our knowledge of evolutionary relations 
Apart from the Forward, Preface, and ic eden the book is as into eight chap- 
ters, two appendices, separate taxonomic and subject indices, and a CD-RO 
pter One, entitled “The Science of Plant Systematics’ immediately ae the phylo- 
aeneice context that pervades the entire text. Although the term synapomorphy is introduced 
sae simplified cladogram showing the shared derived characters uniting the major lineages of 
plants appears on the second page. Phylogeny is the unifying principal throughout the text, as any 
ee treatment should be. 
econd chapter, “Methods and Principles of Biological pene the principles of 
eee systematics are developed in some detail. Unfortunately, the treatment is much 
dense for most students at the introductory level. The excessive detail ee in sas two per- 
meates the text throughout, and from a pedagogical perspective, it is the largest shortcoming of 
the book. As one who used the text for a semester long, upper-division te pexoniomy course at a 
medium-sized public university shortly after its introduction, | suggest wit that 
the level of detail overwhelms most beginning students 
5 such, and with greater perusal, one wonders for whom the text is targeted. For example, a 
number of unrooted networks in chapter two illustrate how character state variation can be or- 
dered, which seems unnecessary at an introductory level. Likewise, a concise summary of long- 
branch attraction appears as an offset box on page 20, which apparently is deemed more impor- 
tant to beginning students than including simple illustrations showing the differences between 
monophyly, paraphyly, and polyphyly (although monophyletic and paraphyletic groups are shown 
in fig 2.23 in the context of phylogenetic and evolutionary classifications). The common use of 
cladograms early in the text is admirable, as for example in the first chapter, where Common names 
are used as terminals of cladograms (raspberries, blackberries, cherries) to illustrate Common an- 
cestry and sister taxa. Soon thereafter (figs. 2.9, 2.12, 2.13), however, cladograms begin to incorpo- 
rate terminal taxa such as Piperales, Winteraceae - Iruruaceae ieee Calycanthaceae, 
other names that mean nothing to the majority of students who likely would be covering this 
re during the second or third week of an introductory taxonomy class. Since thorough com- 
prehension of diagrams generally requires some understanding of word(s) contained therein, ex- 
amples of cladograms containing more accessible names might have been preferable from the 
perspective of beginning students. One topic in which insufficient detail is provided in chapter two 
is the inadequate coverage of phenetic and evolutionary classifications. If students are to under- 
stand the importance of phylogenetic systematics in an historical context — given the phyloge- 
netic unity of the text, how could the authors wish otherwise? — then they must have a greater 
appreciation of the methods of classification that preceded it. 
in contrast, the third chapter, entitled “Classification and System in Flowering Plants: The His- 
torical Background’ seems to approximate more closely the level of coverage towards which an 
introductory text probably should strive. The author of this chapter, P. Stevens, centers the discus- 
sion around numerous historically prominent taxonomists. Sometimes, however, the text presup- 
poses a knowledge that beginning students clearly will lack, such as a reference to Bentham and 
Hooker (p. 36), which would be understood only by those possessing considerable depth in their 
knowledge of taxonomic history. Once again, one wonders for whom the text is written. 


SIDA 19(1): 228. 2000 


BOOK REVIEWS 229 


The fourth chapter,“Taxonomic Evidence: Structural and Biochemical Characters” also exem- 
plifies the excess of detail that characterizes much of the text.The figures accompanying the narra- 
tive are good, but | question the necessity of eight types of marginal teeth in leaves (fig. 4.13), 
eleven variations of cell types in xylem (fig. 4.31), five TEM ee of sieve-element plastids 


(fig. 4.32), nine different stomatal types (fig. 4.35), twelve “common petiole vascular patterns” (fig. 
4.36), and entire sections regarding “Anatomy” and “Embryology’ one specialists need to 
know much of this information, but not ee students. Besides material belongs 


more appropriately in a text covering traditional plant anatomy. One area where detail was appre- 
ciated by this reviewer was in the illustrations of important secondary plant compounds, which are 
not included in so 
The floral ane (Box 4a:57) also seem excessively complex as presented in the text. Al- 
though these need not be of universal consi | think those of Smith (1977), which use a letter 
(e.g., K for calyx, C for corolla, etc.) to anchor che numbers associated with each floral whorl, are 
easier for students to understand and apply. 
tailed discussion of inflorescence types and the fundamental dichotomy between 
ac a indeterminate inflorescences is another situation where excessive detail drowns 
out the basic knowledge beginning students require. The terms panicle, raceme, spike, cyme, um- 
bel, corymb, head, and a few more specialized ones (e.g., spikelet, perigynium) are generally ad- 
equate to key out specimens in regional floras. piougn the basic Gicneromy of ISIE Scenes 
types is necessary for specialists, 1.aspanicle- 
like cyme” and “raceme-like cyme” are cumbersome. A better approach might have Be to intro- 
duce the basic terms denoting floral arrangement, followed by a relatively brief commentary on 
the distinction between indeterminate and determinate inflorescences. Speaking again from ex- 
perience in the classroom, the determinate/indeterminate distinction seemed t to interfere with 
the student's ability to understand the more basic terms associated with inflorescence structure. 
hapter four includes other topics that should have been omitted or relegated to separate 
chapters. Sections covering pollination Cronies coevolution between plant and pollinator, de- 
aie in orchid pollination, self-i hromosomes, and palynology all 
m either unnecessary or out of place. Here the text : seems to drift off course and begins to 
ae a text on reproductive biolo 
hapter Five, entitled “Molecular Systematics is also written at a level of detail considerably 
beyond an introductory text. For example, figs 5.1 and. 5.5 are bewildering in their complexity to 
anyone but an evolutionary biologist or eee specialist. The clarity and simplicity of figs 5.2, 
5.3,and 5.7 are more appropriate for an introductory text.In view of the molecular focus of much 
recent research in plant taxonomy, and given the need for more advanced students to have an 
introduction to the topic, the chapter is a useful component of the book. Graduate students and 
established taxonomists trained before the advent of phylogenetic analysis and the use of mo- 
lecular data may find much of the modern cladistic and molecular terminology more comprehen- 
sible after rev this chapter. To them | recommend the text without reservation 
th chapter is the first one not directly related to detailed as Meco OLOnG Entitled 
“The Evolution of Plant Systematic Diversity’ the text returns to the theme of phylogeny, history 
and evolution. Microevolutionary processes are covered, including the nature of variation in plant 
species. oe is raise discussed in some detail, as are polyploidy and species concepts. Re- 
garding the latter, th rs tly highlight varying perspectives of the Le species 
concept sithouiak it eludes me why the genealogical concept of species is mention Il. 
Starting with Chapter 7, “Phylogenetic peony of Maj ol Groups oF acheophys Ex- 
cluding Angiosperms’ the authors begin their detailed coverage of family treatments, following to 
a considerable extent the classification of the Angiosperm Phylogeny Group (1998). Along with 


SIDA 19(1): 229. 2000 


230 SIDA 19(1) 


Chapter eight, which covers angiosperm families, 145 families are treated in detail. These families 
are summarized on the back inside endplate, and boldfaced in table 8.1. The more inclusive groups, 
whose cladistic relationships are sUmimaniced 0 on the inside en plates, include the “Free-sporing 
acheophytes” (Lycopodiophytes, Psilophytes, Eq ferns), the "Seed 
ae ts" (cycads, Ginkgo, Conifers),and the Angiosperns Non- monocot palecherbs™ 


“Magnoliid com- 
plex’;"Monocots’s.s.,"Basal tricolpates’ Cayophyl and a few smaller lineages;"Eurosids | and II’ 
Cornales and Ericales: and “Euasterids | and II").As indicated above, the book is an excellent vehicle 
for professional taxonomists to acquaint themselves with newer concepts of higher taxa. 

The quality of illustrations of taxa, which are generally recycled from other sources and there- 
fore inconsistent regarding level of detail, ranges from fair (e.g., fig.8.99) to good (e.g.,fig. 7.15). The 
book is inconsistent in providing descriptions for some (e.g., Malvales) but not all (e.g., Apiales) 
Orders. Keys are typically provided to the major families of a given order. In some cases (@.g., 
Brassicales), fairly detailed cladograms are also provided. Although the average student cannot 
grasp this amount of detail, it is a positive sie to see a text using Gacodielns and ae 

re 


moting Se a ee eens Families wl monophyly 

indicated 1 to grasp ae Putative pe 
are boldfaced, athough to reiterate he theme, family descriptions are excessively detailed for an 
oe 


ae of leptosporangiate fern families are subsumed under Polypodiaceae, which 
seems a necessary approach for an introductory text. The coverage of Gnetales lacked any cita- 
tions of Friedman (1990, 1994) pede the evolution of double lea which came as a 
surprise. Judd et al. has a recurring bias towards coverage in the Northern Hemisphere. For example, 
despite an erroneous referral in the Sent Index to page 152 for Araucariaceae, this as is 
not treated. However, its coverage woul have proces an opportunity to mention the excite- 
ment, careful research,and with the recent discovery of eae 
Pine in Australia. The bias returns later in Table 17 in alent 2,where important journals from the 
Southern Hemispher looked, such as Australian Systematic Botany, Austrobaileya, Bothalia 
Darwiniana, New Zea land Journal of Botany, and South ae Journal of Botany, to mention some. 

A ae covers the pag of Bereled OMEN CINE oo in uae to much of the 
book, seems to be written at a level a d. However, table 1, which 
contains ene 300 Latin names of specific he, is capatiesioneble value. In contrast, a normal 
glossary is absent. Since beginning students rely on glossaries constantly, its absence is a promi- 
nent shortcoming 

ee 2, likewise written at an appropriate level for beginners, reviews aspects of making 
collections, conservation laws, plant identification and keys, an overview of the taxonomic litera- 
ture (particularly journals), and a brief mention of herbaria and botanical gardens. 

The M contains some 650 color photographs of plant taxa, which can be selected 
from an siphapetical list (by genus) of species, by families, or by orders. Like Cantino (1999),| ques- 
tion the value of photographs in the teaching of systematics. Most students seem relatively unin- 
terested in the use ge CD-ROMS in the lab, probably because they know the material therein will 
not be don sens the less, anny photos are of high quality, the taxonomic coverage 
is broad, ane some will no dou that some taxa are illustrated by more than one photo 
(e.g., Albizzia ju dal issin, peers a sibiflora, Cocos nucifera, Liriodendron tulipifera). My computer 
seemed unable to call up an image for Plantago major.Included in the CD-ROM are comparisons of 
more are te classifications of Cronquist and Thorne. 

e largest caveat to those who adopt the book for the classroom is to remain hawkishly 
alert to the details of family-level changes in the book, irrespective of whether we agree them all. 
Family circumscriptions sometimes depart considerably from the manner in which they have been 


SIDA 19(1): 230. 2000 


BOOK REVIEWS 231 


taught up to the present time. To cite one of the more spectacular examples, Scrophulariaceae 

sensu Judd et al.is considerably different from its previous circumscriptions sensu lato.M 

era aay placed in the family have been transferred into Plantaginaceae, such as Veronica, 
mon, Mimulus Digitalis, and Antirrhinum. The expanded lantaginaceae however, also now 


ee Callitrichaceae and Hippuridaceae. Other genera such as Pedicularis, Castilleja, and Agalinus 
have been transferred to Orobanchaceae. 

These and ee taxonomic changes will challenge our “knowledge” of families and force us 
to consider how we teach this information. My approach has been to explain the historical con- 
cept of the eae followed by the newer concept presented by the text. This provides a wonderful 
opportunity to champion taxonomy as a dynamic science, as the text admirably does, and stress 
that newer data support the need for the changes. The first time such a major familial level change 
arises | stress to the students that most other biologists will“know" these families in their previous 
incarnations, and that nearly all floras will treat them as such.| also caution that students may be 
challenged by their elders regarding their knowledge of newer familial boundaries. One can easily 
imagine a bright younger student raising the ire of an older biologist by insisting that Penstemon 
belongs in Plantaginaceae rather than Scrophulariaceae. It is also fairly easy to envision the older 
biologist wondering what the nearest tax-supported university is teaching, and perhaps voicing 
disparaging remarks about the competence of the instructor. Revised taxonomies also provide an 
opportunity to discuss the pronounced discontent that can arise in the face of nomenclatural 
changes, and Hike students should be able to explain in non-technical terms why such changes 


ons periodically t 

Apart from decir the empirical basis for specific taxonomic changes, the abundance of 
taxonomic changes in the text can also offer an opportunity to discuss some basics of field diplo- 
macy, such as when and when not to challenge others on the “correct’ oe of specific taxa. 
For example, it would be lacking in common sense to pontificate about the “correct” disposition of 
Penstemon to a private landowner whose land you ene like to collect on, a who possessed 
some understanding of Scrophulariaceae s.|. As an anecdotal aside, some landowners have a so- 
sui aa knowledge of local plants. Two colleagues and | once were dumbfounded when a 

e rancher in southeastern Colorado, wie, when informed we were solute fora ae ee 
os plant technically called Haplopappus, nonchalantly and correctly asked whether we were look- 
ing for Haplopappus fremontii! 

In general, the text highlights some of the pedagogical challenges currently faced by instruc- 
tors.For example, is it better to teach families that have been merged in their newer, more inclusive 
sense (e.g.,Amaranthaceae), or as distinct families in their more traditional sense (e.g.,Amaranthaceae 
plus Chenopodiaceae)? To my mind it seems preferable to present both perspectives, since begin- 
ning students need to know these family names when keying out material in local Floras, and 
those who pursue taxonomy beyond the introductory level eventually will have to become ac- 
quainted with the older concepts. 

It is uncomfortable to say so given favorable impressions of the book as a researcher, but the 
biggest pedagogical challenge for a introductory taxonomy class may be whether to use the vol- 
ume at all. Despite being one who i is eo! ue lovlenng of eueieeleea standards, the book is too 
information rich for many beg evaluations largely echoed this sen- 
timent, even though much of the material was covered in lecture as well. When students are over- 
whelmed ule either umn sala cline ig text or ie Sate for information. The most poignant 
evidence of t reflected bya ho, unprompted from the instruc- 
tor, ae banging textbooks “ Smith (1977) a Walters ana Keil (1996) to lab. In short, from a 
teaching standpoint, the value of the book as an introductory text is questionable if many begin- 
ning students are unable to benefit from its contents 


— 


SIDA 19(1): 231. 2000 


232 SIDA 19(1) 


| hope that the authors will see the need for a distilled version of the book targeted for intro- 
BURT saad and quien produce a volume of perhaps 275 pages under a different title. The 
precedent already exists for Sinauer Associates, Inc.,to publish more detailed (Hart! and Clark 1989) 
and less detailed (Hart 1988) texts covering basically the same content by the same author(s). 

In conclusion, the book is indispensable for graduate students and as a source of updated 
information for well-trained systematists. But the authors need to remember that the great majority 
of students in introductory classes in plant taxonomy are those majoring in wildlife biology, for- 
estry, agronomy, horticulture, biology, and botany as well, but that only a small percentage have any 
meen to pursue taxonomy professionally. In my view the text does not succeed as a phyloge- 

etically oriented introduction to plant systematics for undergraduates, where the need is greatest. 


ACKNOWLEDGEMENTS 


My thanks to Jill Sipes, a recent UNC graduate, for her detailed discussion of the textbook from a 
student perspective. 


REFERNCES 


ANGIOSPERM PHYLOGENY Group. 1998. An ordinal classification of the families of flowering plants. 
Ann. Missouri Bot. Gard.85:531-553. 

Cantino, P. 1999. Review of: Judd, W.S., C.S. Campbell, E.A. Kellogg, P.F. Stevens. 1999. Plant 
systematics: A phylogenetic approach. Sinauer Associates, Sunderland, Massachusetts 
Syst. Biol.48:826-828. 

FricoMAN, W.E. 1990. Double fertilization in Ephedra,a nonflowering seed plant:its bearing 
on the origin of angiosperms. Science 247:951-954. 

FricoMan, W.E. 1994. The evolution of embryogeny in seed plants and the developmental 
origin and early history of endosperm. Amer. J. Bot. 81:1468-1486 

Haart, D.L. 1988. A primer of population genetics, 2nd Edition. Sinauer Associates, Inc., 
Sunderland, MA. 

Hartt, D.L. and A.G. Ciark. 1989. Principles of population genetics, 2nd Edition. Sinauer 
Associates, Inc., Sunderland, MA 

Suit, J.P, Jr. 1977. Vascular plant families. Mad River Press, Inc., Eureka, CA. 

Wacters, D.K.and DJ. Kei. 1996.Vascular plant taxonomy. Fourth Edition. Kendall/Hunt Pub- 
lishing, Dubuque 


—Neil Snow, Department of Biological Sciences, University of Northern Colorado, Greeley, 
CO 80639, USA. 


SIDA 19(1): 232. 2000 


BOOKS RECEIVED 233 


BOOKS RECEIVED 


Rocers McVaucH. 2000. Botanical Results of the Sessé & Mocifho Expedition (1787-1803): 
VII. A Guide to Relevant Scientific Names of Plants. (ISBN 0-913196-68-1, hbk.). 
Hunt Institute for Botanical Documentation, Carnegie Mellon University, 5000 Forbes 
Avenue, Pittsburgh, PA 15213-3890.$55.00.626 pp 

From the preface: The purpose of this paper is not taxonomic, in the sense that it is not mono- 

graphic, but primarily a task of reporting. Most of the identifications reported herein are those 

made by otier workers who have studied the publications of Sessé & Mocifo, or panel i te 


orthe ill nade by their artists.| have not attempted to give modern mes 
except in a few groups en | have special mowers EWS] usvally ve eae to 
comment upon th f those who have made ide tions, but merely to report the 


fact of identification. Naturally many names that were applied in v7 herbarium of Sessé & Mocino, 
in 1936 and soon thereafter, have become obsolete for taxonomic or other reasons. Many new 
species and genera that were based on specimens or icones during the first half of the 19" century 
have since been re-evaluated.” Contents: Preface, Introduction to the Guide, How to use the Guide, 
The use of “ex” attributed to authors, Authors contemporary with Sessé & Mocino, Latin and other 
quotations, Relevant names defined, Unpublished names, Documentation from /cones, Authorship 
of names based on the Sessé & Mocino paintings, Unpublished names applied by de Candolle, 
Documentation from non-pictorial sources, Appendix: Mocino’s list, Selected references, Annotated 
list of names, and Index to scientific names. wo new combinations are published in the volume: 
Dahlia cordifolia (Sessé & Moc.) McVaugh, comb. nov. (p.148) and Bernardia dodecandra (Sessé & 
Cav.) McVaugh, comb. nov. (p. 222) 


HUNT INSTITUTE FOR BOTANICAL DOCUMENTATION. 1998. Sessé & Mocino Biological Illustrations. 
(ISBN 0-913196-60-6, CD-ROM). Carnegie Mellon CD Press, Carnegie Mellon Uni- 
versity, 5000 Forbes Avenue, Pittsburgh, PA 15213-3890.$40.00. Shipping/Handling: 
$4 Domestic; $6 Outside U.S. 

“The collection compris 000 watercolor drawings and sketches; about 1,800 are 

of botanical er ees Full aunuenens for aavigsting the CD- ROM are provided on the disk. Find 

artworks by accession number, subject, transcription or notes. Also offered is the Curator’s Choice, 
an interesting and beautiful selection of 100 paintings that were exhibited at the Hunt Institute 
shortly after the collection was acquired. The CD-ROM requires a color monitor and a World Wide 

Web browser, preferably version 4 or higher of Netscape Navigator or Internet Explorer.” 


SIDA 19(1): 233. 2000 


234 


SIDA 19(1) 
ANNOUNCEMENT 
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UNIVERSIDAD AUTONOMA DE NUEVO LEON 0% 
x 9 g 
IV CONGRESO LATINOAMERICANO DE 3 og 
ESTUDIANTES DE CIENCIAS FORESTALESALECIF 5 : 
Compartiendo Oportunidades y Retos para el ) t Hi 
Manejo Sustentable de los Ecosistemas Forestales" Slermemes 


Linares, México del 9 al 14 de octubre del 2000 


Para: Estudiantes de Ciencias Forestales de América Latina 


Estimados estudiantes forestales: 


La presente tiene como objetivo invitarlos oficialmente al IV Congreso de la Asociacion 
renee os mallee de Ciencias Forestales aed que se llevara a cabo en la Facultad 
d 


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- ; _ : a octubre del 2000. 

tema principal del Congreso es: OimipSsHHenaS Oportunidades y Retos para el Manejo 
Sustentable de los Ecosistemas Forestales 


m™m 


otemas que se decane en el evento son: 1) Silvicultura y Restauracion de 
Ecosistemas Forestales; 2) Operaciones y Técnicas Forestales; 3) Inventario, Monitoreo y Evaluacion 
de los Recursos Forestales; 4) Productos Maderables y no Maderables; 5) Certificacién Forestal y 
alc Internacionales; 6) Fauna Silvestre 
El costo de inscripcidn es de USD $150. 00, los cuales cubren los gastos de alojamiento, 
alimentacion, salidas a campo, refrigerios, diploma y papeleria, entre otros. En cuanto al numero de 
participantes por cada Institucidn no existe restriccion. 
os resumenes de las ponencias deberan ser una breve descripcién de estudios de caso, 
investigaciones y/o experiencias, debiendo contener: nombre del autor(es), direccidn postal y 
electronica, introduccién, metodoloagia, resultados, discusiones y conclusiones, presentados en una 
cuartilla maximo hasta el 15 de agosto del 2000, fecha limite de inscripciones para ponentes y 
participantes 


peramos recibirlos en nuestra Universidad y ofrecerles nuestra caracteristica hospitalidad 
mexicana. Mayores informes, favor de comunicarse a: 


Facultad de Ciencias Forestales, UANL. 
Tels: (52821) 2-48-95 y 2-42-51 y Fax: (52821) 2-42-5 
Email: raranda@ccr.dsi.uani.mx y juvalle@ccr.dsi.uanl.m 


Atentamente 
Rafael Aranda Ramos José |, Uvalle Sauceda 
Presidente del Comité Organizador Comision de Contenido 


SIDA 19(1): 234, 2000 


> 


“You don't have to be a botanist to use it! All the tools you need 


to teach plant taxonomy, family characters, classification, etc. 
Excellent introductory materials on the history and geology of 
North Central Texas. Great for identifying indigenous plants in 
an ecosystem! Essential for student wildflower collection 
projects! Use this one book instead of five, it has it all.” 
—Sharon Foster, 7" grade teacher, 


Fort Worth Country Day School 


“The best available, user-friendly source of information about 
our native and naturalized plants. This book should be in the 


library of every gardener, landscape architect, horticulturist, 


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To be more specific: COMPREHENSIVE, ENCYCLOPEDIC, 
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departure in Texas floras, a breath of fresh air.” 

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Manual of the Vascular Plants of Texas 


“The Illustrated Flora on North Central Texas is a magnificent work. 
This abundantly illustrated volume will not only be of use to 
those people wanting to identify Texan plants, it will be a 
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VOLUME 19, NUMBER 2, PAGES 235-443. 
Copyright 2000 

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ISSN 0036-1488 


TABLE OF CONTENTS 


Ee 


IONEAE) 


A NEW VARIETY AND FOUR NEW COMBINATIONS IN PITTOCAULON AND TELANTHOPHORA (A 
FROM Mexico 
BONNIE L. CLARK—235 


ADDITIONS TO THE FLORA OF CRATER Mr., PAPUA NEW GUINEA 
W. TAKEUCHI—237 


THE INTRIGUING CASE OF CYPRIPEDIUM CROSSII, ITS PRIORITY OVER CYPRIPEDIUM CALLOSUM AND ITS TRANSFER 
TO THE GENUS PAPHIOPEDILUM 
Guido J. BRAEM AND KARLHEINZ SENGHAS—249 


USE OF VARIETY AND SUBSPECIES AND NEW VARIETAL COMBINATIONS FOR STYRAX PLATANIFOLIUS (STYRACACEAE) 
B.L. TURNER AND Guy L. NEsomM— 


NEW SUBTRIBES FOR a AMERICAN ASTEREAE (ASTERACEAE) 
Guy L. Nes 


STYLOGYNE AGUARUNANA (MYRSINACEAE) A NEW SPECIES FROM AMAZONAS, PERU 
JOHN J. PIPOLY II] AND JON M. RICKETSON—269 


Discovery OF ARDISIA SUBGENUS ACRARDISIA (MYRSINACEAE) IN MESOAMERICA: ANOTHER BOREOTROPICAL ELEMENT? 
JOHN J. PIPOLY II] AND JON M. RICKETSON—275 


SENECIO QUAYLE! (ASTERACEAE: SENECIONEAE), A NEW SPECIES FROM NORTH CENTRAL TEXAS, U.S.A 
THEODORE M. BARKLEY —285 


SOLIDAGO VILLOSICARPA (ASTERACEAE: ASTEREAE), A RARE NEW SOUTHEASTERN COASTAL PLAIN ENDEMIC 
R.J. LEBLOND—291 


STRUCTURE OF POPULATIONS OF OTATE (OTATEA ACUMINATA SUBSP. : POACEAE) IN HARVESTED STANDS 
Jose MARTIN VAZQUEZ Lopez, BRUCE F. BENZ, MIGUEL OLVERA VARGAS, AND SERGIO GRAF MONTERO—301 


BLOOMING "BEHAVIOR" IN FIVE SPECIES OF BOERHAVIA (NYCTAGINACEAE) 
RICHARD SPELLENBERG—311 


CYPERUS SANGUINOLENTUS (CYPERACEAE) NEW TO THE SOUTHEASTERN UNITED STATES, AND ITS RELATION TO THE SUPPOSED 
ENDEMIC CYPERUS LOUISIANENSIS 
RICHARD CARTER AND CHARLES T. BRYSON—-325 


CyYPERUS (SUG. QUEENSLANDIELLA) HYALINUS (CYPERACEAE) NEW TO THE UNITED STATES AND THE WESTERN HEMISPHERE 
RICHARD CARTER AND RANDY L. MEARS— 


LEGUMES FROM THE CENTRAL PART OF THE STATE OF CHIHUAHUA, MEXICO 
A. EDUARDO ESTRADA C. AND ALFONSO Martinez M.—351 


A VASCULAR FLORA SURVEY OF CALCASIEU PARISH, LOUISIANA 
Ray NEYLAND, BILLIE J. HOFFMAN, MARK MAYFIELD, AND LOWELL E. URBATSCH—361 


ESTABLISHING ETHNOBOTANICAL CONSERVATION PRIORITIES: A CASE STUDY OF THE KALLAWAYA PHARMACOPOEIA 
KEVIN D. JANNI AND JOSEPH W. BASTIEN—387 


CRYPTOCORYNE BECKETTH (ARACEAE), A NEW AQUATIC PLANT IN TEXAS 
David J. ROSEN—399 


THYMELAEA PASSERINA (THYMELAEACEAE) NEW TO TEXAS 
WALTER C. HOLMES, JOHN F. PRUSKI, AND JASON R. SINGHURST—403 


REDISCOVERY OF RIBES NIVEUM (GROSSULARIACEAE) IN COLORADO 
TIMOTHY W. CHUMLEY AND RONALD L. HARTMAN—407 


pass 


ALOPECURUS MYOSUROIDES SCLEROCH DURA (POACEAE) NEW TO LOUISIANA 


JOHN K. SAICHUK, CHARLES M. ALLEN, AND WILLIAM D. Reese—411 


GAMOCHAETA SIMPLICICAULIS (ASTERACEAE: GNAPHALIEAE) IN GEORGIA 
Guy L. Nesom—413 


GEORGE JONES GOODMAN (1904-1999). A MEMOIR 
CHERYL A. LAWSON—415 


Nesta DUNN Ewan (1908-2000). A MEMOIR 
ANNE S. BRADBURN—42 1 


BOOK REVIEWS AND NOTICES 248, 256, 274, 284, 290, 324, 344, 402,414,424 


INDEX TO NEW NAMES AND NEW COMBINATIONS IN 19(2), 2000. 


ASTERACEAE SUBTRIBE ASTRANTHIINAE NESOM, SUBTR. NOV.— 265 
ASTERACEAE SUBTRIBE BOLTONIINAE NESOM, SUBTR. NOV.—266 
ASTERACEAE SUBTRIBE CHAETOPAPPINAE NESOM, SUBTR. NOV.— 264 
ASTERACEAE SUBTRIBE PENTACHAETINAE NESOM, SUBTR. N 
CANARIUM ACUTIFOLIUM (DC.) MERR. VAR. PIORIVERENSIS eet VAR. NOV.—237 
PAPHIOPEDILUM CROSSII (MORREN) BRAEM & SENGHAS, COMB. ET STAT. NOV.—251 
Laituiecatis CROSSII VAR. POTENTIANUM (GRUB & ROTH) BRAEM & SENGHAS, COMB. ET STAT. NOV.—254 
1 VAR, SUBLAEVE (RCHB.F.) BRAEM & SENGHAS, COMB. ET STAT. NOV.—254 
PITTOCAULON HINTONI! H. Ros. & ane VAR. CERROGRANDENSIS B.L. CLARK, VAR, NOV.— 
nee VELATUM (GREENM.) H. Ros. & BRETTELL VAR. TZIMOLENSIS (T.M. BARKLEY) B.L. CLARK, COMB, NOV.—235 
RKLEY, SP. NOV.—286 
Sones VILLOSICARPA LEBLOND, SP. NOV,— 292 
STYLOGYNE AGUARUNANA PIPOLy & RICKETSON, SP. NOV.—269 
STYRAX PLATANIFOLIUS VAR. MOLLIS (PW. FRITSCH) B.L. TURNER, COMB. ET STAT. NOV.—261 
STYRAX PLATANIFOLIUS VAR. TEXANUS (CORY) B.L. TURNER, COMB. ET STAT. NOV.—2 
STYRAX PLATANIFOLIUS VAR. YOUNGIAE (CORY) B.L. TURNER, COMB. ET STAT. NOV.—261 
TELANTHOPHORA COBANENSIS (J.M. COuLt.) H. Ros. & BRETTELL VAR. MOLINAE (H. Ros. & BRETTELL) 
MB. ET STAT. NOV.—236 
Gnas GRANDIFOLIA (LESS.) H. Rog. & BRETTELL VAR. SERRAQUITCHENSIS (GREENM.) 
K, COMB. ET STAT. NOV.—236 
een SUBLACINIATUS (GREENM.) B.L. CLARK, COMB. ET STAT. NOV.— 236 


¥ 


A NEW VARIETY AND FOUR NEW COMBINATIONS IN 
PITTOCAULON AND TELANTHOPHORA 
(ASTERACEAE: SENECIONEAE) FROM MEXICO 


Bonnie L. Clark 


8201 Hauser Drive 
Lenexa, KS 66212-2542, U.S.A. 


ABSTRACT 
Anew variety, Pittocaulon hintonii var. lensis (A S ioneae) is described, and 
Our! velatum var. tzimolensis, felanthophora cobanensis 
var. mone Telanthophora aiandifciane var. fe aauichoiss and Telanthophora sublaciniatus. 


Key Woros: Senecio, Telanthophora, Pittocaulon, Mexico, Central America 


RESUMEN 


Se describe una variedad nueva, Pittocaulon eae var. CeO IRGSHA x se ee cuatro 
combinaciones nuevas, Pittocaulon velatum var. tzimolens 
Telanthophora grandifolia var. serraquitchensis, er aanes sublaciniatus. 


var. molinae, 


Recent studies in certain Mexican and Central American segregates of Senecio, s.|. (Clark 
1996) have led to the recognition a new variety and the need for four new nomencla- 
tural combinations. These are recorded here to make the names available for curatorial 


purposes. More complete explanations will be presented when the revisionary studies 
are published. 


Pittocaulon hintonii H. or & ial var. eeu oot andenss B.L.Clark, var. Nov. Tyee: MEXICO: 


5 of Cerro Grande on road from Lagunillas 
to Campo Cuatro & juluamcene on n limestone cf fs above camp 4, dry, ecaue woodland, 
together with the gigantic ij nd 
Pedilanthus in full sun, lat. 19° 232 25 N and long. 103° 51' 30"-52' 30" W, elev. 1480 m, S ve 
1987, HH. Iitis et al, 29713 (HoLoryee: WISI; isotypes: IBUG, KSC!, TEX, ZEA). 


A var. hintonii cujusque capituli flosculis radiatis 14 (nec 7-9) perspicue diversa. 


Pittocaulon hintonii var. cerrograndensis differs notably from var. hintonii in having capitula 
with 14 ray florets instead of 7-9. It is apparently restricted to the Sierra de Manantlan, a 
region known for a number of endemics (Vazquez et al. 1995). The nomenclaturally typi- 
cal var. hintonii occurs farther to the north, in the region of Sierra Naranjillo, Michoacan. 


Pittocaulon velatum (Greenm,) H.Rob.& Brettell var.tzimolensis (1.M. Barkley) B.L.Clark 
nov. Senecio praecox (Cav.) DC. var. tzimolensis T.M. Barkley, Phytologia 69:142. 1990 
Type: MEXICO: CHiapas:Tzimol, tropical deciduous forest 15 km S of Comitan on road to Tzimol 

and Tuxtal Gutierrez, elev. 1200 m, 20 Mar 1981, D.E. Breedlove 50266 (HoLotyre: CAS!). 


SIDA 19(2): 235 — 236. 2000 


236 SIDA 19(2) 


Telanthophora cobanensis (J.M.Coult.) H.Rob.& Brettell var. molinae (H. Rob. & Brettell) 
B.L. Clark, comb. et stat. nov. Telanthophora molinae H. Rob. & Brettell, Phytologia 27:428. 
1974. Type: GUATEMALA: DeparTMENT OF SAN Marcos: near Alta Fraternidad, between San Rafael 
Pie de la Cuesta and Palo Gordo, W facing slope of the Sierra Madre Mountains, wet moun- 
tain forest,elev. 1800-2400 m, 10-18 Dec 1963, L.0.Wiliams, A.Molina & LP Williams 26271 (HOLOTYPE: 
USI isorypes: F!, GH!, MO!, NY!) 


ae grandifolia (Less.) H. Rob. & Brettell var. li abate (Greenm,) BLL. 
Field M at. Hist., Bot. Ser, 2(6):286 
1907. Telanthophora serraquitchensis (Greenm. H.Rob.& Brettell, a 27:428.1974. Tyee: 
CONE DEPARTMENT OF ALTA VerapAZ: Serraquitche, elev. 760 m, Apr 1889, J. Donnell Smith 


|! ,WOTYFe,& HI!). 


ra 


Telanthophora sublaciniatus (Greenm.) B.L. Clark, comb. et stat. n io cob 
.Coult. var. sublaciniatus Greenm., Ceiba 1(2):120. 1950. Tyee: | LA: Dene OF 
eile hills of finca Carmona Sout ast of Antigua, in damp forest, elev. 1590-1800 m 
27 Jan 1939, PC. Standley 63680 NO!; isotype: FI), 


ACKNOWLEDGMENTS 
This study derives from a doctoral dissertation prepared at Kansas State University, Man- 
hattan, Kansas, under the direction of T.M. Barkley. Rupert Barneby of the New York Bo- 
tanical Garden kindly rendered the Latin diagnosis. 


REFERENCES 
Ciark, B.L. 1996. A study of variation in Senecio Sect. Terminales Greenm. (Compositae; 
Senecioneae). Ph.D. dissertation, Kansas State University, Manhattan, Kansas 
VAzquez G., J.A.,R. Cuevas G., T.S. COCHRANE, H.H. IutIs, FJ. SANTANA M., and L.GuzmAn H. 1995. Flora 
de Manantlan. Sida, Bot. Misc. 13, Botanical Research Institute of Texas, Fort Worth. 


ADDITIONS TO THE FLORA OF CRATER MT, 
PAPUA NEW GUINEA 


W. Takeuchi 


BOranied Research Institute of Texas ane 


dl Danii Ne W (sy inea Fo re coagrch Inctithjte 
PO. Box 314, Lae, Morobe Province 411, PAPUA NEW GUINEA 
ABSTRACT 


The floristic checklist for the Crater Mt. Wildlife Management Area (CMWMA) is updated. Canarium 
acutifolium var. pioriverensis (Burseraceae) is newly described. 


Key Worbs: Botanical survey, Canarium, Papuasia 


ABSTRACT (JAPANESE) 
AVAF-U BEE SEM OMMF ryFPVALERMODOOICKHS. 
Canarium acutifolium var.pioriverensis (FIbv+2 DE) (SFA ICEMAN TS. 


INTRODUCTION 


As one of Papuasia’s largest wilderness environments, the Crater Mt. Wildlife Manage- 

ment Area (CMWMA) was recently a focal venue for an intensive program of floristic 

exploration and documentation. In the following account, an earlier botanical summary 

of the CMWMA (cf. Takeuchi 1999) is updated with several additions and corrections. 

BURSERACEAE 

Canarium acutifolium (DC.) Merr. var. pioriverensis Takeuchi, var. nov. Type: PAPUA NEW 
GUINEA. CHimeu Province: Crater Mt.Wildlife Management Area, Pio River, alluvial forest,06°47'S, 
145°02'E, 500 m, 25 Mar 1997 (stam. fl.), W. Takeuchi 11,886A (Hovoryre: LAE; soryees: A, BRIT, L). 


Differt a C. acutifolio (DC.) Merr. var. aemulans folioliis longioribus usque ad 43 cm longis, stipulibus 
majoribus usque ad 25 x 6 mm, fructibus ellipsoideis 24-27 x 12-14 mm 
Canopy tree, 25-30 m tall, deciduous, plank-buttressed to ca. 1 m height, outer bark fur- 
rowed, pallescent, slash ferruginous to ochraceous, clear-resiniferous, wood white, very 
dense. Branchlets longitudinally corrugate, ca. 1.5 cm diam., minutely furfuraceous, pithy, 
not hollowed with age; vascular strands peripheral, appressed to the pith. Leaves spiral, 
phyllotaxy 2/4, 6-8 jugate, 1.0-1.4 m long, the flush emerging simultaneously with flow- 
ering; leaflets membranaceous, (ovate), elliptic-oblong, or oblong, decrescent on the ra- 
chis, (11-)20-31(-43) x (6.5-)9.0-14.0 cm, apex rounded or (obtusely acuminate), base 
truncate to (cuneate), margins entire; venation camptodromous, bifacially prominulous, 
secondaries 10-15(-16), at the leaflet center diverging 55-70° from costulae, gradually 
arcuate or turning abruptly at the margin, crossing nerves scalariform, reticulum dense; 
laminar surfaces adaxially dark dull green, abaxially rnedium green, on both sides 
pusticulate, all parts with an indument of appressed furfuraceous hairs; petiolule terete 


SIDA 19(2): 237 — 247. 2000 


238 SIDA 19(2) 


or compressed, pulviniform at both ends, 9-25 x 1.5-2.5 mm; rachis cylindrical, shal- 
lowly sulcate, purple-green; petiole 250-390 x 7-10 mm, dilate at the branchlet, 
articulative or not, broadly concave on the upper side, rounded beneath; stipules persis- 
tent, inserted at the base of the petiole, acuminate, 14-16(—25) x 3-4(-6) mm, flat, paired, 
evenly tapered to the apex. Staminate inflorescence axillary (lateral), thyrsoid, pyramidal, 
51-60 cm long, rachide branches to 15 cm long, all axes glossy green or reddish-green 
and with an indument of pale scalelike hairs; rachis bracts narrowly acuminate, the larger 
ones 7-10 mm long, higher order bracts ovate, 1-2 mm long, undulate; peduncle 170- 
240 x 4-5 mm. Staminate flowers ellipsoid-oblongoid in bud, perianth 3-merous; calyx 
tubiform, ca.5 x 3 mm, 1/2-2/3 connate, lobes ovate, densely sericeous on inner sur- 
faces, glabrate or puberulent outside; corolla distinct, imbricate, petals lanceolate-ob- 
long, 6-7 mm long, adaxially glabrous, externally with lax sericeous hairs; stamens 6, 
uniseriate, equal, united at the base into a glabrous column ca. 1 mm high, filaments 
plane, 0.5-0.75 mm long, anthers basifixed, sagittate, mucronulate, oblongish, ca. 2.5 x 
0.5 mm, erect, pilosulous; disk dome-shaped or globular, rugose, glabrous, ca. 1 mm high 
and not exceeding the column, 6-lobulate, recessed at the summit: pistillode absent. 
Pistillate inflorescence unknown. Infructescence axillary (lateral), occasionally from defoli- 
ate nodes, ebracteate, puberulous, ca. 20 cm long, branching only at the top, peduncle 
170 x 4 mm. Drupe ellipsoid or (subovoid), 24-27 «x 12-14 mm, obtusely 6-angled in 
cross-section, apex acute or bluntly rostrate, locules 3,2 of the cells sterile and rudimen- 
tary, exocarp with scattered subappressed hairs; calyx persisting, spreading, not accrescent, 
lobes deltate or rounded, 3.0 x 3.5 mm, adaxially densely sericeous, externally glabrate. 

Distribution and habitat.—Known thus far only from lowland forest in the CMWMA. 

Etymology.—The varietal name commemorates the type locality. 


Paratype: PAPUA NEW GUINEA. CHimeu Province: Crater Mt. Wildlife Management Area, Pio River, foot- 
hill forest, 06°47' S, 145°02' E,500 m, Jan 1998 (fr.), W. Takeuchi 11,886B (LAE). 

Malesian Canarium has been treated by Leenhouts (1955a, 1955b, 1956, 1959: cf. also 
Steenis 1972). There are 20 species reported from New Guinea (ibid). 

The new taxon belongs to section Pimela (Lour.) DC. and is related to the complex 
consisting of C. acutifolium (DC.) Merr. and its allies. Because of the 6-staminate flowers, 
the novelty is comparable to C. acutifolium var.aemulans but is more robust, with leaflets 
to 43 x 14cm and with larger stipules to 25 x 6 mm. Although var.acutifolium has similar 
panicles, the fruits of var. pioriverensis are 24-27 mm long (up to 15.0-17.5 mm for the 
other varieties of A. acutifolium). The deciduous phenology of var. pioriverensis is also 
distinctive. According to village respondents the new tree is characteristically leafless 
immediately prior to flowering. Their report is substantiated by the fact that other indi- 
viduals of the same variety were seen in leafless condition near the type collection and 
later exhibited synchronous flowering and leaf flush. The correspondence between 
leaf and flower emergence is clearly reflected on the type, which has denigricant mem- 
branaceous leaves unlike the coriaceous blades usually seen on herbarium sheets of 
congeners. 


TAKEUCHI, ADDITIONS TO THE FLORA OF CRATER MOUNTAIN 239 


The new variety will key to the species using the modified sequence in Steenis 
(1972) and can be incorporated into the varietal conspectus by replacing lead 2b 
(Leenhouts 1956: 292) with the following continuation. 


2.Nerves 12-15 pairs 
Leaves 2-4-jugate, leaflets 7-18 x 3.5-8.5 cm, abruptly acuminate; stipules subulate, 

to 17 x 1.5 mm; fruits subglobose, to 17.5 x 15mm var.aemulans 
Leaves 6-8-jugate, leaflets (11-)20-31(-43) x peer te cm,apex obtuse or (gradu- 

ally subacuminate), never abruptly devel ipul 


acuminate, flat, broader, to 
25 x 6mm; fruits ellipsoid, 24-27 x 12- 14t mm var. pioriverensis 


OTHER COLLECTIONS 
ADIANTACEAE 
Coniogramme macrophylla (BI) Hieron,; coll. 12,797. Initially reported as ‘sp. nov. aff. 
macrophylla’ (Takeuchi 1999: 953), the CMWMA plants have once- pinnate fronds with 
pinnae to 36 x 5 cm. Comparisons against extra-Papuasian material indicate that the 


survey collections are merely a vigorous growth form. The lax venation noted previously 
(ibid) is actually within the range of variation for the species. 
MIMOSACEAE 


Archidendron hispidum (Mohlenbr.) Verdc,; coll. 17,270. (Fig. 1). Archidendron hispidum 
was formerly recorded only from Northern and Milne Bay Provinces (Nielsen et al. 1984: 
95:Nielsen 1992: 133). The Crater Mt. provenance places the species further west,and on 
the southern side of the Central Divide. The plant is apparently rare throughout its 
range. 

There are two species in the series Ptenopae Nielsen to which A. hispidum \s as- 
signed (ibid). Although the congener A.ptenopum Verdc. has terminal inflorescences, it is 
now apparent that A. hispidum is strictly cauliflorous. The Crater Mt. specimen confirms 
that series Ptenopae is characterized by an inflorescence of pedunculate umbels. How- 
ever, unlike A. ptenopum, the gynoecium in A. hispidum consists of a single ovary, so the 
pluricarpellate condition is not salient to the series. 

An accessory description is provided for the previously unknown flowers of A. 
hispidum: 

Inflorescence a panicle of umbels or (corymbs), cauligerous, pendulous, diffuse, to 
15 312cm,all axes filiform, nitid green, + densely hirtellous; floral bractlets chartaceous, 
oblongish or widest above the middle, ca. 1.0 3 0.3 mm, falling early, aglandular, hairy; 
pedicels 5-11 mm long;rachide branches usually alternate, 20-50 3 0.2-0.4 mm, bracte- 
ate at the base, the rachis bracts scarious or not, acuminate, 0.7-2.0 mm long, persistent 
adaxially glabrate, externally hirtellous; peduncle 5-52 3 0.4-0.5 mm. Flowers (measure- 
ments from spirit material) seen in bud only, up to 11 per cluster, bisexual, Peel pen- 
bfleshy, qreen, hirtellous;calyx obconic, 1.0-1.5 mm long, n subtruncate, 


wd 


tamerous, 
erulose, or denticulate; corolla gamopetalous but connate only near the base, the lobes 
lanceolate or ovate, ca. 2.5 3 1.0-1.5 mm; androecium polyandrous (ca. 40-50), stamens 
glabrous, entirely white, column obsolete, ca.0.5 mm long, filaments delicate, contorted, 


SIDA 19(2) 


centimeters inches 


Enea TTT ETT 


a 


a 


pnp ae 


a 


Fic. 1. Archidendron hispidum (Mohlenbr.) Verdc, Flowering habit. Scanned from Takeuchi 17,210. 


TAKEUCHI, ADDITIONS TO THE FLORA OF CRATER MOUNTAIN 241 


anthers minute, globular, 2-locellate; gynoecium glabrous, sessile, ovary solitary, 
columnar, slightly reduced near the top, style cylindrical, equal to the ovary, stigma dilate, 


obconic to disciform. 


MYRTACEAE 
Syzygium hylochare (Diels) Merr. & Perry; coll. 11,847. (Fig. 2). The survey voucher was 
previously cited as ‘Syzygium aff. roseum Merr. & Perry, possible sp. nov.’ but has been 
rekeyed to the given binomial. Although the Crater Mt. specimen has leaves with very 
prominent, evenly-spaced, and more numerous lateral veins, it is otherwise referable to 
the new result. The stipitate base (4-5 mm long) on dried flowers and the comment of 
the stipes being very noticeable on immature fruits (Hartley & Perry 1973: 168) are de- 
ciding. These latter features are well-expressed on the survey number. Syzygium hylochare 
is also apparently characterized by the appearance of large pustules on the lamina and 
inflorescence after drying, in the manner of S. malaccense (L.) Merr. & Perry sens. lat. This 
identification aid can be advantageously appended to the couplet sequence for the 
species. 

Syzygium hylochare is much closer to S.roseum Merr. & Perry than is apparent from 
the artificial key in Hartley and Perry (ibid). Specimens belonging to either facies should 
be compared against the congener, especially for collections from the southern districts. 


PIPERACEAE 
Piper arfakianum C.DC;coll. 12,453.The survey collection differs from the typical facies 
inter alia by the much sparser indument and the pendulous spikes with unusually fili- 
form peduncles. The delicate peduncles are only 0.1-0.2 mm diam.(0.5-1.0 mm in other 
provenances) and the rachis is similarly narrowed to 0.5-0.6 mm diam. (normally 1.0-2.2 
mm diam.).On the fully mature spike the individual fruits are elongate-ellipsoid rather 
than the more typical squat-obovoid shape, and laxly disposed so the rachis is occasion- 
ally visible between the berries. At 12 x 6.5 cm, the laminae are at the maximum end of 
the size range for the species. Although the Crater Mt. voucher was listed as ‘Piper sp. 
?nov.'in the CMWMA checklist (Takeuchi 1999: 977) the collections at hand suggest that 
the survey number can be accommodated within the variable concept for Parfakianum 
established in specimen annotations by Chew (cf. also Jebb 1987).While the character 
states for the Crater Mt. population appear distinct from the species sens. str., future 
collections should speculatively be able to unite the herbarium variation into a single 
continuum. If this does not eventuate, then the Crater Mt. plants will be deserving of 
future recognition as a separate subspecies or variety. 

Piper is well-represented in the Crater Mt. Wildlife Management Area, with 17 spe- 
cies now documented for the locality. 


ADDITIONS AND EMENDATIONS TO THE CRATER MT. PLANT CHECKLIST 
The following taxa are listed as additions and emendations to the earlier compilation in 
Takeuchi (1999). Voucher source for occurrence record: LAE = staff collections from the 
Lae National Herbarium series; UPNG = institutional series from the University of Papua 


242 SIDA 19(2) 


centimeters inches 


Fic. 2 5 gi hyloct (Di Is)M & Perry. Flowering habit. Scanned from Takeuchi 11,847. 


TAKEUCHI, ADDITIONS TO THE FLORA OF CRATER MOUNTAIN 243 


New Guinea Herbarium;WT = W. Takeuchi. Other collectors indicated by name. Determi- 
nations by WT unless otherwise noted. 


FERNS AND FERN ALLIES 

ADIANTACEAE 

Coniogramme macrophylla (BI.) Hieron.; WT 12,197 
DICOTS 


ASCLEPIADACEAE (all dets. by Pl. Forster) 
Hoya aff. microphylla Schltr; WT 12 
Hoya sussuela i Merr., WT 12,669 


Hoya sp; WT 12 

Marsdenia sp., probably nov., WT 12,469 

BURSERACEAE 

Canarium acutifolium (DC.) Merr. var. pioriverensis Takeuchi; WT 11,886A (type), 11,8868 
ERICACEAE 


Vaccinium acrobracteatum K. Schum.; WT 12,451, originally cited as ‘Gaultherieae indet.’ 

EUPHORBIACEAE 

ete ar ides Teijsm. ex Hassk.; WT 12,689, the voucher previously cited (WT 12,702) is ac- 
ually Codiaeum variegatum (L.) B Bl. 

MELASTOMAT. 

Medinilla na Bak.; J. Croft et al. LAE 61,178, det. J.Regalado 


MIMOSACEAE 
Archidendron hispidum (Mohlenbr.) Verdc.; WT 11,210 
MONIMIACEAE 
Steganthera sp.nov. ined.; WT 12,742 (type) 
MYRTACEAE 

Syzygium hylochare (Diels) sel & Perry; WT 11,847 
a sp.nov.ined.; WT 7 9 (type) 
Sates 

per rane C.DC; WT 12,453 

RUBIACEA 
Myrmecodia tuberosa Jack, entity ‘mueller’; UPNG 3,481, cited in Huxley & Jebb (1993: 285). 
Myrmecodia tuberosa Jack, entity ‘versteegii’; UPNG 3, 479, (ibid: 287) 
SAPINDACEA 
Rhysotoechia - nov.ined.; WT 12,694 (type) 


MONOCOTS 


PANDANACEAE (all dets. by K.-L. Huynh) 

Freycinetia acuta Huynh, sp. iy in oe WT 12,875 (type) 
Freycinetia biroi Warb,; WT 12 

Freycinetia craterensis Huynh, | nov. y le WT 11,955 (type) 
Freycinetia lagenicarpa Warb.; WT 11,7 

Freycinetia obtusiacuminata Huynh, ee nov.in press; WT 12,107 (type) 


DISCUSSION 


The flora of the Crater Mt. Wildlife Management Area (CMWMA) has been addressed, at 
least in part, by a succession of papers based on the 1997-1998 plant surveys. These 


244 SIDA 19(2) 


surveys were originally intended as part of a comprehensive assessment through a con- 
tinuous elevational sequence, on a schedule eventually to culminate in the preparation 
of a plant identification guide. Because of the truncation of our itinerary in duration and 
scope, the major objectives are now out of reach. However the existing documentary 
base provides considerable opportunities for future investigators. As noted earlier, the 
lowland environment at Crater Mt.is still relatively unsurveyed and almost certainly pre- 
sents rich prospective opportunities for discovery (Takeuchi 1999). Even though some 
work has already occurred in the low elevation zone, previous efforts have been brief 
and spatially restricted. The existing herbarium documentation is also highly skewed, 
with past collectors tending to focus on easy-to-obtain taxa such as understory herbs 
and shrubs.A conscious attempt to secure fertile gatherings of canopy/subcanopy phan- 
erophytes and high epiphytes should prove rewarding. The previously reported discov- 
eries of new plants from the ecotone contact with the Pio alluvial plain, are suggestive 
of the possibilities. 

The Crater Mt.tract is positioned on the leading edge of the Australian craton, form- 
ing part of a southern geoprovince which is floristically depauperate in comparison to 
the speciose northern orogen of New Guinea (inter alios Pigram & Davies 1987;Welzen 
1997).The contrasting species content on north-south orientations has been attributed 
to the orogenic etiology of northern New Guinea environments, and to increased spe- 
ciation induced by the orogeny (Balgooy et al. 1996; Welzen 1997). At least part of the 
difference however, is an artifact of the uneven state of plant exploration in PNG, of which 
a salient element is the lesser number of collections from the southern districts. Since 
mountain-building processes appear to drive floristic endemism in New Guinea, the high- 
est species density from its austral geoprovince should be expected on the latter's north- 
ernmost margin, where the plate has been forced upwards by tectonic collision with the 
island arc terranes. The novelties from Crater Mt.can be rationalized on this basis. Further 
exploration of the south descending side of the Dividing Ranges can be expected to 
produce similar results. 

The CMWMA represents a previously unexplored expanse of wilderness forest. Re- 
cent discoveries from this tract are indicative of the necessity of surveying such areas as 
part of an overall program for developing veridical phytogeographic estimates of the 
New Guinea flora. The existing CMWMA checklist consists of a diverse mixture of 
allochthonous and autochthonous taxa, including many distributional records. While 
there are bases for anticipating that future findings will follow patterns suggested by 
geological correlation, new refinements are likely to arise fram exhaustive work on the 
Papuan side. Additional surveys toward the lowland alluvial zone could eventually con- 
nect to the elevational anomalies discerned at Lakekamu (cf. Takeuchi & Kulang 1998; 
Takeuchi in press). At the latter site for example, Palmeria gracilis Perkins was found in 
lowland communities corresponding to Australian provenances, though the species is 
ordinarily strictly montane in Papuasia. A complete elevational series may be demon- 
strable at Crater Mt., where P gracilis is already documented from its more characteristic 
montane stations. 


TAKEUCHI, ADDITIONS TO THE FLORA OF CRATER MOUNTAIN 245 


Several generalizations can be offered from our ongoing inventory work in wilder- 
ness areas, many of them previously little-explored or unexplored. 1) Substantial num- 
bers of undiscovered species are sequestered in lowland habitats, even in accessible 
places, having been unrecorded because of the poor documentation from low eleva- 
tions (Takeuchi 2000). 2) Although montane areas are better documented than lowland 
ones, additional discoveries are more likely to result from working the south-descend- 
ing ranges, on the margin of the Austro-craton, rather than the Mamose-descending 
side. To be sure, virtually any sort of serious effort will result in substantive discovery. 
Even at this late date in Papuasian botany, every major expedition comes back with new 
collections. 3) It is an imperative on inventories, for survey botanists to collect uncritically 
by taking everything encountered, in multiple bers rather than being selective (Pipoly 
pers.comm.). Many of the most consequential findings from current investigations were 
of taxa whose significance was only revealed in the herbarium. This is especially true of 
novelties from speciose genera. If investigators are too choosy in what they gather, these 
sort of records will be missed. 4) In traditional PNG cultures, there is a characteristic em- 
phasis on vegetative markers for plant identification, which not unexpectedly results in 
highly erratic nomenclatural systems. The significance and discriminatory value of ver- 
nacular names are considerably overrated. At Crater Mt.,this is exemplified by the Pawaian 
‘way-e-be,’ which was described as monotypic by village guides, but actually encom- 
passes ranthera, Horsfeldia,and Myristica;in fact virtually the entire Myristicaceae. 
Numerou mples of comparable imprecision were documented during the CMWMA 
inventory. Especially when compilations are based on limited sampling, the local names 
will appear to be specific merely because of discontinuities in the polling. While it is 
often standard practice for investigators to report local names in revisionary work, these 
reports have little to acquit themselves unless they are cast in the context of a compre- 
hensive census accompanied by comparative evaluation of the vouchers. Our experi- 
ence is that such inquiry will reveal numerous examples of nomenclatural circumscrip- 
tion grossly incompatible with formal science. The quirkiness of local naming systems 
has also been shown in the ethnobotanical polling from the recent Josephstaal surveys 
(Takeuchi 2000). 

The Integrated Conservation and Development (ICAD) strategy at Crater Mt. con- 
joins community-based conservation initiatives with low-impact socioeconomic devel- 
opment. Nearly all conservation programs in Papua New Guinea are now founded on 
this principle. As one of the largest of the ICAD experiments, the CMWMA also ranks 
among the floristically richest sites presently subsumed under this operational para- 
digm.Whether or not the ICAD philosophy can achieve programmatic success in places 
such as the CMWMA is still open to question (cf. Saulei & Ellis 1998). What the Crater 
surveys demonstrate is that the long-term viability of CAD at Crater Mt. could have dra- 
matic implications for a remarkable and precinctive flora. The nature of future scientific 
contributions to be forthcoming from this area is in many respects an imponderable, 
but the results achieved thus far clearly point to a considerable promise for further floristic 


discovery. 


246 SIDA 19(2) 


ACKNOWLEDGMENTS 


The botanical surveys of Crater Mt. were supported by principal funding from the Liz 
Claiborne and Art Ortenberg Foundation, and by the John D.and Catherine T.MacArthur 
Foundation. Staff of the Research and Conservation Foundation of PNG (Robert Bino, 
John Ericho, Paul Hukahu, Paul lgag, and Arlyne Johnson) assisted with logistics and com- 
munity liaison. 

The Lae National Herbarium provided facilities for processing and identification of 
collections. Curators Emily Wood (A) and Debra Trock (BRIT) facilitated loans of critical 
specimens. Jin Komatsuzaki and Keiko Hanlon provided the Japanese translation. 

| especially thank John Pipoly Ill for his constructive criticism and Barney Lipscomb 
for his generous assistance. 


REFERENCES 


Ba.cooy, M.M.J. van, P. Hovenkame, and P. van Wevzen. 1996. Phytogeography of the Pacific — 
floristic and historical distribution patterns in plants. In: A. Keast and S. Miller, eds. The 
origin and evolution of Pacific island biotas: New Guinea to Eastern Polynesia: pat- 
terns and processes. SPB Academic Press, Amsterdam, the Netherlands. Pp. 191-214. 

Harte, [.G.and L.M. Perry. 1973.A provisional key and enumeration of species of Syzygium 
(Myrtaceae) from Papuasia. J. Arnold Arbor. 54:160-227. 

Huxtey, C.R.and M.H.P. Jess. 1993. The tuberous epiphytes of the Rubiaceae 5:A revision of 
Myrmecodia. Blumea 37:271-334. 

Jess, M.H.P. 1987. Key to Piper species in Papuasia. In: RJ. Johns. The flowering plants of 
Papuasia. Dicotyledons. Part 1: Magnoliidae. PNG University of Technology Forestry 
Department, Lae. Pp. 103-106. 

LeenHouts, PW. 1955a. The genus Canarium in the Pacific. B.P. Bishop Mus. Bull. 216:1—-53. 

. 1955b. Florae Malesianae Precursores X!. New taxa in Canarium. Blumea 8: 
181-194. 

. 1956. Burseraceae. Flora Malesiana ser. |, 5:209-296. 

. 1959. Revision of the Burseraceae of the Malaysian area in a wider sense. Xa. 
Canarium Stickm. Blumea 9:275-647. 

Nietsen, |.C. 1992. Mimosaceae (Leguminosae-Mimosoideae). Flora Malesiana ser. |, 11: 
]-226. 

_ T. BARETTA-Kuipers, and P. Guinet. 1984. The genus Archidendron (Leguminosae- 
Mimosoideae). Opera Bot. 76:5-120. 

Picram, C.J. and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea 
orogen. J. Austr. Geol. Geoph. 10:193-211. 

SauLel, S.M. and J.-A. Evuis (eds). 1998. The Motupore Conference: ICAD practitioners’ views 
from the field.A report of the presentations of the second ICAD conference. Motupore 
Island (UPNG), Papua New Guinea 1-5 September, 1997. Dept. of Environment & Con- 
servation, Papua New Guinea/United Nations Development Programme PNG/93/G31 
Biodiversity Conservation & Resource Management. 


TAKEUCHI, ADDITIONS TO THE FLORA OF CRATER MOUNTAIN 247 


STeenis, C.G.G.J. VAN. 1972. Addenda, corrigenda et emendanda. Canarium. Flora Malesiana 
ser. |, 6:921-928. 

TAKEUCHI, W. 1999.New plants from Crater Mt., Papua New Guinea, and an annotated check- 
list of the species. Sida 18:961-1006. 

2000. A floristic and ethnobotanical account of the Josephstaal Forest Man- 
agement Agreement Area, Papua New Guinea. Sida 19:1-64. 

er Rhodod I hi (Ericaceae) from mainland New Guinea. 
A distributional record and new subspecies. Edinb. J. Bot. 57(3). 

.In press. New and noteworthy plants from recent exploratory surveys in Papua 
New Guinea, 7. Edinb. J. Bot. 58(1). 

and J. KuLANG. 1998. Vegetation Part 2: Botanical survey. In: A. Mack, ed. A bio- 
logical assessment of the Lakekamu Basin, Papua New Guinea. Rapid Assessment Pro- 
gram Working Papers no. 9. Conservation International, Washington, D.C. Pp. 36-39, 
105-130. 

Weuzen, PC. van. 1997. Increased speciation in New Guinea:tectonic causes? In:J.Dransfield, 
M.J.E. Coode, and D.A. Simpson, eds. Plant diversity in Malesia Ill. Proceedings of the 
Third International Flora Malesiana Symposium 1995. Royal Botanic Garden, Kew. Pp. 
363-387. 


248 SIDA 19(2) 


BOOK REVIEW 


sai oo BULMAHN and Jack Becker. 1998. American Garden Literature in the 
mbarton Oaks Collection (1 785- 1900): From The Newengland Farmer to 
hy. (ISBN 0-88402-253-6, pbk.). Dumbarton 
Oaks Research Library and Collection, , Washington D.C. U.S.A. $35.00 pbk. ix + 243 
pp., b/w illustrations. 


This annotated bibliography summarizes 410 books on horticulture, botany, gardening, garden 
esign, cemeteries, parks, rural architecture, and landscape architecture in America. The titles were 
published in the United States between 1785 and 1900 and are housed in the collections of the 
Garden Library of Dumbarton Oaks. Traditionally, the library's dominant holdings have been in the 
areas of English, French, and Italian garden literature, in keeping with the interests of garden histo- 
rians. In recent years, as increasing numbers of scholars have addressed the American landscape, 
saa at Oaks expanded its holdings in 19th-century American literatur 
though, as noted by the authors, the traditional form or biography no longer has the 
same i it did in the past, because more and more libraries place their complete hold- 
ings online, and printed pibiodap hier are a valuable for scholars because they often provide 
importa ory essays,annotations, illustrations, indices, and other 
supplement material. |t is precisely in aspect that the value of the present bibliography lies. 
ok is organized in five sections: I. Introduction; Il. Annotated list of titles; II Chrono- 
ese list of titles; IV. Index; and V. Bibliography. The introductory essay presents a history of the 
literature of American gardening, HOIIeUtUra) and landscape, cemetery and park design. The an- 
notations are succinct but comments on each title’s contents. The index is extensive and 
provides one of the greatest ie of the volume. One wishes for more and better illustrations 
(none are in color); however, publishing constraints may have placed unwanted limitations on the 
bibliographers. 
The bibliography of A den | t th { Oaks ene 1900) 
will be a useful addition to the iibeary of a scholar of hag genre. Soe; H. Swai 


SIDA 19(2): 248. 2000 


THE INTRIGUING CASE OF CYPRIPEDIUM CROSSII, ITS 
PRIORITY OVER CYPRIPEDIUM CALLOSUM AND IIS 
TRANSFER TO THE GENUS PAPHIOPEDILUM 


Guido J.Braem Karlheinz Senghas 
Schlechter Institute and Botanischer Garten der Universitat Heidelberg 
California Academy of Sciences Im Neuenheimer Feld 340 
Naunheimer Str. 17 D-6900 Heidelberg, GERMANY 
D-35633 Lahnau, GERMANY 
ABSTRACT 


Cypripedium crossii Morren was first mentioned in La Belgique Horticole for 1865 and described in 
the same journal for 1883. Its priority over Cypripedium callosum Rchb.f. 1886 is discussed and es- 
tablished. The taxon and two of its known varieties are transferred to the genus Paphiopedilum 
Pfitzer. 


L ft} 


ee Ae iy callosum, crossii 


Key Worbs: Orchidaceae, Cyr ypripedium, Pap 
barbatum, sublaeve, potentianum, Taxonomy, Systematics, Orchid Hybrid Registration International 
Code of Botanical Nomenclature, Saint Louis Code. 


RESUMEN 


Cypripedium crossii Morren se menciond Bol primera vez en La Belgique Horticole en 1865 y se 
describid en la misma ite: en ee Se discute y establece su peHOnaas sibs Cypripedium callo- 
sum Rchb.f.1 1886.E LOLE la tty MA ar Paphi iopedilum 


Pfitzer. 


INTRODUCTION 


During the preparatory work for the third volume of Braem, Baker & Baker, The genus 
Paphiopedilum — Natural History and Cultivation, the literature regarding Paphiopedilum 
callosum (Rchbf.) Stein was reviewed. The taxon referred to as Cypripedium crossii Morren 
has been interpreted as a synonym of Paphiopedilum barbatum by Braem (1988) and 
considered to be a nomen nudum within the synonymy of Paphiopedilum callosum by 
Cribb (1987, 1998).A review of the original literature has conclusively revealed that both 
interpretations are erroneous. 


DISCUSSION 


Cypripedium crossii was first mentioned by Morren (1865) in a short article in volume 15 
of La Belgique Horticole,21 years before the publication of Cypripedium callosum (Rchbf. 
1886).In his 1865 article, Morren simply states that the Cypripedium originates from Peru 
and that it carries the name of its discoverer, Mr. Cross. No description or any further 
information is made available. However, plate 17, which is part of the article, shows a 


SIDA 19(2): 249 -255. 2000 


250 SIDA 19(2) 


plant labeled as Cypripedium crossii. This illustration leaves no room to doubt that the 
taxon published by Morren as Cypripedium crossii is identical with the plant described as 
Cypripedium callosum by the younger Reichenbach in 1886. The Morren “note” has been 
interpreted as insufficient to be regarded as a valid and effective publication of Cypripe- 
dium crossii as an autonomous taxon, an interpretation generally based on article 42.3 of 
the International Code of Botanical Nomenclature (ICBN) generally referred to as the 
“Code.” Article 42.3 (taken from the Saint Louis Code [Greuter et al. 2000], which is the 
effective version) reads, 


= 


"Prior to I January 1908 an illustration with analysis, or for non-vascular plants, a single figure showing 
details aang identification, is acceptable, for the purpose of this article, in place of a written description 
or diagnosis 


And article 42.4 clarifies what is to be understood as an euuee 


“For the purpose of Art. 42, an analysis is a figure or group of figure separate from the main 
illustration of the plant (though usually on the same page or ee showing: details aiding identifica- 
tion, with or without a caption.” 


The Morren note was accompanied by a color plate of Cypripedium crossii (Fig. 1). There 
can be no doubt about the identity of the plant published as Cypripedium crossii by 
Morren in 1865 is identical with the taxon we have hitherto referred to as Paphiopedilum 
callosum (Rchb.f) Stein. The plate contains details that allow for the identification; two 
flowers are shown. The color plate clearly depicts the different aspects of the plant and 
flower such as shape and tesselation of leaves, shape and color of all parts of the corolla, 
from the front as well as from the back, detailed shape of staminodal shield, the ovary, 
and the floral bract.In other words all aspects relative to the identification of a slipper 
orchid. Thus the plate satisfies the requirements of Art. 42.4, especially as that article 
clearly states (see above) that the “figure” or“group of figures” do(es) not necessarily have 
to be“separate from the main illustration of the plant.” If this were not so, the inclusion of 
the word “commonly” in article 42.4 would make no sense. 

It is ludicrous to disqualify the publication of Cypripedium crossii by Morren in the 
scientific journal La Belgique Horticole knowing that the valid code accepts publication 
in trade catalogues or non-scientific newspapers before 1 January 1953, and in seed- 
exchange lists before 1 January 1973 (see Code, Article 30.3). As every botanist knows, a 
publication in a seed list generally means simple mention of the botanical (Latin) name. 
Thus, Cypripedium crossii is for all purposes to be regarded as validly and effectively pub- 
lished in 1865. 

In 1883, in volume 33 of the same journal (La Belgique Horticole), Morren wrote an 
extensive article entitled “Cypripedium barbatum, Lindl.and its major varieties, crossi [sic], 
warnerianum, etc." This article was illustrated with a colored plate (Fig 2). 

The plate reveals that the plant Morren described is identical with the plant he had 
depicted in 1865, thus the plant we have hitherto generally addressed as Paphiopedilum 
callosum (Rchb.f) Stein. Morren writes, “Cypripedium crossi [sic.] has a very wide, some- 
what trilobed dorsal sepal which is white, veined with green and crimson. It is transversally 


251 


divided in the middle by a crimson stripe. The petals are half-green and half-white with 
green veins and rose extremities. The lip is dark rose-brown." 

Thus, in 1883, Morren published a description of Cypripedium crossii, although he 
considered this plant to be a variety of Paphiopedilum barbatum Lindley. 


CONCLUSIONS 


In view of the facts delineated above, there is no reason to deny Cypripedium crossii 
Morren full taxonomic validity.Whereas the validity of Morren’s publication of 1865 could 
possibly be a matter of discussion (but see below), the taxon was distinctly described in 
Morren’s 1883 article, and clearly identified as an autonomous taxon identical to the 
species hitherto generally referred to as Paphiopedilum callosum (Rchb.f) Stein. Both 
Morren publications predate the publication of Cypripedium callosum by the younger 
Reichenbach and therefore, Cypripedium crossii Morren is to be given priority in accor- 
dance with the rules of nomenclature. The taxon is to be transferred to the genus 
Paphiopedilum. The authors are well aware of the fact that conservative growers and 
hybridizers will argue that the name “Paphiopedilum callosum” should be retained be- 
cause a multitude of hybrids has been registered indicating “Paphiopedilum callosum” to 
be part of their ancestry. This argument must be rejected. The registration of orchid hy- 
brids (and any other hybrids for that matter) is no concern to botanical taxonomy. Fur- 
thermore, the registration authority for orchid hybrids (Royal Horticultural Society, Lon- 
don) is by no means a taxonomic ruling body. Furthermore, if the horticultural argument 
were to be followed, the name “Cypripedium” would have to be re-instated for the genus 
Paphiopedilum (and other genera), etc. Last, but not least, the identity of (at least) some 
of the hybrids must be questioned. If we would reject a valid name because its use is 
considered to be inconvenient to horticulture, it would, indeed be best to ignore all 
rules of botanical taxonomy. 


TAXONOMY 
aaa apt crossii (Morren) Braem & Senghas, comb. et stat. Nov. Basionym: Cypripe- 
crossii Morren in Belgique Horticole 15:226,t. 17, 1865; Belgique Horticole 33:96-98 [97— 
a .7,1883. 

Cypripedium callosum Rchbf, Gard. Chron. ser. 2, 26:326. 1886. Cordula callosa (Rchbf.) 
Rolfe, Orchid Rev. 20:2. 1912. 

Cypripedium schmidtianum Kraenzl., Bot. Tidsskr. 24:13. 1901. Paphiopedilum callosum 
var. schmidtianum (Kraenzl.) Pfitzer in Engler, Pflanzenr. IV. 50. Orchidaceae- 
Pleonandrae: 93. 1903. 

Paphiopedilum callosum var. angustipetalum Guillaumin, Bull. Soc. Bot. France ser. 4, 
24:551.1924. 


The decision to transfer the following two varieties was a difficult one as their botanical 
validity is by no means established. The differentiation between the type variety (thus 
the autonym Paphiopedilum crossii var. crossii) and Paphiopedilum crossii var. sublaeve 


SIDA 19(2) 


252 


pripedium - Crossit. 


(Cy 


le, 1865. 


ay 


Fic. 1.P, 


La Belg. hort. 


X83, PL VII, CYPRIPEDIUM BARBATUM var. 


Fic. 2. P, phi 4 ii (M 1D ° Senghas, fi RA pala Uartical 1883. 


254 SIDA 19(2) 


consists merely in var. sublaeve having somewhat smaller flowers with a smaller dorsal 
(cf. Cribb 1987, 1998). Reichenbach fil. (1888), in his original publication of var. sublaeve, 
simply notes that the plant came out of a batch of “Cypripedium callosum,” and that “it 
might be supposed to be a natural hybrid.” Cribb also states (loc. cit.) that the petals of 
var. sublaeve “usually bear warts only on the upper margin." This, however, also applies to 
the plant he depicts as Paphiopedilum callosum var. callosum on page 333 of his book 
(Cribb 1998). 

It is amazing that the validity and effectiveness of the Reichenbach fil. publication, 
rendered in a British horticultural newspaper and in English, without any description or 
illustration, has never been questioned, whereas the publication of Cypripedium crossii 
by Morren, admittedly in a Belgian scientific journal, and admittedly in French, should be 
denied acceptance although Morren’s plant can be positively identified by the explicit 
illustration that is part of the publication. 

Variety (?) potentianum differs merely by the lack of the marginal warts on the up- 
per margin and its narrower dorsal sepal (see Cribb 1998). 

It may very well be argued that these are phenotypes that are well within the nor- 
mal range of a natural species. 


Paphiopedilum crossii var.sublaeve (Rchbf) Braem & Senghas,comb.et stat. NOV. Basionyo: 
Cypripedium oa var. sublaeve Rchb.f, Gard. Chron. Ser. 3, 3:331. 1888. Paphiopedilum cal- 
losum subsp (Rchb.f) Fowlie, Orchid Digest 36:145.1972;38:187.1974.Paphiopedilum 
sublaeve (Rchbf) Fowlie, Orchid Digest 43:224. 1979. Paphiopedilum callosum var. sublaeve 

Rchb.f) Cribb, Genus Paphiopedilum 188. 1987. 


=a 


Paphiopedilum thailandense Fowlie, Orchid Digest 43:220. 1979, nomen nudum. 
? Cypripedium callosum var. warnerianum T. Moore in Warner, Select Orchid. Pl, 3, t. 11, 


Paphiopedilum crossii var. potentianum (Gru & R6th) Braem & Senghas, comb. et stat. 
nov. Basionym: Paphiopedilum potentianum GruB & Réth., Caesiana 5:39. 1995. Paphiopedilum 
callosum var. potentianum (Gru & Réth) Cribb, Genus Paphiopedilum, 2nd ed. 337.1998. 


ACKNOWLEDGMENT 


The authors are indebted to Peter H. Peeters for making the slides of the pertinent plates 
available. 


REFERENCES 


Braem, GJ. 1988. Paphiopedilum — A monograph of all tropical and subtropical Asiatic 
slipper-orchids. Brucke Verlag, Hildesheim, Germany. 

BraeM, G.J.,C.O. Baer, and M.L. Baker. 1998. The genus Paphiopedilum — natural history and 
cultivation, vol. 1. Botanical Publishers, Miami, FL. 

Braem, G.J.,C.O. Baker, and M.L. Baker. 1999. The genus Paphiopedilum — natural history and 
cultivation, vol. 2. Botanical Publishers, Miami, FL 


255 


Bram, G.J., C.O. Baker, and M.L. Baker (in preparation). The genus Paphiopedilum — natural 
history and cultivation, vol. 3. Botanical Publishers, Miami, FL. 

Crips, PJ. 1987. The genus Paphiopedilum. Collingridge, London, UK. 

Cris, PJ. 1998. The genus Paphiopedilum. Natural History Publications (Borneo), Kota 
Kinabalu, Sabah 

Greuter, W., J. McNeil, E.R. Barrie, H.M. Burber, V. DeMouLIN, T.S. FiGueiras, D.H. Nico.son, O.C. SILVA, 
J.E. Skoa, P. TREHANE, N.J. TURLAND, and D.L. Hawkswortu. 2000. International code of botani- 
cal nomenclature (Saint Louis Code). Koeltz Scientific Books, K6nigstein, Germany. 

Gruss, O and J. RotH. 1995. Paphiopedilum potentianum, a new species from Thailand. 
Caesiana 5:9. 

Moreen, C.J.E. 1865.Le Franciscea lindeniana Pl.et le Cypripedium crossii. Belgique Horticole 
15:226,t.17. 

Moreen, C.J.E. 1883. Note sur le Cypripedium barbatum, Lindl. et ses principales varietes, 
crossi, warnerianum, etc. Belgique Horticole 33:96—98, t. 7. 

REICHENBACH, H.G. 1886. Cypripedium callosum, n. sp. Gard. Chron. ser. 2, 26:326 

REICHENBACH, H.G. 1888. Cypripedium callosum (Rchb.f) sublaeve, n. var. Gard. Chron. ser. 3, 
3:331. 

Stein, B. 1892. Stein's Orchideenbuch. Breslau, Germany. 


256 SIDA 19(2) 


BOOK REVIEW 


Haron W. Ketter and Karl L. Braun, 1999. Myxomycetes of Ohio: Their Systematics, Biol- 
ogy, and Use in Teaching. (ISBN 0-86727-133-7, pbk.).Ohio Biological Survey Bul- 
letin New Series volume 13 Number 2 (ISSN 0078-3994). Ohio Biological Survey, 

1315 Kinnear Road, Columbus, OH 43212-1192,U.S.A.(615-292-9645, 614-688-4322 
fax; http://www-obs.biosci.chio-state.edu). $35.00 pbk.(Wire-O Binding or Perfect 
Binding) xvi + 182 pp., 16 color plates. 


“Half animal-half plant!” A fungus and a protozoan? A naked mass of colorful, slimy,“snot-like” pro- 
toplasm, sometimes several inches or more across, creeping in mass or in a vein-like network over 
rotten logs and leaves in the woods - or in the lab on a bowl of oatmeal. Under the microscope 
beautifully ornamented globe-shaped balls, the wind-borne spores that in dew break open to re- 
lease minute, swarming, sperm-like creatures that swim with rapidly moving flagellar paddles, these 
zoospores changing into white blood cell-like amoebas that glide and feed by engulfing bacteria 
and other minute morsels, these reproducing themselves into an army of amoebas that eventually 
reunite and fuse into masses of streaming, multinucleated siete he like the beautiful 
butterfly that emerges from the drab cocoon, may change overnight into dozens of minute, mar- 
velously structured and colorful spore producing bodies that we can see easily with a 10 X hand 
lens and sd contain the tiny balls that we saw first under the microscope. 

What are Myxomycetes? Mycology has been the traditional home for this mysterious group 
of “animal- ie plants.” How do mycologists study this living stuff where there is no such thing as an 
individual organism that we can separate and count or experiment with such as an ant,a single 
great ape, or a single bean plant? Does it really matter where in their hierarchical categories the 
biosystematists place these creatures? Wherever Myxomycetes are Classified, they are wonderfully 
fascinating life 

One of the most important kinds of biosystematic research is that which results in practical, 
useful products that can be used at a local level by teachers, students, and other scientists to learn 
about and identify the specific organisms in their immediate surroundings. This is especially true 
for microorganisms, which are much less popular than macroorganisms and have had very few 
useable publications written about them at a local level. 

Keller and Braun's book is the kind of product that gives practical, useful, thorough, colorful 
one about the biology, morphology, and one of Myxomycetes. It also includes valu- 
able i about tech RIGUE> for Staying these fantastic ena | recommend this 
book especially because it not only pre ell-organized scientific data about Myxomycete 
species, but also because of its Beene uch otk the authors, their teachers, their students, and 
other scientists who have contributed to knowledge of Ohio Myxomycetes, But this book is valu- 
able in a much wider area than the state of Ohio. Nearly 35 % of the world’s 600 species and 57 
genera of eae are included, and the contributions of the scientists mentioned has had 
influence kane e 

Every state eae an agency like the Ohio Biological Survey that supports field research on 
groups of A biota and publication of useful, interesting, practical information about the specific 
things that live in our own backyards.Where in the United States of America is there a county that 
has all of its species biodiversity catalogued? In what locality does anyone know all of the living 
things, ean es the microorganisms, that one might find in a handful of soil from nearby parks, 


prairies, woodlands, schoolyards, or even flower pots? Keller and Braun have produced an excellent 
book that is a necessary step toward achieving the goal of an“all-taxa inventory” of at least a small 
part of the world.—Joe F Hennen, Resident Research Associate, Botanical Research Institute of Texas, 


Fort Worth, TX 76102-4060, U.S.A. 


SIDA 19(2): 256. 2000 


USE OF VARIETY AND SUBSPECIES AND 
NEW VARIETAL COMBINATIONS FOR STYRAX 
PLATANIFOLIUS (STYRACACEAE) 


B.L. Turner 
Section of Integrative Biology 
niversity of Texas 
Austin, TX 78713, U.S.A. 


Guy L.Nesom 


North ae Botanical Garden 
Uni of North Carolina 
cel an ne 27599-3280, U.S.A. 


ABSTRACT 


The term “variety” has historical precedence over “subspecies” and is either prescribed or recom- 
mended by the ICBN as the rank to be first used in the description of infraspecific taxa. The rank 
"subspecies" is then used to cluster related varieties. Accordingly, to replace earlier combinations at 
subspecific rank, the following new combinations in Styrax platanifolius are proposed: var. mollis 
(PW. Fritsch) B.L. Turner, comb. et stat. nov,; var. texanus (Cory) B.L. Turner, comb. et stat. nov.;and var. 
youngiae (Cory) B.L. Turner, comb. et stat.nov.No subspecies are recognized here in S. platanifolius. 


RESUMEN 


El término“variedad" precede histéricamente al de"subespecie’y es ordenado o recomendado por 
el ICBN como el primer range a ser usado - Ie descripcién de taxa infraespecificos. El rango 
“subespecie” lacionadas. De acuerdo con ésto,se proponen 
ie sjguleiites nuevas ; combinaciones se reemplazar combinaciones previas en el rango 

ollis (PW.Fritsch) B.L.Turner,comb.et stat.nov.;var.texanus 
(Cory). B.L. Turner, comb. et stat. nov.; y var. youngiae (Cory) B.L. Turner, comb. et stat. nov. No se 
reconocen subespecies aqui en S. platanifolius. 


Fritsch (1997) has provided a much needed revision of Styrax for Mesoamerica, Mexico 
and western Texas, recognizing 19 species. One of these, S. platanifolius Engelm. ex Torr., 
was treated as having five subspecies, bringing to 24 the number of formal taxa of Styrax 
recognized for the region concerned. Fritsch recognized infraspecific taxa as “subspe- 
cies” rather than “varieties,” noting (p. 711) that his usage was “in accordance with the 
concepts of Hultén (1967) and Thorne (1978),” who “use the subspecies category for 
infraspecific taxa that are geographically as well as morphologically distinct." 

Of course, most current botanists who employ “variety” also use it in reference to 
infraspecific taxa that are geographically and morphologically distinct, and we do not 
accept what is essentially the equating of these two categories of infraspecific classifica- 
tion. The use of variety in plant taxonomy for the first infraspecific rank dates back to 
Linnaeus and has historical precedence over the term subspecies. More significantly, 


SIDA 19(2): 257 — 262. 2000 


258 SIDA 19(2) 


this usage of variety is set forth by the current International Code of Botanical Nomen- 
clature (Greuter et al. 2000) in the same format and language as other conventions that 
are clearly prescriptive. 


Article 4.1."The secondary ranks of taxa in descending sequence are tribe (tribus) between family 
and genus, section (sectio) anes series (series) between genus and species, and variety (varietas) 
and form (forma) below species. 


Use of the “sub” prefix provides added ranks. 


Article 4.2."If a greater number of ranks of taxa is desired, the terms for these are made by adding the 
prefix sub- to the terms denoting the principal or secondary ranks. A plant may thus be assigned to 
taxa of the following ranks (in descending subregnum, divisio or phylum, sub- 
divisio or subphylum, classis, subclassis, ee subordo, familia, subfamilia, tribus, subtribus, genus, 
subgenus, sectio, subsectio, series, subseries, species, subspecies, varietas, subvarietas, forma, subforma.” 


Philosophical and interpretive differences regarding use of infra-taxon categories are 
magnified by this tension in the ICBN: variety and/or forma are me ne to be used 
first in describing infraspecific taxa (Article 4.1), but sub the term first in 
hierarchical rank below species (Article 4.2). Use of ipeaes and variety, however, is 
analogous to that for general use of kingdom and division, class and order, family and tribe, 
and genus and section.|n each of these pairs the first used sub-rank (the second term) is 
code-prescribed as such (4.1), with the orthographically subsidiary and immediately hi- 
erarchical ranks subkingdom, subclass, subfamily, and subgenus generally used for an ad- 
ditional rank (4.2),as is subspecies. 


Varieties may be clustered by use of the subspecies category 

Varieties are recognized within a species when it is desirable to refer by name to mor- 
pho-geographically differentiated entities comprising that species. In our concept and 
experience, varieties usually are closely similar allopatric entities that intergrade over a 
relatively short distance in regions of contact (as opposed to gradual, broadly regional 
intergradation), if they intergrade at all.In addition to morphology and geography, other 
factors may be used in evaluation of taxonomic status of such entities—genetic diver- 
gence, likelihood of natural hybridization, and fertility of hybrids (Stuessy 1990). 

In a species where several varieties are recognized, two or more varieties may be 
grouped within a subspecies.|n this sense, use of the subspecies rank may point to larger 
patterns of variation and/or coherence within the species. This use of infraspecific cat- 
egories finds support in the ICBN, which implies that the term subspecies is used for 
clustering varieties. 


Recommendation 26A.2."A subspecies not including the type of the correct name of the species 
should, where there is no obstacle under the rules, be given a name with the same final epithet 
and type as a name of one of its subordinate varieties.” 

Use of the subspecies rank in clustering varieties is not pervasive, but it is currently 
found across various families and genera in the North American flora (e.g., Agastache, 
Arenaria, Arnica, Artemisia, Cerastium, Chamaecrista, Chrysothamnus, Ericameria, Erigeron, 


259 


Eriogonum, Heterotheca, lpomopsis, Lathyrus, Lupinus, Machaeranthera, Monarda, Prunus, 
Ptelea, Ruellia, Salix, Salvia, Scutellaria, Sidalcea, Silene, Solidago, Streptanthus, 
Symphyotric hum, and Tetramolopium). 


Other perspectives on the use of infraspecific categories 

The issue of “variety vs. subspecies’ in infraspecific classification has been discussed re- 
peatedly. Cronquist (1988) and Stuessy (1990) gave detailed overviews of the issue and 
Hamilton and Reichard (1992) provided a review of current practice in the use of in- 
fraspecific categories. Our commentary does not break new ground, but it emphasizes 
primary considerations and it is a reminder that usage of these categories remains in- 
consistent and commonly without explicit rationale. 

Views similar to ours have been well-expressed by Kapadia (1963) and Holmgren 
(1994). In a contrasting view, Raven (1974) proposed to simplify infraspecific terminol- 
ogy by using only the term subspecies, pomenc easy equating the term variety, this 
proposal accompanied by detailed lizat n the ICBN. Another 
form of this latter solution is to use apece: as the first category A eee taxa— 
then to use varieties (at lower rank) for subsequent subdivisions of subspecies (see Stuessy 
1990, Fig. 12.1). For Thorne (1978, p. 190), “Genetic variants without well-defined geo- 
graphic ranges are treated as varieties ...."Such proposals, however, to formally displace 
“variety” as the first infraspecific category have not been accepted, presumably because 
many botanists find utility in maintaining two classificatory units at infraspecific rank 
and because the basis for use of “variety” is historical and currently codified. 

The rank of variety has been used to describe taxa over a range of evolutionary and 
morphological differentation. Some taxa are more strongly differentiated than others. 
Traditional views of boundaries between taxa at specific and infraspecific rank also differ 
among genera and families. Replacing variety with subspecies would not change this. 

The International Code of Zoological Nomenclature (International Commission on 
Zoological Nomenclature 1999) does not provide for the term “variety” for classificatory 
purposes. The latter term, as used by most botanical taxonomists and as prescribed by 
the ICBN, is essentially equivalent to the subspecies rank of zoologists. This is acceptable. 
The botanical code is for plant workers, the zoological code for animal workers and the 
two codes need not become one. Actually, it is informative to see the term “variety” in a 
title or abstract: one knows that the organisms concerned are most likely plants. 

It might be argued that the term “variety” has been misapplied by various workers, 
especially horticulturists and plant breeders, to designate mere forms. But most profes- 
sional plant taxonomists use the term “forma” for such population variants, while the 
International Code of Nomenclature for Cultivated Plants (Trehane et al. 1995) refers to 
such an individual plant or genetic strain as a “cultivar” (cultivated variety). The horticul- 
tural taxonomists coordinate their taxonomy with the ICBN, providing an adjunct system, 
not one that contradicts (Brickell & Trehane 1997). In short, horticultural usage is not a 
tenable rationale for generally adopting the rank of subspecies to the exclusion of variety. 


260 SIDA 19(2) 


In contrast to these more pragmatic arguments for using variety as first choice of 
infraspecific rank, guidance of the ICBN is more ambiguous. Is the nomenclatural con- 
vention codified in Article 4.1 a prescription, or is it merely a suggestion or recommen- 
dation? In either case, what is its relationship to Article 4.2? And what bearing, if any, 
does Article 5.1 have on this interpretation? 

Article 5.1. “The relative order of the ranks specified in Art. 3 and 4 must not be altered (see Art. 
33.7 and 33.8).” 

Articles 4 and 5 are not included among those whose conditions must be met for valid 
publication, according to Article 32. But, then, how should they be interpreted? What 
bearing, if any, does Article 2.1 have on this interpretation? 

Article 2.1.“Every individual plant is treated as belonging to an indefinite number of taxa of con- 
secutively subordinate rank, among which the rank of species (species) is basic.” 


Infraspecific taxa in Styrax platanifolius 

Evidence from morphology and isozyme analysis indicates that only a single species 
should be recognized among the populations of Styrax in west Texas and adjacent 
Mexico—S. platanifolius,a morphologically and graphically distinct taxon, apparently 
most closely related to the California endemic S. redivivus (Torr) Wheeler (Fritsch 1997). 
While variation in pubescence quantity in other New World Styrax species is essentially 
random, Fritsch (1997, p.741) found that “trichome morphology or abundance within S. 
platanifolius is distinctly regional and facilitates the delimitation of nearly or completely 
allopatric taxa. Therefore, | have recognized five subspecies within S. platanifolius based 
on minor but distinctive differences. Trichome characters are those most reliable for the 
delimitation of these subspecies; characters of secondary importance include leaf form, 
calyx gland density, and surface features of the stem," characters considered by Fritsch 
“not taxonomically reliable” or “taxonomically inconsequential” within many other spe- 
cies of the genus. 

Three of the Styrax platanifolius subspecies (platanifolius, stellatus, texanus) form a 
relatively compact geographic cluster on the Edwards Plateau of Texas; the other two 
(youngiae and mollis) are longitudinally aligned in sierran localities from trans-Pecos Texas 
into northeastern Mexico. Fritsch did not mention any aspect of intergradation among 
these taxa—while it apparently is true that the distinctions are fairly discrete, the popu- 
lations are rare, composed of very few individuals, and each population can be seen as 
completely isolated within the rugged terrain. Fritsch’s comment (p. 743) that “subspe- 
cies stellatus resembles subspecies mollis more closely than does subspecies youngiae” 
might suggest that [subsp] ste/latus and mollis could be considered together as a larger 
infraspecific unit, but the geographic disjunction between these two would render this 
a peculiar concept. 

In sum, the morphological and evolutionary status of infraspecific taxa of Styrax 
platanifolius (sensu Fritsch 1997) corresponds to what we conceive of here as varieties. 
Varietal rank is generally used as the first infraspecific taxonomic category throughout 


261 


the Texas flora, especially in the developing “Atlas of the Flora of Texas” (Turner in prep.). 
In this context, the following varietal combinations in S. platanifolius (a, d, e) are pro- 
posed in order to provide for their use in matters systematic. 


a. Styrax platanifolius var. mollis (PW. Fritsch) B.L. Turner,comb.et stat. nov.B St 
platanifolius subsp. mollis PW. Fritsch, Ann. Missouri Bot. Gard. 84:742. 1997, 


b. Styrax platanifolius Engelm.ex Torrey var. platanifolius. Automatically established with 
the publication of Cory’s var. stellatus, as listed below. 
Styrax platanifolius subsp. platanifolius. Automatically established by Fritch’s subspe- 
cies, as listed below. 


c. Styrax platanifolius var. stellatus Cory, Madrono 7:111.1943. 
Styrax platanifolius subsp. stellatus (Cory ) PW. Fritsch, Ann. Missouri Bot. Gard. 84:743. 


d. Styrax platanifolius var. texanus (Cory) B.L. Turner, comb. et stat. nov. Basionym: Styrax 
texanus Cory, Madrono 7:112. 1943. 


Styrax platanifolius subsp. texanus (Cory) P.W. Fritsch, Ann. Missouri Bot. Gard. 84:744. 


e. Styrax platanifolius var. dae av B.L. Turner, comb. et stat. Nov. Basionym: Styrax 
youngiae Cory, Madrono 7:113 
Styrax platanifolius subsp. youngiae (Cory) PW. Fritsch, Ann. Missouri Bot. Gard. 84:744. 
1997 


ACKNOWLEDGMENTS 


We are grateful to Peter Fritsch for comments on an early draft of the manuscript, to 
John Strother for perspectives on terminology and parsimony, and to Rogers McVaugh 
for a discussion of the issues involved. 


REFERENCES 


BrickeLL, C. and P. TREHANE. 1997.The RHS Advisory Panel on nomenclature and taxonomy. 
New Plantsman 4:115-119.Also <http://www.rhs.org.uk/science/mn_APONAT 1.asp>; 
accessed October 2000. 

Cronauist, A. 1988. The evolution and classification of flowering plants. Ed. 2. New York 
Botanical Garden, New York. 

Fritsch, PW. 1997. A revision of Styrax (Styracaceae) for western Texas, Mexico, and 
Mesoamerica. Ann. Missouri Bot. Gard. 84:705-761. 

Greuter, W., FR. Barrie, H.M. Burbet, W.G. CHALONER, V. DEMOULIN, D.L. HAwksworTH, P.M. JORGENSEN, 
D.H. Nicotson, PC. Sinva, P TREHANE, and J.McNett (eds.). 2000. International code of botani- 
cal nomenclature (Saint Louis Code). Koeltz Scientific Books, Konigstein, Germany. 


262 SIDA 19(2) 


Hamilton, C.W. and S.H. ReicHarb. 1992. Current practice in the use of subspecies, variety, 
and forma in the classification of wild plants. Taxon 41:485-498. 

Hotmoren, N.H. 1994. Redefinition of Dodecatheon dentatum (Primulaceae) and rationale 
for use of varietal rank. Brittonia 46:87-94. 
Hutten, E. 1967. Comments on the flora of Alaska and Yukon. Ark. Bot., n.s. 7:1-147. 
INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1999. International code of zoological 
nomenclature (ed. 4). International Trust for Zoological Nomenclature, London, U.K. 
Kapania, Z.J. 1963.Varieties and subspecies: A suggestion towards greater uniformity. Taxon 
12:257-258. 

Raven, PH. 1974. Proposal for the simplification of infraspecific terminology. Taxon 23: 
828-831. 

Stuessy, T.F. 1990. Plant taxonomy: The systematic evaluation of comparative data. Colum- 
bia Univ. Press, NY. 

THorne, R.F. 1978. New subspecific combinations for southern California plants. Aliso 
9:189-196, 

TREHANE, P., C.D. BrickeLt, B.R. Baum, W.L.A. HeTterscueip, A.C. Lestir, J. MCNEILL, S.A. SPONGBERG, and 
F.VrucTMan, eds. 1995. The international code of nomenclature for cultivated plants (ed. 
6). Quarterjack Publishing, Wimborne, UK. 


NEW SUBTRIBES FOR NORTH AMERICAN 
ASTEREAE (ASTERACEAE) 


A L.Nesom 


aroalina | 


Nor AI olina Botar nical WUT Ciel 
Coker Hall : 3280, University ofNort Carolina 
Chapel Hill, NC 27599 


ABSTRACT 


Recent molecular evidence indicates that most genera of North American Astereae comprise a 
mienopmyletic eemelage Within this North American clade are three Se generic groups 
reviously hypothesized to be most closely related to subtribes of the Southern Hemisphere—the 
Monoptilon group, the Pentachaeta group, and the es ia ele xaionaly eee 
and molecular evidence indicates that the gener a,Chl ely 
related amend) Meiielves ali consume: a eonetent gieup: ve four peu American ae 

Mae naetopappa 


Monoptilon), Pentachaetinae subtr. nov. (Pentachaeta Rigiopappus Tracyina), A 
nov. (Astranthium, Dichaetophora, Geissolepis, Townsendia), and Boltoniinae (Batopilasia, Boltonia, 
Chloracantha). Homochrominae is placed synonymy under subtribe Bellieae. 


—m 


RESUMEN 


Recientes evidencias moleculares muestran que la mayoria de los géneros norteamericanos de 
Astereae comprenden un conjunto monofilético. Dentro de este clado norteamericano hay tres 
qrapoe Genetics Soule los que se piper previamente que fuesen los mas relacionados con 
| grup noptilon,el grupo eee y el grupo Townsendia. 
Adicionalmente, la evidencia morfolégica y molecular indican que los Ces ene 

Chloracantha, y Batopilasia estan muy relacionados entre ellos y caen 
paw cuatro Gieeoo sn norteamericano se reconocen aqui con nombres formales: en el rango: 
v.(Chaetopappa Monoptilon) ), Pentachaetinae subtr. nov. (Pentachaeta, 

Astranth Dichaetophora, Geissolepis, Townsendia) 


nee Taine) 
y Boltoniinae (Batopilasia, Boltonia, Chloracantha). Se lectotipifica Eisenecnrominde colocandola 
en la sinominia de la subtribu Bellieae. 


A primarily morphological overview of the tribe Astereae (Nesom 1994a) recognized 14 
subtribes, four of which were hypothesized to be primarily North American 
(Machaerantherinae, Chrysopsidinae, Solidagininae, and Symphyotrichinae). Three dis- 
tinctive North American generic groups were hypothesized to be most closely related 
to subtribes of the Southern Hemisphere—the Monoptilon group (subtribe Feliciinae = 
Bellieae, primarily Africa and western North America), the Pentachaeta group (subtribe 
Bellieae), and the Townsendia group (subtribe Brachyscominae, primarily Australia and 
North America). The genus Boltonia was hypothesized to be related to genera of subtribe 
Asterinae, primarily an Eurasian group (Nesom 1994a, 1994b). 

Noyes and Rieseberg (1999) used nucleotide sequence data from nuclear riboso- 
mal DNA representing a broad range of Astereaean genera to test various hypotheses of 


SIDA 19(2): 263 — 268. 2000 


264 SIDA 19(2) 


relationship and classification in the tribe. Their results showed that all genera examined 
of North American Astereae comprise a single, strongly supported clade—thus mor- 
phological parallelism rather than homology underlies hypotheses suggesting that dis- 
persal from the Southern Hemisphere and Asia accounts for the origin of the Monoptilon, 
Pentachaeta,and Townsendia groups and Boltonia. 

Notwithstanding these unexpected patterns of relationship, the infra-tribal generic 
groups under consideration have consistently been recognized as coherent (see cave- 
ats below regarding Aphanostephus, Geissolepis, and Boltonia). The morphological dis- 
tinctiveness of these groups is equal to others already recognized at subtribal rank, and 
with recognition of their origin from within the broad North American clade, they are 
provided here with formal names at the same rank. 


Chaetopappinae Nesom, subtr. nov. Tyre cenus: Chaetopappa DC. 


Herbae annuae vel perennes vitae brevis radice palari. Capitula plerumque solitaria; phyllaria 
marginibus late hyalinis. Corollae radii albae vel caeruleae, valde circinnatae. Pappus setarum, 
squamarum vel palearum, setarum ac squamarum, vel carens. Numerus basicus chromosomatum, 
x=8 

Small, annual or short-lived perennial herbs, taprooted, decumbent (Monoptilon, some 
Chaetopappa) to erect. Leaves entire, oblong to oblanceolate-spatulate, alternate. Heads 
mostly solitary; phyllaries flat to convex, with broad, sharply delimited, hyaline margins. 
Ray corollas blue to white, strongly coiling. Disc flowers sometimes with sterile ovaries; 
style branches with obtuse or truncate to triangular collecting appendages. Cypselae 
eglandular or glandular, terete and multinerved (most Chaetopappa) or obovate, flattened, 
and 2-nerved (Monoptilon, some Chaetopappa); pappus of persistent bristles, or scales, 
or pales, or of bristles and scales, commonly in multiples of 5 (in Chaetopappa), or absent. 
Base chromosome number, x = 8.Genera included: Chaetopappa DC., Monoptilon Torr. & 
A. Gray ex Gray. Distribution primarily in arid habitats in the southwestern and south- 
central USA and northern Mexico. 

This is essentially the “Monoptilon group,” earlier placed in subtribe Feliciinae = 
Bellieae (Nesom 1994a) and as “Incertae sedis” (Nesom 2000). The two genera of 
Chaetopappinae form a monophyletic group sister to Futhamia Nutt. ex Cass.in the Noyes 
and Rieseberg analysis. In the analysis of Lane et al. (1996), the phyletic origin of 
Chaetopappa lies immediately between Astranthium and Townsendia, although it does 
not do so in the original analysis from which the DNA data were drawn (Morgan 1990). 

The southern European genus Bellium L. was earlier included in the Monoptilon 
group (Nesom 1994a), but with the strong indication that Monoptilon and Chaetopappa 
are North American in origin, it is probable that the closest relationships of Bellium are 
with the other Old World genera. Similarities between Bellium and Amellus L.were noted 
by Nesom (1994a). 


Pentachaetinae Nesom, subtr. nov. Type Genus: Pentachaeta Nutt. 


Herbae annuae radice palari. Folia filiformia vel linearia vel anguste oblanceolata. Capitula solitaria 
longipedunculata; phyllaria marginibus hyalinis. Corollae radii flavae vel rubescentes, minus 


NESOM, NEW SUBTRIBES IN ASTEREAE 265 


plerumque albae, Naas pl Miee ORENGIEES collectentes: lineari-lanceolatae Jammorunn sul 


florum disci 
pappus setarum vel squamarum. * Nuraanus basicus chroniocomatunn x= 9, 


Annual herbs, taprooted. | It te, entire, filiform or linear to narrowly oblanceolate. 
Heads solitary and long-pedunculate; phyllaries with hyaline margins. Ray corollas yel- 
low to reddish, less commonly white, strongly coiling (lamina absent in some Pentachaeta). 
Disc flowers: style branches with linear-lanceolate collecting appendages. Cypselae ter- 
ete to slightly compressed, narrowly oblong to oblanceolate in outline, beaked in Tracyina, 
eglandular; pappus 1-seriate (1-2 seriate in Tracyina), of persistent bristles frequently in 
multiples of 5s, sometimes flared at the base and partially connate, or sometimes com- 
pletely lacking, or of long scales (Rigiopappus). Base chromosome number, x = 9.Genera 
included: Pentachaeta Nutt., Rigiopappus A.Gray, Tracyina S.F.Blake. Distribution primarily 
in grassland habitats of California, USA, with several taxa of Pentachaeta reaching Baja 
California, Mexico; Rigiopappus also occurs into Oregon, Washington, Idaho, and Nevada. 

This is the “Pentachaeta group,"earlier placed in subtribe Feliciinae = Bellieae (Nesom 
1994a) and as “Incertae sedis” (Nesom 2000). The close resemblance and relationship 
among these three genera have been noted by Blake (1937), Ornduff and Bohm (1975), 
and Robinson and Brettell (1973); they were placed as a coherent unit within the 
“Chaetopappa group" by Bremer (1994).The Pentachaetinae form a monophyletic group 
sister to Ericameria Nutt.in the Noyes and Rieseberg analysis. 


Astranthiinae Nesom, subtr. nov. Tyee cenus: Astranthium Nutt. 


Herbae annuae biennes vel perennes plerumque radice palari. Capitula solitaria, plerumque 
longipedunculata; phyllaria marginibus late hyalinis; receptacula convexa vel conica. Corollae radii 
plerumque albae vel caeruleae, midfascia abaxiali lavandula, non reflexae aut circinnatae. Corollae 
discii tubo OIevt ey pseiae oblanceolatae sa ppovarer: Complahates, a 3)- pervane, aves vel 
papillata 


setarum et squamarum. Numerus basicus chromosomatum, X= 9 et xX = 3,4,5. 


Annual, biennial, or perennial herbs, taprooted, often with a branching caudex, rarely 
fibrous-rooted, mostly strigose with short, white hairs. Leaves alternate, spatulate to linear, 
entire or few-toothed. Heads solitary, mostly long-pedunculate, rarely sessile; phyllaries 
with broad, hyaline margins; receptacles convex to conical. Ray corollas white to bluish or 
pinkish above (rarely yellow in Townsendia), usually with a lavender to blue or pink abaxial 
midstripe, not reflexing or coiling. Disc corollas short-tubed; style branches with triangular- 
lanceolate collecting appendages. Cypselae oblanceolate to obovate, flattened, 2(—3)- 
ribbed, the surfaces smooth or papillate, glabrate or usually pubescent with duplex hairs 
with glochidiate, bifurcate, or entire apices (achenes winged and fringed-ciliate in 
Dichaetophora); pappus 1-seriate, of barbellate bristles or a low crown of setae/bristles 
and scales (2-awned in Dichaetophora).Base chromosome number, x = 9 (or x = 3,4,and 5 
in Astranthium;x = 3 in Dichaetophora).Genera included:Astranthium Nutt, Dichaetophora 
A. Gray, Geissolepis B.L. Rob., Townsendia Hook. Distribution mostly in western North 
America north of Mexico (Townsendia), Mexico and south-central USA (A hium), Texas 
and adjacent Mexico (Dichaetophora), and east-central Mexico (Geissolepis). 


266 SIDA 19(2) 


This is the “Townsendia group,” earlier placed in subtribe Brachyscominae (Nesom 
1994a) and as"Incertae sedis” (Nesom 2000).Aphanostephus DC. also was earlier included 
in the Townsendia group (by Nesom and various others), but molecular data from sev- 
eral sources (Morgan 1990; Lane et al. 1996; Noyes and Rieseberg 1999; Noyes 2000) con- 
sistently indicate that Aphanostephus arose from within the Conyzinae. Astranthium, 
Dichaetophora,and Townsendia comprise a monophyletic group essentially sister to the 
Conyzinae and Chrysopsidinae in the Noyes and Rieseberg analysis. Geissolepis is phyl- 
etically interposed between the Townsendia group and these two related subtribes, but 
a set of morphological features places it closer to the Townsendia group than the 
Conyzinae or Chrysopsidinae, and the genus can reasonably be included in the 
Astranthiinae. Alternately, it presumably would be treated as a monophyletic subtribe. 

Geissolepis is similar to the Astranthiinae “core” genera in its solitary heads, white, 
straight ray corollas, conical receptacles, and glochidiate cypselar vestiture. It is distinct 
within the subtribe in its combination of a prostrate habit with creeping, fibrous-rooted, 
lignescent stolons, succulent leaves, resin canals on the phyllaries, cypselae, and disc 
corollas, paleate receptacles, ray corollas without an abaxial midstripe, gradually ampliate 
disc corollas, subterete cypselae with 8 resinous ribs, and pappus of short scales with 
uncinate-ciliate margins. The chromosome number has been reported as 2n = 16 (Ralston 
et al. 1989) and 2n = 18 (Lane and Li 1993). 


Boltoniinae Nesom, subtr. nov. Tyee Genus: Boltonia L'Hérit. 


Herbae vel subfrutices perennes rhizomatosae caulibus ac foliis persistente viridi-glabratis. Folia 
m omnino caulina. Capitula solitaria vel laxe aogieceta) oe herbacea, apicem 
+ Pee | Lae 
IAL Ia ka \ Sa -res Osis SG vel leniter 
circinnatae. Appendices collectentes eas ramorum styli florum disci. Numerus basicus 
chromosomatum, x 


Perennial, herbs or subshrubs (Chloracantha), rhizomatous, with persistently green- 
glabrate stems and leaves, thorny in Chloracantha. Leaves essentially all cauline, entire or 
few-toothed. Heads solitary or very loosely corymboid to paniculate; phyllaries primarily 
herbaceous, apically rounded to obtuse, with three orange-resinous nerves. Ray corollas 
white to slightly bluish, coiling. Disc corollas orange-veined; style branches with deltate 
collecting appendages. Cypselae terete and multinerved or flattened, 2-nerved, and 
winged (Boltonia).Base chromosome number, x = 9.Genera included: Batopilasia Nesom 
& Noyes, Boltonia L'Hérit., Chloracantha Nesom, Suh, Morgan, Sundberg, & Simpson. Dis- 
tribution in northwestern Mexico (Batopilasia), eastern USA (Boltonia), and Mexico and 
the southwestern USA to Louisiana (Chloracantha). 

Batopilasia, Boltonia,and Chloracantha apparently are closely related among them- 
selves (summary of ideas and evidence in Nesom & Noyes 2000). Batopilasia and Boltonia 
are sister genera in the Noyes and Rieseberg analysis, the pair in a sister relationship to 
the Symphyotrichinae and Machaerantherinae. Chloracantha was not included in this 
molecular analysis but is morphologically similar to Batopilasia,as observed in the origi- 
nal description of the species (Sundberg & Nesom 1990). 


NESOM, NEW SUBTRIBES IN ASTEREAE 267 


Boltonia is set apart from the other two genera by its conical or convex receptacles, 
short-tubed disc corollas, and flattened, 2-nerved, orange-veined, often winged cypselae 
with an abbreviated pappus. These specializations prompted the observation that 
"Boltonia is morphologically isolated in the New World” (Nesom 1994b, p. 163), but mo- 
lecular evidence has found its close relatives. It is related neither to Old World Asterinae 
(Nesom 1994b) nor to genera of the Townsendia group (Bremer 1994). 

With the formal recognition of the four subtribes above, six North American genera 
of Astereae remain without a clear hypothesis of subtribal affinity. These are among the 
“orimitive’ Asters” (Nesom 2000), placed by Nesom (1994a, 1994b) mostly in subtribes 
Asterinae or Symphyotrichinae: Doellingeria Nees, Eucephalus Nutt., lonactis Greene, 
Oclemena Greene, Oreostemma Greene, and Tonestus A. Nelson. 


Note on Homochrominae.—My interpretation (Nesom 1994a) of subtribe 
Homochrominae of Bentham & Hooker as invalid was incorrect. This name was validly 
peer was provided with a description and it can be assumed to have been 
based on a legitimate genu (Homochroma DC.) included by Bentham and Hooker 
among those genera ey placed in the subtribe. Homochrominae presumably formed 
the nomenclaturallysister”taxon to the illegitimate Heterochrominae Bentham & Hooker 
(this name not based on an included genus), but a reasonable lectotypification for 
Homochrominae already has been effected by Solbrig (1963), who specified the type. 


Homochrominae Bentham & Hooker, Gen.PI.2:174.1873.Lectorye: (Solbrig 1963): Homochroma 
DC. (= Zyrphelis Cass.). 

With this typification, Homochrominae becomes a synonym of Bellieae DC. ex Godr. (in 

Gren. & Godr,, Fl. France 2:83, 104. 1850; type, Bellium L.) along with Feliciinae Nesom 

(Phytologia 76:205. 1994; type, Felicia Cass.). 


ACKNOWLEDGMENTS 
| appreciate the helpful comments of Ted Barkley and K.N. Gandhi. 


REFERENCES 


Bake, S.F. 1937. Tracyina, a new genus of Asteraceae from northern California. Madrono 
4:73-77. 

Bremer, K. 1994. Asteraceae: cladistics and classification. Timber Press, Portland, Oregon. 

Lane, M.A.and J.L.1993.Chromosome number reports in Compositae with emphasis on 
tribe Astereae of the southwestern United States and Mexico. Sida 15:539-546. 

Lane, M.A., D.R. Morcan, Y. SUH, B.B. Simpson, and R.K. Jansen. 1996. Relationships of North 
American genera of Astereae, based on chloroplast DNA restriction site data. In DJ.N. 
Hind and H.J. Beentje (eds.). Compositae: Systematics. Vol. 1, pp. 49-77. Proc. Interntl. 
Compositae Conf. Kew, 1994. 

MorcAn, D.R.1990.A systematic study of Machaeranthera (Asteraceae) and related groups 


268 SIDA 19(2) 


using restriction site analysis of chloroplast DNA and a taxonomic revision of 
Machaeranthera section Psilactis. Ph.D. dissertation, Univ. of Texas, Austin. 

Nesom, G.L. 1994a. Subtribal classification of the Astereae (Asteraceae). Phytologia 76: 
193-274. 

Nesom, G.L. 1994b, Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), 
emphasizing the New World species. Phytologia 77:141-297. 

Nesom, G.L.2000.Generic conspectus of the tribe Astereae (Asteraceae) in North America, 
Central America, the Antilles, and Hawaii. Sida, Bot. Misc. 20:1-100. 

Nesom, G.L. and R.D. Noyes. 2000. Batopilasia (Asteraceae: Astereae), a new genus from 
Chihuahua, Mexico. Sida 19:79-84. 

Noyes, R.D. 2000. Biogeographical and evolutionary insights on Erigeron and allies 
(Asteraceae) from ITS sequence data. PI. Syst. Evol. 220:93-114. 

Noyes, R.D. and L.H. Rieseserc. 1999. ITS sequence data support a single origin of North 
American Astereae (Asteraceae) and reflect deep geographic divisions in Aster s.l. Amer. 
J. Bot. 86:398-41 2. 

Ornourr, R.and B.A. Boum. 1975. Relationships of Tracyina and Rigiopappus. Madrono 23: 
53-55. 

Ratston, B.,G.L. Nesom, and B.L. Turner. 1989. Chromosome numbers in Mexican Asteraceae 
with special reference to the tribe Tageteae. Sida 13:359-368. 

Rosinson, H.and R.D. Brettet. 1973. Tribal revisions in the Asteraceae. V. The relationship of 
Rigiopappus. Phytologia 26:69—70. 

Sovsric, O.T. 1963. Subfamilial nomenclature of Compositae. Taxon 12:229-235. 

SunpeerG, S.D. and G.L. Nesom. 1990. A new species of Erigeron (Asteraceae: Astereae) from 
Chihuahua, Mexico. Phytologia 69:278-281. 


STYLOGYNE AGUARUNANA (MYRSINACEAE) 
A NEW SPECIES FROM AMAZONAS, PERU 


John J. Pipoly Ill 


Fairchild Tropical Garden Research Center 
11935 o ld Cutler ned Miami, fl, 331 oe saad, U.S.A. 


OF ad td 


Jon M. Ricketson 
Missouri Botanical Garden 

PO. Box 299, St. Louis MO 63166-0299, U.S.A. 
jon.ricketson@mobot.org 


ABSTRACT 


Preparation of our treatment of the family Myrsinaceae for the florula of the “Rio Cenepa’” region 
resulted in the discovery of a heretofore undescribed species in the genus Stylogyne. Stylogyne 
aguarunana Pipoly & Ricketson is described, illustrated, mapped and its phylogenetic relations are 
discussed 


RESUMEN 


Estudios para preparar un tratamiento taxondmico de la Myrsinaceae para la fldrula de la regidn 
“Rio Genepa resultaron en el oe ul ae de una nueva especie perteneciente al género 
Stylogyne. S g Pipoly & Ri describe, se ilustre, se mapea y se discute su 
parentezco. 


INTRODUCTION 


The neotropical genus Stylogyne A.DC. comprises approximately 60-70 species, a num- 
ber of which remain undescribed, owing to lack of adequate material. Understanding 
the systematic biology of this genus has long been problematic because of its sexual 
liability. Androdioecious, bisexual, polygamo-dioecious and dioecious species of Stylogyne 
have been documented (Pipoly 1989, 1991), as well as the consequent morphological 
variation due to sex expression. Since no comprehensive study of the genus Stylogyne 
has occurred since C. Mez’s treatment in Engler’s Das Pflanzenreich (1902), the genus is 
in need of additional study. Our current studies continue in the genus Stylogyne (Pipoly 
& Ricketson 1999; Ricketson & Pipoly 1997), as well as the entire family for our treatment 
of the Myrsinaceae for Flora Neotropica. 

During preparation of a manuscript for the Florula of the Rio Cenepa Drainage Ba- 
sin Project of the Missouri Botanical Garden, a new species was found and is described 
herewith. 


Stylogyne aguarunana Pipoly & Ricketson, sp.nov. (Fig. 1). Tye. PERU. Amazonas: Prov. Bagua; 
Distrito Imaza, NW Region of Rio Maranon; Comunidad Yamayakat, 04° 55' S,078° 19' W, 320 


SIDA 19(2): 269 — 273. 2000 


270 SIDA 19(2) 


m,5 Aug 1994 (pist.fl.), N. Jaramillo, A. Pera, R.Apanu & S. Katip 296 (HoLotype: MO; isotypes: CPUN 
nv., F, FTG). 
Quoad inflorescentiam racemosam bractea floricina numerosa S. brancteolatam valde arcte affinis 

b ea habito arbreo (non fruteo), ramulis crassis (non tenuis), laminis subsessilibus (non 

longipetiolatis), 32-39 (nec 9.5-18) cm longisque praeclare distat. 
Tree or small tree 4-8 m tall, to 5.7 cm in diam. Branchlets ca. 10 mm in diam., terete, the 
bark yellowish-brown, longitudinally ridged, glabrous, hollow, der isely lenticellate. Leaves 
pseudoverticillate; blades chartaceous, oblong to oblanceolate, 32-39 cm long, 5.5-13.4 
cm wide, apically acute to obtuse, basally tapering gradually to petiole base, the midrib 
channel prominently raised above, decurrent on the petiole/stem junction, the midrib 
prominently raised below, the secondary veins numerous, brochidodromous, 
prominulous above and below, smooth above, densely and conspicuously red punctate 
below, the hydropotes scattered, bright orange below, the margin flat, entire; leaf base 
and petiole, when distinguishable, deeply canaliculate, obsolete to 10 mm long,the petiole 
margin often abruptly tapered at petiole apex to almost appear auriculate, deeply 
canaliculate and marginate, glabrous. Staminate inflorescence and flowers unknown. Pis- 
tillate inflorescence lateral, a condensed raceme, 3-10 mm long; floral bract girdling pe- 
duncle, chartaceous, elliptic, 2.7-3 mm long, 1.3-1.5 mm wide, apically acute, densely 
and prominently orange punctate and punctate-lineate, the margin irregular, somewhat 
erose apically otherwise entire; pedicel cylindrical, 2.5-3 mm long. Pistillate flower 5- 
merous, white; calyx carnose, membranaceous, 2.5-2.7 mm long, the tube ca. 0.2 mm 
long, the lobes nearly free, ovate, 2.3-2.5 mm long, 1.5-1.7 mm wide, apically obtuse, 
with one or two orange punctations medially, glabrous, the margin hyaline, entire; co- 
rolla membranaceous, 3.8-4.2 mm long, the tube 2-2.2 mm long, the lobes connate 
basally, ovate to lanceolate, 1.8-2.2 mm long, 1-1.2 mm wide near the base, apically acute, 
conspicuously orange punctate and punctate-lineate, glabrous, the margins entire, hya- 
line;stamens 3.7-4.3 mm long, the filaments 2.8-3.2 mm long, filamentous, free, epunctate, 
glabrous, the anthers free, narrowly ovate to lanceolate, 1.2-1.3 mm long, 0.5-0.6 mm 
wide at the base, apically emarginate, basally cordate, longitudinally dehiscent by slits, 
the connective conspicuously punctate; pistil obturbinate, 3-3.3 mm long, 1.3-1.5 mm 
in diam, glabrous, the ovary 1.5-1.7 mm long, the style 1.3-1.5 mm long, the stigma 
punctiform, the placenta cotyliform, with 4 open chambers above, the ovules 4,exposed. 
Fruit globose, 5—6.2 mm long, 5-6.2 mm diam., apically truncate, red at maturity, densely 
and prominently pellucid lineate-punctate, the exocarp thin. 

Distribution.—Stylogyne aquarunana is known only from Imaza District, Bagua Prov- 
ince, Amazonas, Peru, in the Rio Marahon Drainage Basin around the Comunidad 
Yamayakat (Fig. 2), from 300-480 m. 

Ecology and conservation status.—Stylogyne aguarunana is know from only four 
collections and is thus considered rare. It occurs in primary transitional forests at the 
junction of the lowland with the premontane forests on sandstone-derived soils. 

Etymology.—We dedicate this species to the Aguaruna people who inhabit the area. 


PIPOLY AND RICKETSON, A NEW SPECIES OF STYLOGYNE FROM PERU 271 


fl D. Detail of pistillate fl E. Fruit and fruiting infl A-D drawn from holotype, N. Jaramillo 
et al. 296 (MO). E drawn from C. Diaz et al. 7895 (MO). 


Fig. 1. Styl Pipoly & Ricketson. A. Fl ing b h.B. Detail of ak leaf surf. C. Detail of pistil- 


Pararypes. PERU. Amazonas: Prov. Bagua, Distrito Imaza, Comunidad Aguaruna Yamayakat, camino 
hacia Temashnum, bordes de la quebrada, 300-480 m, 17 Ago 1996 (fr), C. Diaz et al. 7895 (CPUN nwv., 
FTG, MO); Prov. Bagua, Distrito Imaza, Comunidad Aguaruna de Putuim, arriba Rio Shimutaz, 04° 55' 
S, 078° 19' W, 480 m, 20 Jun 1996 (young bud), £. Rodriguez et al. 1167 (CPUN nw., FTG, MO); Prov. 
Bagua, Distrito Imaza, Yamayakat, 05° 03' 20"S, 078° 20' 23"W, 380 m,6 Nov 1996 (ster.), R. Vasquez et 
al. 21594 (CPUN nvv., MQ). 


272 


SIDA 19(2) 


oe 
7 
- 
- 
oe 


te 


La 


~~ Sl 
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_ 
7 
P 


Rio Cenepa 
= 


ECUADOR 


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. 

. 


e : 
Comunidad ‘ 
' / Y amayakat : 
4 H ' 
a / : 
PERU ‘ 4 ry 
- ‘ ( 
‘ ' 
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‘ : A LoRETO 
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‘ ‘ 
ry ‘ \ 
‘ ‘ \ 
‘ ‘ AMAZONAS his 
< i : eS 
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Pd @ Bagua 4 ae 
Fig ? ee eee € Cadeci ne ly 0° Di-Lne LV teh ~ earn 5 saa 
the Rio Marafién, in the D t tofA , Peru 


The long leaf blades, that are nearly sessile and pseudoverticillate, clearly distinguish 
Stylogyne aguarunana from all other members of the genus. The multibracteate inflores- 
cence rachis is similar to those of Stylogyne bracteolata, but can be easily distinguished 
from that species by its smaller or obsolete petioles, and 5-merous flowers. Stylogyne 
aguarunana belongs to a group of taxa with 5-merous flowers and anthers and 
antherodes on long spindly filaments, generally 3 or more times longer than the anthers. 


ACKNOWLEDGMENTS 


We thank the Missouri Botanical Garden, the Flora Mesoamericana Project, and the 
Fairchild Tropical Garden, for funding that allowed J.Ricketson (MO) to visit J. Pipoly (FTG) 
and vice versa.We are also grateful to those who have been so instrumental in assisting 


PIPOLY AND RICKETSON, A NEW SPECIES OF STYLOGYNE FROM PERU 273 


us in our work, including Gerrit and Jeany Davidse, Linda Oestry, Mary Bard, and Catherine 
Mayo (MOQ), Barney Lipscomb, Debra Trock and Jim Rivers (BRIT). Illustrations were pre- 
pared by the junior author. Reviews of the manuscript by Gerrit Davidse and Roy Gereau, 
and meticulous copy editing by Barney Lipscomb, improved the presentation of the 
paper. 


REFERENCES 


Mez, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich IV. 236(Heft 9):1-437. 

Pipoty, J.J. 1989. Notas one 2 gener ylegyiae A. DC. (Myrsinaceae). Ernstia 53:1—9. 

Pipoty,J.J.1991. Styl (Myrsinaceae):a new androdioecious species from 
Amazonia. on 1:202- 203, 

Pipovy, J.J.and J. Ricketson. 1999. Novelties in the Myrsinaceae from the Venezuelan Guayana. 
Sida 18:1167-1174. 

Ricketson, J. and J. Pipoty. 1997. Nomenclatural notes and a synopsis of Mesoamerican 
Stylogyne (Myrsinaceae). Sida 17:591-597. 


— 


274 SIDA 19(2) 


BOOK REVIEW 


Karsten H.K. Woprich. 1997.Growing South African Indigenous Orchids. (/SBN 90-5410- 
650-6, hbk.). Rotterdam (in USA: A.A. Balkema Publishers, Old Post Road, 
Brookfield, VT 05036-9704, U.S.A (Fax 802-276-3837, e-mail info@ashgate.com, 
http://www.balkema.nl) $85.00 hbk., 253 pp., 94 color and 109 b&w photos and 
drawings 


This beautifully illustrated volume is a self-contained monograph on how to grow South African 
orchids, from summarizing the habitats in which the orchids are found to providing plans for set- 
ting up a tissue culture lab at home. Although, the focus is on species native South Africa, much of 
the general discussion can be applied to the culture of any orchids. Early in the book, the discus- 
sion moves from general aspects of growing orchids to the various habitats and climates in South 
Africa. Next is a general discussion of design of culture space around one’s home, orchid physiol- 
ogy and nutrient requirements, and orchid pests and diseases. The next chapter looks at the three 
major growth forms of orchids and the horticultural requirements common to each. Within the 
treatment of the respective growth habits, each genus and species is broken out for specific in- 
struction on natural occurrence and horticultural needs. Most species are illustrated with a color 
photograph of its inflorescence or individual flower. There are sections on orchid breeding and 
propagation using culture media both with and without fungal symbionts. Step-by-step photos 
and recipes guide the novice through sterile culture methods using only kitchen equipment and a 
pH meter. An appendix even includes designs for building one’s own sterile seeding box and a 
laminar- peel pees Tie etek does ne longs conservation issues and includes copies of conser- 

tion legisl d well-designed that | found only one negative point: 
the inks contain a & Welatile comipeund { that has the odor of oven lie Guava This book is a must- 

or orchid {would make a 


library.—Roger W. Sanders, Associ ate Collections Manager, Botanical Research Institute of Texas. 


10! ticulturist ‘5 


SIDA 19(2): 274, 2000 


DISCOVERY OF ARDISIA SUBGENUS ACRARDISIA 
(MYRSINACEAE) IN MESOAMERICA: ANOTHER 
BOREOTROPICAL ELEMENT? 


John J. Pipoly Ill 


faiiena Oe aes Garden 
11935 Old Ses nodg, Co Cae! (Mia n Lae 331 7 4299, U.S.A. 


JPOP 


Jon M.Ricketson 
Missouri Botanical Garden 

PO. Box 299, St. Louis MO 63166-0299, U.S.A. 
jon.ricketson@mobot.org 


ABSTRACT 


Ardisia rarescens, native to the border area of SE Chiapas, Mexico and SW Guatemala, was studied in 
preparation for a Mieaenens of Ws for the Flora oe plolect Its Cerone 
he 


imbricate petals,e tremely Pies! Oe Le Hlo ft ame i) wide longitu 
dinal slit from each theca meet apical to Sain a subeontnuous 0 opening at panthesie short and 
road| nical flower buds, and relatively rather thick plac enta, precluded 


its placement in any known neotropical subgenus of Ardisia. Guienst the aforementioned fea- 
tures comprise the diagnostic character states unique to the Indo-Malesian Ardisia subgenera 

Acrardisia and Stylardisia. Because subgenus Stylardisia is defined by its protogynous flowers (the 
style piercing the bud apex long before the bud opens), Ardisia rarescens can be excluded from 
that group and therefore placed in subgenus Acrardisia Mez.We suggest that Ardisia rarescens is a 
member of Ardisia subgenus Acrardisia and that this represents another species whose distribu- 
tion may be attributable to the logical extension of the Boreotropics Hypothesis, previously ex- 
plained in our work on the discovery of the genus Hymenandra in Mesoamerica. Ardisia rarescens is 
described, newly illustrated, its distributions, phylogenetic relationships, ecology and conservation 
status, are discusse 


RESUMEN 


Ardisia rarescens fue estudiada para el tratamiento taxondmico de la familia Myrsinaceae para la 
Hele Meson Mean Sus ae Sex Opa vente MON ages) porcione® apical libres ce ee 


nchas y continilds en el snes botones corto- y anchamente cénicos en perfil, y Ovulos 
er eae nae insertados en una pl nta gru para indicar que no 
fue posible ubicar ésta especie dentro de un suagene previamente canecice en he useieletces 


Sin embargo, los caracteres susodict 


Acrardisia, y Stylardisia,am ientes d la region Indo-Malesia Debido e! hecho de ne se 
define el subgénero cas ee fl alidad, 
ubicamos la especie HeNHO oc subgénero Acrardisia Surgerimos - Ardisia rarescens es otro 
elemento, tant | ya distribuci6n se atribuya 


fiiaok ee] 


ala extension logica de lat ede sails se ee y se discute su distribuci6n, 
parentezco, ecologia y eee en cuanto a la conservacion se refier 


SIDA 19(2): 275 — 283. 2000 


276 SIDA 19(2) 


INTRODUCTION 


The pantropical genus Ardisia Sw. is by far the largest in the family Myrsinaceae, contain- 
ing perhaps as many as 500 species (Chen & Pipoly 1996). Its circumscription has been 
problematic owing to a lack of comprehensive treatment since that of Mez (1902) in 
Engler’s Pflanzenreich, almost a century ago.While reviewing the status of Ardisia rarescens 
Standl., we observed the combination of: dextrorsely imbricate petals, extremely short 
free apical portions of the filaments; anthers whose longitudinal slits from each theca 
meet apically to form a subcontinuous opening at anthesis; short, broadly conical buds, 
and relatively numerous ovules on the rather thick (thicker than long) placenta, all char- 
acteristics of Ardisia subgenus Acrardisia Mez.With subgenus Acrardisia’s distribution from 
Sri Lanka eastward through Malaysia and Indonesia to the Philippines and New Guinea, 
it is strikingly similar to that of Hymenandra (A. DC.) A. DC. ex Spach, a group that we 
suspect is of boreotropical origin (Pipoly and Ricketson 1999).A distribution such as this, 
most closely fits those groups cited as partial evidence to support what Wendt (1993) 
discovered in his study of lowland Mexican wet forests, and what Lavin and Luckow 
(1993) attributed to the Boreotropics Hypothesis, proposed by Wolfe (1975) and Tiffney 
(1985a, b) to explain the distribution of the “boreotropical flora.” The hypothesis pro- 
poses that the biotas of North America and Europe, including tropical North America, 
were once more widespread in the northern hemisphere and transgressed the North 
Atlantic by direct land connections or over limited water gaps until the late Eocene or 
early Oligocene (Lavin & Luckow 1993). A logical extension to Wolfe's and Tiffney’s con- 
cept, presented by Wendt (1993) and Lavin and Luckow (1993) is that the boreotropical 
flora not only existed in the Eocene, but also left a significant number of direct descen- 
dant lines in present lowland tropical floras of northern Latin America. Therefore, we 
would expect that many of the early Tertiary fossil taxa from both North America and 
Europe were most closely related to extant species from tropical southeast Asia, and to 
some extent, Central America and the Greater Antilles (Lavin & Luckow 1993). It is the 
latter notion that is congruent with the Amphipacific distributional pattern like that found 
in our new concept of Ardisia subg. Acrardisia. This extension to the boreotropics hy- 
pothesis was cited by Wendt (1988, 1989, 1993), in discussing the relationships of 
Chiangiodendron (Flacourtiaceae), and by Zona (1990) in discussing the biogeography 
of Sabal (Arecaceae). A similar distribution for the genus Alstonia (Apocynaceae) was 
cited by Gentry (1983), but he did not invoke the hypothesis per se. Conran (1995), in his 
study of the Liliiflorae, found that three taxa defining the Southeast Asian/northern 
Australasian clade (Stemonaceae, Hanguanaceae and Uvulariaceae) were widespread 
northern taxa that have spread southwards. 

The geographic distribution of subgenus Acrardisia is entirely consistent with the 
area cladogram presented by Lavin and Luckow (1993, Fig. 1), where Central and South 
American elements are ultimately derived from among diverse North American lineages, 
these lineages having a sister group relationship to paleotropical groups. While Wolfe's 


277 


(1975) hypothesis could be correct even if no modern descendants of the boreotropical 
flora were found in the Neotropics owing to extinction, the three criteria proposed by 
Lavin and Luckow (1993) to test the hypothesis were: 1) a center of diversity in North 
America (including “tropical North America” as they define it), 2) an early Tertiary fossil 
record in North America, and 3) a pantropical distribution. 

For the first test criterion, “North America” includes both tropical and temperate 
elements, the tropical ones south of the Tropic of Cancer.In the case of subgenus Acrardisia 
it is clear that only one species is presently known, but until we have tested hypotheses 
of phylogenetic relationship among other related Ardisia species, in particular the group 
described by Lundell as /barrea, and containing the rather common species Ardisia 
paschalis Donn. Sm., we cannot be sure. Subgenus Acrardisia itself may be paraphyletic, 
because the only characteristics separating it from the extremely closely related subge- 
nus Stylardisia Mez are that it does not have a stigma that perforates the flower bud and 
is receptive before the flower opens (thus protogynous), and that its ovules are pluri- 
rather than uniseriate. A group even partially defined by the lack of a structure can be 
problematic because it is not clear if protogynous flowers have occurred more than 
once within the genus Ardisia as a whole. 

Unfortunately, no data is available for use in the second test criterion owing to lack 
of fossils known for the group. As for the last criterion, a pantropical distribution, the 
closest related group, Ardisia subgenus Stylardisia, is concentrated in Borneo, but occurs 
from northern India and Bangladesh through Indochina, eastward through Malesia as 
far as Sulawesi, with no neotropical members known at this time. However, the entire 
genus Ardisia is truly pantropical, as is the tribe Ardisieae. Until a phylogenetic analysis is 
complete for the tribe, a rigorous test cannot be performed. Despite the absence of a 
cladogram, the pattern of distribution among the genera of the tribe Ardisieae fit the 
general pattern one would expect if ule coon was poe cones 

In summary,we hypothesize that Ardisia subger rs to fit the overall 
pattern consistent with a boreotropical dctibutlor: We r must snienepa te that until a 
phylogenetic analysis among the subgenera of the genus Ardisia,and more importantly, 
among the genera of Myrsinaceae is complete, there is no reliable way to unequivocally 
determine if the group is boreotropical or Gondwanan. However, at this point in our 
work, we find it useful to point out the strong correlation and call attention to the value 
of examining generic limits on a worldwide basis when preparing treatments for a large 
flora such as Flora Mesoamericana. 

Ardisia rarescens is known only from 27 herbarium specimens, and apparently is 
restricted to the eastern slope of the Sierra Madre de Chiapas, from central-southern- 
most Chiapas, Mexico, along the s| dint thern San Marcos and Quetzaltenango 
Departments, Guatemala. This region, including the famous Volcan Tacana, houses many 
endemics and a myriad of taxa with boreotropical affinities, particularly in the pine-oak 
and Liquidambar forests in the region. 


278 SIDA 19(2) 


NOTES ON DESCRIPTION AND TERMINOLOGY 


Quantitative and qualitative data in the description for floral parts and bracts were taken 
from organs rehydrated from herbarium specimens by boiling in water. Measurements 
from these range from 10% to 15% greater than those measurements taken directly 
from dried material. Data regarding stem diameters, inflorescence rachises, pedicels, leaf 
and fruit shape and size were taken from dried herbarium specimens. 

Morphological terms in this treatment follow Lindley (1848) and Pipoly (1987, 1992) 
for the inflorescence, rachis pedicels and floral parts. Description of leaf morphology 
follows Hickey (1984), trichome description follows Theobald et al. (1984) and basic cell 
and tissue terminology follow Metcalfe (1984). 


TAXONOMIC TREATMENT 
Ardisia Swartz subgenus Acrardisia Mez, Pflanzenr. lV. 236(Heft 9):116. 1902. 


Subshrubs to small trees. Branchlets glandular-papillate, glandular-hirtellous, ferrugine- 
ous tomentose or rarely, glabrous. Leaf blades mostly entire or obscurely crenulate, but 
never regularly serrate. Inflorescence terminal, subtended by a foliaceous bract, the 
branches corymbose to umbellate or eye racemose in fruit. Flowers 5-merous; — 
short and broadly conical;sepals and p ly imbricate;st 
short apical portions of the ene anthers dehiscent by wide, apically confluent lon- 
gitudinal slits; pistils not protogynous, the ovules numerous, pluriseriate 

Distribution —Approximately 35 species, throughout Indo-Malesia, with one disjunct 
at the border of Mexico and Guatemala. 

Ecology.—Premontane to montane humid to wet forest, often on calcareous or sand- 
stone-derived soils, 500-2400 m elevation. 


Ardisia rarescens Stand. (Fig. 1), Publ. Field Columbian Mus., Bot. Ser.4:248.1929.Amatlania 
rarescens (Standl.) Lundell, Wrightia 7:40. 1982. Type. MEXICO. CHiapas:Cerro del Boquerén, 
without elevation, Sep 1913 (fl) CA. Purpus 7032 (HoLotype: F, F neg.n0.68247; IsoTyYPEs: 
BM, GH, MO, NY, US) 


Shrub to small trees 3-12 m tall. Branchlets slender, terete, 2-5 mm in diam, densely glan- 


dular-papillate, glabrescent. ly pseudoverticillate; blades chartaceous, elliptic 
to slightly oblanceolate, 3.5-16.2 cm long, 1.1-5.3 cm wide, apically acuminate, the acu- 
men 0.5-1.4 mm long, basally acute to cuneate, decurrent on the petiole, midrib im- 
pressed above, prominently raised below, the secondary veins 15-27 pairs, slightly raised 
above and below, prominently black punctate and conspicuously punctate-lineate, gla- 
brescent above and below, the margins entire, revolute; petioles slender, canaliculate, 
3.5-10.2 mm long, glabrescent above and below. Inflorescence terminal, erect, bipinnately 
or tripinnately paniculate, 6—22.5 cm long, 5.5-20 cm wide, pyramidal, usually longer 
than the leaves, densely, erect glandular-papillate, the branches terminally congested 
into 7-12-flowered corymbs; peduncles 0.3-2.2 cm long; inflorescence bract unknown; 
inflorescence branch bracts foliaceous, chartaceous, ovate to oblong, 2.3-3.7 cm long, 


279 


Fic. 1. Ardisy, C+ dil A FI . kL hBD 1 fab a nT rd £. C Detail Linfl D Detail of 


flower. E. Detail of stamen f F. Detail of stamen, st in. G. Detail of stamen, 


showing abaxial surface. H. Fruit. A-B drawn from isotype, C. Purpus 7032 (BM).C—G drawn from isotype, C Purpus 7032 
(GH). H drawn from L. Williams et al. 26002 (F). 


280 SIDA 19(2) 


1.3-1.9 cm wide, apically acute, early caducous; secondary branches similar to the pe- 
duncles, but 0.9-2.2 cm long; floral bracts persistent, membranaceous, ovate to oblong, 
0.7-1.8 mm long, 0.3-0.7 mm wide, apically acute, the midrib inconspicuous, the sec- 
ondary veins obscure, densely and prominently punctate and punctate-lineate, glabrous 
above, sparsely erect glandular papillate below, the margins entire, hyaline, sparsely glan- 
dular-ciliate; pedicels slender, terete, 6.7-10.3 mm long, inconspicuously punctate and 
punctate-lineate, densely, erect glandular papillate. Flowers 5-merous, membranaceous, 
pink to reddish-violet; calyx 1.6-1.8 mm long, the tube 0.3-0.5 mm long, the lobes ovate 
to lanceolate, 1.1-1.5 mm long, 0.6-0.8 mm wide near the base, asymmetric, apically 
acute to rounded, prominently punctate and punctate-lineate, glabrous within, tomen- 
tum of scattered, erect glandular papillae, the margins entire, minutely erose, hyaline, 
sparsely glandular-ciliolate; corolla 5-5.3 mm long, the tube 0.9-1.1 mm long, the lobes 
connate basally, ovate to lanceolate, 3.9-4.4 mm long, 2.5-2.8 mm wide near the base, 
apically acute to rounded, prominently punctate and punctate-lineate, glabrous through- 
out, the margins entire;stamens 3.1-3.3 mm long;the filaments 1.9-2.1 mm long, apically 
free, 0.3-1.7 mm long, connate basally into an elobate tube, 0.4-0.6 mm long, free from 
the corolla tube, epunctate, glabrous, the anthers free, ovate, 1.4-1.7 mm long, 0.8-0.9 
mm wide near the base, apically apiculate, basally sagittate, dehiscent by wide, apically 
continuous longitudinal slits, the connective epunctate,; pistil obturbinate,4—4.2 mm long, 
glabrous; ovary 0.9-1.1 mm long, the style 2.9-3.3 mm long, slender, erect, inconspicu- 
ously punctate; stigma punctiform; ovules 11-15, pluriseriate. Fruit globose, 5-6.4 mm in 
diam., conspicuously and prominently punctate, the style base persistent. 

Distribution.—Ardisia rarescens is restricted to the extreme SE corner of Chiapas in 
the Sierra Madre Mountains, and in the adjacent area in the Departments of San Marcos 
and Quetzaltenango in ae oe at 1,300-2,400 m elevation. 

Ecology and el n status. occurs in montane rain forest and 
cloud forests. oe of it its restricted distribution, it should be considered threatened. 

Etymology.—The specific epithet comes from the combination “rare” meaning far 
apart, sparsely and “-escens’ to indicate a process of becoming without full attainment 
reached, referring to the sparse tomentum. 

Common Name.—"Cereza morada” UU. Steyermark 33656) 


Specimens examined. MEXICO. Cxiapas: Mpio. Motozintla de Mendoza, 45-50 km NE of Huixtla 
along road to Motozintla, 1,900 m, 17 Nov 1971 (fr), D. Breedlove & A. Smith 22592 (LL, MO): 28 Dec 
1972 (fr), D. Breedlove & R. Thorn 31017 (LL, MO); Municipio of Motozintla de Mendoza, SW side of 
Cerro Mozotal, 11 km NW of the junction of the road to Motozintla along the road to El Porvenir 
and Siltepec, 2,100 m, 21 Nov 1976 (fr), D. Breedlove 41608 (MO); 23 Nov 1981 (fr), D. Breedlove & B. 
Bartholomew 55740 (LL, NY); Municipio of Motozintla de Mendoza, between El Rosario and Ojo de 
Aqua along road to Niquivil, 1,770 m, 10 Jun 1988 (fl), D. Breedlove 691 14 (CAS); Above El Rosario, 8 mi 
S of Motozintla, 1,800 m, 10 Jul 1977 (fl), 7. Croat 40740 (LL, MO); Mt. Ovando, without elev, 24 Dic 
1936 (fr), E. Matuda 0680 (US); Mt. Pasitar, without elev., 3-4 Aug 1937 (fl), E Matuda 1641 (LL);4 Aug 
1937 (fl), E. Matuda S-206 (A, K, LL, MO, NY, US); Mt. Ovando, 2,100 m, 14-18 Nov 1939 (fl), & Matuda 
3942 (A, NY); Mt. Ovando, Escuintla, without elev., 1-16 Jul 1940 (fl), E. Matuda 4180 (A, GH, LL, MO, 
NY); Saxchanal, Sierra Madre, 2,700 m, 1 Jul 1941 (fl), &. Matuda 4306 (A, F, LL, MO, NY); Carlas, near 


— 


281 


Motozintla, 2,176 m, Apr 1945 (fl), E. Matuda 55717 (LL 2-sheets, TEX); San Juan Panama, Escuintla, 
1,834 m, 23 Jul 1948 (fl), E. Matuda 18148 (F); Along the dirt road to Siltepec, past Ejido Benito Juarez, 
ca. 12 km from the turnoff from Mexican highway 190, S of Motozintla, 15° 20' N, 92° 15'W, 2,100 m, 
9 May 1987 (fl), J. Miller & J. Myers 2778 (BRIT, F, MEXU, MO);Tapachula, Finca Chinincé, 1,500 m,17 Aug 
1961 (fl, fr), H. Schwabe s.n. (B 2-sheets); Municipio of Motozintla de Mendoza, track from Ejido 
Boquerén to Cerro Boquerén, 15°?15' N, 92° 17' W, 2,400 m, 9 Feb 1990 (fr), P Stafford et al. 347 (BM, 
MO); Municipio of Motozintla, Buenos Aires, 1,900 m, 31 Jul 1986 (fl), £. Ventura y E. Lopez 3985 (MO). 
GUATEMALA. Quezattenanco: Volcan Zunil, 6,100 ft [1,859 m],5 Aug 1934 (fl), A. Skutch 948 (F, NY); 
Lower S-facing slopes of Volcan Santa Maria, between Santa Maria de Jesus and Calahuaché, along 
great barranco between as Pirineos a San Juan Patzulfn, 1,300-1,500 m, 6 Jan 1940 (fr), J 
Steyermark 33656 (F, LL). San Marcos: 6 mi SW of town of Tajumulco, NW slope of Volcdn ‘iplaiulcs 
along Rio eee m,26 ate a (ster.), J. Steyermark 36666 (F); Above Finca El Porvenir, 
between “Todos Santos Chiquitos” and “Loma de la Paloma:, S-facing slopes of Volcan Tajumulco, 
1,400-1,700 m, 8 Mar 1940 (ster.), J. Steyermark 37283 (F, LL); 1,300-1,500 m, 16 Mar 1940 (ster), J. 
Steyermark 37981 (F, LL); Near Aldea Fraternidad, between San Rafael Pie de la Cuesta and Palo Gordo, 
W-facing slope of the Sierra Madre Mountains, 1,800-2,400 m, 10-18 Dec 1963 (fr), L. Williams et al. 
26002 (F); (fl, fr), 26004 (F 2-sheets); (fr), 26707 (NY); Outer slopes of Tajumulco Volcano, Sierra Madre 

Mountains about 8-10 km W of San Marcos,ca.2,300 m, 31 Dec 1964-1 Jan 1965 (fr), L. Williams etal 
267994 (US). 


— 


~~ 


ACKNOWLEDGMENTS 


We thank the Missouri Botanical Garden and the Flora Mesoamericana Project, for fund- 
ing that allowed J. Ricketson (MO) to travel with J.Pipoly (FTG) on visits to the C.L.Lundell 
Herbarium (LL), housed at the University of Texas at Austin. Without access to that critical 
collection, assembled by C.L. Lundell over a period of nearly 60 years, the present study 
would not have been possible.We also thank the curators of the herbaria cited for loans 
of specimens. We are grateful to the staff of LL and TEX, especially Billie L. Turner, Tom 
Wendt, Carol Todzia, Beryl Simpson, and José Panero, for their cooperation and hospital- 
ity. We are also grateful to those who have been so instrumental in assisting us in our 
work, including Gerrit and Jeany Davidse, Linda Oestry, Mary Bard, and Catherine Mayo 
(MO), Barney Lipscomb, Debra Trock and Jim Rivers (BRIT). Illustrations were prepared by 
the junior author. Reviews of the manuscript by Gerrit Davidse and Roy Gereau and 
meticulous copy editing by Barney Lipscomb, improved the presentation of the paper. 


REFERENCES 


Cuen, J.and JJ. Pipoy. 1996. Myrsinaceae. In:Wu Zheng-yi and P. Raven, eds. Flora of China. 
Science Press, Beijing, People’s Republic of China, and Missouri Botanical Garden, St. 
Louis, MO, U.S.A. 15:1-38, Illus. 1-20. 

Conran, J. 1995. Family distributions in the Liliiflorae and their biogeographical implica- 
tions. J. Biogeog. 22:1023-1034. 

Gentry, A. 1983.Alstonia (Apocynaceae): A 
Ann. Missouri Bot. Gard. 70:206-207. 

Hickey, L. 1984. A revised classification of the architecture of dicotyledonous leaves. Pp. 
25-39.In: C.R. Metcalfe and L. Chalk (editors). Anatomy of the Dicotyledons. Vol. 1. Sys- 


r paleotropical genus in Central America. 


282 SIDA 19(2) 


tematic anatomy of leaf and stem, with a brief history of the subject. Clarendon Press. 
Oxford, U.K. 

Lavin, M. and M. Luckow. 1993. Origins and relationships of tropical North America in the 
context of the Boreotropics Hypothesis. Amer. J. Bot. 80:1-14. 

Linotey, J. 1848. Illustrated dictionary of botanical terms. Excerpt from illustrated dictio- 
nary of botanical terms by John Lindley. Pp. 346-383. (Reprint, with an introduction by 
Alice Eastwood, Stanford University, School of Earth Sciences 1964]. 

Metcatre, C.R. 1984.Some basic types of cells and tissues. Pp.54—62.|n:C.R. Metcalfe and L. 
Chalk, eds. Anatomy of the Dicotyledons .Vol. 1. Systematic anatomy of leaf and stem, 
with a brief history of the subject. Clarendon Press. Oxford, U. K. 

Mez, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich lV. 236(Heft 9):1-437, 

Pipoty, J.J. 1987. A systematic revision of the genus Cybianthus subgenus Grammadenia 
(Myrsinaceae). Mem. New York Bot. Gard. 43:1—76. 

Pipoty, J.J. 1992. The genus Cybianthus subgenus Conomorpha (Myrsinaceae) in Guayana. 
Ann. Missouri Bot. Gard. 79:908-95 7. 

Pipoty, J.J. and J. Ricketson. 1999. Discovery of the Indo-Malesian genus Hymenandra 
(Myrsinaceae) in the Neotropics, and its Boreotropical implications. Sida 18:701-746. 

THEOBALD, W.L., J. KRAHULIK, and R.C. RoLuins. 1984.Trichome description and classification. Pp. 
40-53.In:C.R. Metcalfe and L. Chalk (editors), Anatomy of the Dicotyledons. Vol. 1. Sys- 
tematic anatomy of leaf and stem, with a brief history of the subject. Clarendon Press. 
Oxford, U.K. 

TirFNey, B. 1985a. Perspectives on the origin of the floristic similarity between eastern Asia 
and eastern North America. J. Arnold Arbor. 66:73-94. 

TirFNey, B. 1985b. The Eocene North Atlantic land bridge: its importance in Tertiary and 
modern phytogeography of the northern hemisphere. J. Arnold Arbor. 66:243-273. 
Wenot, T. 1988. Chiangiodendron (Flacourtiaceae: Pangieae),a new genus from southeast- 
ern Mexico representing a new tribe for the New World flora. Syst. Bot. 13:435-441. 
Wenort, T. 1989. Las selvas de Uxpanapa, Veracruz-Oaxaca, Mexico: evidencia de refugios 
floristicos cenozoicos. Anales Inst. Biol. Univ. Nac. Auton. México, Ser. Bot. 58:29-54. 
Wenot, T. 1993. Composition, floristic affinities and origins of the canopy tree flora of the 
Mexican Atlantic slope rain forests. Pp. 595-680. In:T. Ramawoorthy, R. Bye, A. Lot and J. 
Fa, eds. Biological diversity of Mexico: Origins and Diversity. Oxford University Press., 

New York 

Wot re, J. 1975.Some aspects of plant geography of the northern hemisphere during the 
late Cretaceous and Tertiary. Ann. Missouri Bot. Gard. 74:264-279. 

Zona, S. 1990.A monograph of Saba! (Arecaceae: Coryphoideae). Aliso 12:583-666. 


LIST OF EXSICCATAE 


Breedlove, D. 41608; 69114. Breedlove, D. & Bartholomew, B. 55740. Breedlove, D. & Smith, A. 22592. 
Breedlove, D. & Thorn, R. 31017. 


Croat, 1.40740. 


Matuda, E. S-206; 0680; 1641; 3942; 4180; 4306; 5511; 18148. Miller, J.& Myers, J.2778. 


Purpus, C. 7032 (type) 

Skutch, A. 948. Stafford, P. et al. 347. Steyermark, J. 33656; 36666; 37283; 37981. 
Ventura, E.y Lopez, E. 3985. 

Williams, L. et al. 26002; 26004; 26101; 26799A. 


284 SIDA 19(2) 


BOOK REVIEW 


VIRGINIA Scott JeNkiNs. 2000. Bananas. An American History. (ISBN 1-56098-966-1, pbk.). 
Smithsonian Institute Press, 470 L’Enfant Plaza Suite 7100, Washington, DC 20560- 
0950, U.S.A.$16.95 pbk., 232 pp., 36 b&w photos. 


From the author of 1994's The Lawn: A History of an American Obsessi a book about Amer 
can culture, politics, and trade as related to al things, the banana.From its days of an exotic fruit for 
the riches (late 1800's), to its days of being cheap food for the poor, to its modern day ubiquity and 
subsequent absorption into American culture and everyday life, the banana has been a dynamic 
part of the last century of American histo 

From the introduction," ...the ees leas was : see a bial ited growth in the 
banana-importing industry with the rise of giant mult ananas became the 
cheapest fruit in the grocery store throughout the year and taken for ae by consumers. The 
banana lost its exotic image and disappeared from the formal dinner table as it became the most 
widely eaten fruit in the United States.” Not to indicate that this book is about the rise of the ba- 
nana and its fall to mediocrity, however, this book is entertaining reading about the history of a 
_ fruit in the American diet. 

n eight chapters, Jenkins covers everything you ever wanted to know apoul bananas. The 

first s.chapte covers the introduction of the banana to America in the ninet ntury. The sec- 

chapter traces the Sena of the major American ea lon eonnoanies Chap: 
ter 2 discusses the growth of modern transportation systems that helped put the banana in more 
homes nationwide. Chapter 4 is about the marketing and selling of bananas. Chapter five looks at 
public health and sanitation issues in relation to ue spopiiailzetion of the ae et 6,“Eat- 
ing Bananas,’ Mao wea evening fom! he 

anana split. “The ere sHaotee: is devoted to two Ameri- 

can cities, Filet Kentucky) and South ae Tennessee, the banana ae of the wolle: thanks, 
of course, to a connection in Central America. The final chapter looks at how | 
rooted in American life despite the fact that they have never been a selection of the fruit of the 
month club. Closing the book is an appendix of songs about bananas, notes, and an index. 

Highlights of the book include accounts of the history of import trading of bananas with 
Central American countries and integration of this exotic fruit as a staple of an American diet. Ba- 
nanas came from Jamaica, Honduras, various parts of Latin America and the Hawaiian islands and 
due to its versatility and nutritional value became a part of the American diet. Corporations that 
paid for advertisements and teaching items discussing oe a eee importance of the banana, 
its germ free packaging, and importantly its taste, no doubt had a small part to play in the popular- 
ization of the fruit. Marketed to children (read: future face and the elderly, bananas became a 
necessity, and mostly, a part of breakfasts and desserts. 

This book is intended for lay readers and it would be safe to presume that’s where you will 
find the bulk of the readership. While it is extensively researched, it doesn’t carry with it the socio- 
cultural insight and interpretation of Sidney Mintz’s Tasting Food, Tasting Freedom: Excursions into 
Eating, Culture and the Past or his works on sugar and the Caribbean. It could work in classes of 
American history, political science, and anthropology due to it being both short and inexpensive, 
but only in conjunction with other more scientific books. In summary, it is enjoyable reading and 
anyone interested in pence culture international trade and politics and, of course, bananas will 
be interested.—Kevin D. Janni, | Research Institute of Texas kjanni@brit.org. 


SIDA 19(2): 284. 2000 


SENECIO QUAYLEI (ASTERACEAE: SENECIONEAE), 
A NEW SPECIES FROM NORTH CENTRAL TEXAS, U.S.A 


Theodore M. Barkley 
Botanical Research Institute of Texas 
09 Pecan Street 
Fort Worth, TX 76102-4060, U.S.A. 
barkley@brit.org 


ABSTRACT 


Senecio quaylei, a new species from Texas, U.S.A.,is described. 


RESUMEN 


Se describe Senecio quaylei, una especie nueva de Texas, U.S.A. 


Jeffrey Quayle, an astute amateur naturalist in north central Texas, recently submitted an 
unusual Senecio to BRIT for determination. It was presumed that the plant was simply a 
recent or otherwise unrecorded introduction, but a survey of the literature and several 
herbaria, plus the opinions of several colleaques, suggested no possible matches, so it is 
offered here as a new species, Senecio quaylei. 

The only known occurrence of the plant is a conspicuous colony of about 15 indi- 
viduals, growing waist high in a weedy roadside ditch, along with Cirsium, Geranium, 
Sonchus, Verbena, and much dead vegetation from the previous year. The new species 
vaguely resembles Senecio ampullaceus Hook.,a Texas endemic of disturbed open sites 
in the central part of the state, however, S. quaylei is glabrous and notably coarse, to 12 
dm tall, while S$. ampullaceus is conspicuously hairy but unevenly glabrate in age and 
typically only 3-7 dm tall. In addition, the new species has large, broad, and clasping 
cauline leaves. These characteristics combine to give the new species a distinctive gross 
aspect. Four of the plants were collected to make the specimens needed for documen- 
tation, and they were divided into eleven herbarium sheets. 

Structurally, the stems are hefty, with the proximal third some 10-15 mm in diam- 
eter, but they are hollow and thin-walled. There is a conspicuous purplish-red layer in the 
sub-epidermal region of freshly cut proximal stems; the red color fades as the specimen 
dries. The leaves have unevenly scattered light-brownish spots that are caused by a spe- 
cies of Coleosporium,a heteroecious rust, fide Dr. Joe Hennen,a mycologist at BRIT and an 
authority on rust fungi. 

The biology of S. quaylei is unknown and only suggested from inference. It appears 
to be an annual, with a short, narrow taproot and a tuft of abundant, thin, fibrous roots 
that are weakly branching. The most mature ovaries in the specimens collected are 
wrinkled and unfilled, indicating that they are sterile. The pollen is of uneven size and 


SIDA 19(2): 285 — 289. 2000 


286 SIDA 19(2) 


shape, suggesting that it is of reduced or doubtful viability. The chromosome number is 
undetermined. 

The resemblance of Senecio quaylei to Senecio ampullaceus and other typical sene- 
cios (Barkley 1999) in both structure and aspect clearly places the new species in Sene- 
cio s. str; rigorous quantitative analyses are yet to be done. It is possible that additional 
data from cytology, pollen morphology, and comparative biochemisty, may suggest that 
the new species is allied to Packera, but that notion seems remote. Senecio quaylei is of 
poor fit in the keys to Senecio in the treatment in the North American Flora (Barkley 
1978), for there the possesion of leaves that are progressively reduced upwards carries 
it to the Senecio integerrimus assemblage, a group with unbranched fleshy-fibrous roots 
and of very difetent gross aspect. If the initial couplet of the key is ignored, it falls into 
the Annui group, along with Senecio ampullaceus. 

Recognition of Senecio quaylei generates speculation on its significance in the flora. 
The possibility that it is merely an exotic waif cannot be discounted, but neither can it be 
said that it is not a rare member of the regional flora. This notion is compatible with the 
review by Ertter (2000) on the occurrences and recognition of distinctive species of lim- 
ited occurrence in North America. 


Senecio quaylei [.M. Barkley, sp. nov. tFIgS: 1, 2 TPE U. S.A. TEXAS. PARKER Co.: waste ground 
adjacent to Lake sates State Trailway ( of-way), at Holder 
IR 


Chape n,ca.2 mi N of Hwy 180, Eo Mineial Wels ou 45 N,98° 02'W, 29 Apr 
2000, T.-M. saan M. Barkley, RJ.O’Kennon, & BRIT,M 
TEX, UC) 


A Senecioni ampullaceo similis sed differt altitudine ampliore (7-12 dm vs. 3-7 dm) et foliis caulinis 
glabris amplectentibus ovati-lanceolatis folia basalia amplitudine fere aequans. 


Annual, 8-12 dm tall, glabrous throughout or with a few inconspicuous hairs on the 
peduncles and phyllaries. Stems single, striate, the proximal third 10-15 mm in diameter, 
narrower distally, hollow, with a conspicuous reddish subepidermal layer when freshly 
cut; arising from a short, thin taproot surrounded by abundant, thin and sparingly 
branched fibrous roots. Basal and proximal cauline leaves with blades ovate, mostly 12- 
24 cm long and 8-12 cm wide, margins wavy, with a few scattered minute denticles, 
midvein prominent and lateral veins less conspicuous in dried specimens, tapering or 
gently contracted to a distinct petiole, ca. 2/3 the length of the blade. Middle cauline 
leaves with blades nearly as large as the basal leaves, ovate to broadly lanceolate, sessile 
and clasping. Distal cauline leaves lanceolate to linear-lanceolate, 6-14 cm long, sessile, 
the distal most further reduced and bractlike. Capitulescences terminal or arising from 
the axils of the upper leaves; fundamentally corymbiform cymes of 20-40 capitula, or a 
close cluster of corymbiform cymules. /nvolucres cylindrical or turbinate to weakly cam- 
panulate, each subtended by a weakly defined calyculus of 2-7 linear bracteoles, 1-3 
mm long, margins hairy. Phyllaries mostly 13,+ 8 mm long, green with hyaline margins, 
abaxial surfaces with short hairs distally, the apex with a minute tuft of hyaline hairs. 
Corollas yellow. Ray florets mostly 8, pistillate; corollas ca. 11 mm long, tubes 4 mm and 


BARKLEY, A NEW SENECIO FROM NORTH CENTRAL TEXAS 


Fic. 1. Senecio quaylei A. habit, B capitulum C. disk floret, D. ray floret E 


by Linny Heagy. 


288 SIDA 19(2) 


BARKLEY, A NEW SENECIO FROM NORTH CENTRAL TEXAS 289 


laminae ca.7 mm long, 2.5 mm wide. Disk florets 20-40, bisexual; corollas 7(-10) mm long, 
tube and limb (including lobes) of about oe length, corolla lobes triangular, + 0.5 mm 
long. Cypselae not seen, oldest ovaries wrinkled, empty, 1-3 mm long, pubescent through- 
out. Pappus of abundant white, minutely barbellate bristles in a single series, 6-7 mm 
long. Chromosome number unknown. 

Etymology.—The specific epithet quaylei commemorates Jeffrey Quayle, the dis- 
coverer of the new species. 

Distribution.—Endemic to Texas. There are no known collections other than the type 
collection. 


KEY TO SPECIES OF SENECIO S. STR. IN NORTH CENTRAL TEXAS 


This key covers the range of the recently published Illustrated Flora of North Central 
Texas (Diggs et al. 1999) 


1. Capitula discoid; ray florets rarely present and then with laminae of corollas less 
than 1 mm long; phyllaries and calyculate bracts with prominent triangular — 
tips; leaves shallowly pinnate. Introduced weeds. 1.8. vulgaris 
1. Capitula radiate; laminae of ray corollas 7-15 mm long; phyllaries and calyculate 
bracts green or gray-green tipped; leaves shallowly toothed to subentire. Native or 
apparently so 
2. Herbage woole pubescent, or at most unevenly glabrat 
leaves lanceolate and rarely more than 10 cm long, seen ee stems 3- 


iddl |; 


7(+) dm tall. S.ampullaceus 
2. Herbage glabrous or nearly so, middl line | te or broadly lanceolate, 
10-20 cm long, prominently clasping; stems mostly 8-12 dm tall. S. quaylei 
ACKNOWLEDGMENTS 


It is a pleasure to acknowledge the assistance of Joe Hennen, Barney Lipscomb, Guy 
Nesom, Robert O’Kennon, John Pruski, Peter Raven, John Strother, Debra Trock, and oth- 
ers who offered advice on the nature of this plant.Guy Nesom kindly translated the Latin 
diagnosis and Linny Heagy prepared the illustration. 


REFERENCES 


Barktey, I.M.1999.The segregates of Senecio, s./. and Cacalia,s.l. in the flora of North America 
North of Mexico. Sida 18 

Barkley, T.M. 1978. Senecio. In: C.T. Rogerson et al., eds. North American flora ser. 2, 10: 

0-139 

Diacs, G.M., B.L. Liescoma, and R.J. O’KeNNon. 1999. Shinners & Mahler’s Illustrated flora of 
north central Texas. Sida, Bot.Misc. 16. Austin College and Botanical Research Institute 
of Texas, Fort Worth. 

Ertter, B. 2000. Floristic surprises in North America north of Mexico. Ann. Missouri Bot. 
Gard. 87:81-109. 


290 SIDA 19(2) 


BOOK REVIEW 


JupiTH Sumner, 2000, The N | Hi of Medicinal Plants. (ISBN 0-881 92-483-0, hbk.). 
Timber Press. Inc., The Haseltine Building, 1 33 S.W. Second Avenue. Suite 450. Port- 
land, OR 97204-3527,U.S.A.(503-227-2878,503-227-3070 fax; www.timberpress.com). 
$24.95 hbk. 235 pp.Line drawings, 30 color plates. 


Despite the modest title, The Natural History of Medicinal Plants, is much more than just a historical 
account of the uses of plant medicine. The author, Judith Sumner of the Arnold Arboretum, covers 
that and in doing so reviews everything from medicinal chemistry to zoopharmacognosy. Though 
not necessarily designed as such, this book is an excellent overview of - contemporary issues 
concerning medicinal plants. Complete with a foreword by the ubiquit Aark Plotkin and being 
just over 200 pages, this book should attract a wide readership and peciaee so. The writing 
within bares all the wisdom of a historian, botanist, and enthusiast of a subject whose time has 
come. Indeed, westerners are experiencing a renaissance of interest in medicinal plants and their 
potential to treat modern diseases. And new drugs aren't the only concern; conservation of global 
biodiversity is an increasingly salient issue for academic and lay readers.While | refer to the title as 
eing modest, the author's goal to,”...connect the human concerns of botanical medicines and 
ethnobotany with the role of medicinal plants and their secondary compounds in nature” is any- 
thing but and she admirably succeeds 
The book is laid out in ten chapters: 1) A Brief History of Medicinal Botany, 2) Acquiring Knowl- 
edge, 3) Medicinal Plants in Nature, 4) Toxins and Cures: A Cabinet of Plant Chemicals, 5) Defensive 
Strategies and Plant Chemistry, 6) erean Discoveries, 7) Zoopharmacognosy and Botanical 
Toxins, 8) Chemical Prospecting and New Plant Medicines, 9) Protecting Medicinal pene and 
Knowledge, 10) Herbal Histories, Considerations, and Caveats. A brief- 1/2a page- glossary, three 
pages eon reading and an ne Ow ee ete etiowean oe book ae covers 
plant m dispersal 
and ie of medicinal plants, alkaloids, curare plants, coca chimpanzees ‘and self: medica- 
tion, cancer drugs from plants, and tropical conservation- just to name a few 
Str points are in chapters four through six in which the author covers plant compounds, 
ee See and significant discoveries in the field. Chapters eight and nine are of interest 
to the eae erinobetans aviong US, aS eouENNE! GIbGLS ES new plant drugs, future prospects, 
ownership of ti ut, Sumner does more 
than just cover big- “ticket issues of ethnobotany; this is a “a history book after all. Chapter’s one, three, 
and ten are exceptionally written historical discussions of plant medicines, their discovery and 
uses. It is perhaps the author's skillful writing that makes this book so valuable. While written for a 
lay audience, this book is sure to hold the attention and interest of any scholar. The breadth of 
academia covered in this relatively small book is remarkable and for twenty-five dollars, a bargain. 
imply put, anyone interested in medicinal eel, an history, anthropology and 
conservation should have this book on their shelves. The book is not only accessible to a large 
academic audience, but also lay readers of a wide ales of interests. The see aes 
alone is enough to say that it is only a matter of time before this book becomes a classic. The 
Natural History of Medicinal Plants is well suited for college level classes covering sere es 
pology, Ethnobotany, and Medical Botany, as an introduction to a field with a myriad of contempo- 
rary concerns.—Kevin D. Janni, B | Research Institute of Texas, kianni@brit.org. 


SIDA 19(2): 290. 2000 


SOLIDAGO VILLOSICARPA (ASTERACEAE: ASTEREAE), 
A RARE NEW SOUTHEASTERN COASTAL PLAIN ENDEMIC 


R.J.LeBlond 


North Carolina Natural Heritage Program 
C ain Office 


PO. Box 787 
Richlands, NC 28574, U.S.A. 
ABSTRACT 


Solidago villosicarpa, sp. nov., is described from the outer coastal plain of southeastern North Caro- 
lina, where collections have been made from four counties. Although it apparently is extant in only 
two of these four counties, its habitat and evident adaptation to disturbance suggest that the new 
species may be more frequent along the Carolina coast than is currently known. Seg? villosicarpa 
is placed as amember of sect.and subsect. Solidago and is distinctive in it f pubes 
cent stems, glabrous to glabrate leaves, thyrsoid inflorescence, large heads with bright eau 
low rays, densely villous achenes, and late flowering. 


RESUMEN 


Se decribeSolidago villosicarpa, sp. nov., de la llanura costera externa del sudeste de Carolina del 
Ne ore se han mene ‘eOISECIONE en cuatro condados. Aunque aparentemente existe en 
,su habitat y evidente adaptacidn a las perturbaciones sugieren 
que la nueva especie puede ser mas frecuente a lo largo de la costa de Carolina de lo que se 
conoce actualmente. Solidago villosicarpa se coloca como un miembro de la sect. y subsect. Sol- 
idago y es distintiva su compInacien de tales orbital eles aleelas fe) ge DIescenles 
inflorescencia tirsoide, cay g gulas amarillo lim te villosos 
y floracion tardia. 


INTRODUCTION 


Early collections of a distinctive goldenrod species were made from “live-oak scrub”on a 
barrier island in Brunswick County, North Carolina, in 1949 and 1950. Specimens from 
the 1950 collection were annotated as S. sciaphila Steele (Fox et al. 1952), a plant other- 
wise known only from sandstone and calcareous habitats in the Upper Mississippi River 
region (Minnesota, Wisconsin, lowa, and Illinois). A subsequent collection of the North 
Carolina entity was made in 1963 from a sandy roadside in neighboring New Hanover 
County,N.C.,and annotated as S.erecta Pursh (NCU). Three more populations were found 
1991-1998 in Onslow County, N.C., during a natural area inventory of Camp Lejeune 
Marine Corps Base by the North Carolina Natural Heritage Program (NCNHP),and a popu- 
lation was found in Pender County in 1998 during another NCNHP inventory. Only the 
Onslow and Pender county populations are known to be extant. 

The original collections of the North Carolina entity from 1949 and 1950 have a 
curious history. According to Fox et al.(1952) the 1950 collection (Godfrey 50963 & Boyce) 
was identified as Solidago sciaphila“by Dr. Arthur Cronquist and it was rechecked by him 


SIDA 19(2): 291 - 300. 2000 


292 SIDA 19(2) 


after several duplicate specimens were sent to him. Godfrey also checked specimens of 
the collection against material of the species at the Gray Herbarium, and he concurs 
with Dr. Cronquist in his determination. We are at a complete loss to account for the 
occurrence of this goldenrod so far from the hitherto known range of its distribution.” 
The 1949 collection (Godfrey 50132) is not mentioned in the 1952 Rhodora article. A speci- 
men from this collection was found in the folder for undetermined Solidago specimens 
at the N.C. State University herbarium (NCSC) during the current investigation. The label 
of this specimen (herbarium #19998) has a typed and penciled portion. The typed por- 
tion reads “Solidago ...|n live oak scrub on the sand dunes, Long Beach." The penciled 
portion reads “sp. aff. S. sciaphila Steele, less closely aff. S. glutinosa sens. lat. May need a 
name. A.C. 3-22-50."A.C." undoubtedly is Arthur Cronquist. These circumstances suggest 
that the 1949 collection had been forgotten by the time the 1950 collection was deter- 
mined. Adding to the mystery, the three specimens from the 1950 collection at NCSC 
(herbarium #’s 26549, 26550, 34039) were annotated as S. sciaphila by H.E. Ahles, not 
Cronquist. Yet there is no mention of S. sciaphila occurring in the Carolinas in Radford et 
al. (1968) (Ahles was responsible for the treatment of Asteraceae), nor apparently in any 
other flora, treatment, or checklist since that time. No specimens from the 1949 and 
1950 collections are known from other herbaria, including the New York Botanical Gar- 
den (Kallunki 1998), where Cronquist worked. 

This distinctive, rare,and narrowly endemic North Carolina plant does indeed “need 
aname,"and is here described as a new species. 


Solidago villosicarpa | eBlond, sp.nov. (Figs. 1-6). Type: UNITED STATES. NortH Carotina. Onslow 
Co.:in pine-oak-hickory forest with open hardwood understory, Camp Lejeune oe sae 
Base near Salliers Bay, about 0.6 mi W of Tactical Landing Zone Albatross, 13 Oct 2000, RJ. 
LeBlond 5435 and B.A. Sorrie (HoLotyee: US; isoryees: BRIT, FLAS, FSU, GA, GH, MO, NCU, NY, aan 
A ibus di li | il lil b ibratis, infl tia thyrsoidea, 


capitulis grandibus corollis radii vivide citreis, sehen dense Glee: et Fenner serotina. 


Roots wiry, elongate; caudex stout. Stems usually solitary, occasionally loosely cespitose 
by short rhizomes, 4.5-15 dm long, 2.5-6 mm wide 2 cm above caudex, ribbed and 
grooved throughout, the ribs rounded to angled, some decurrent from leaf bases; pu- 
bescent with short stiff spreading or appressed trichomes 0.1-0.3 mm long, many unci- 
nate; stem color medium brown to dark brownish purple below, and light brown, 
stramineous, or purple ae Basal leaf rosette present; basal and lower cauline leaves 
1; | blades 7-14 cm long (not including petiole) by 4-7 
cm wide, elliptic, broadly elliptic, or elliptic-obovate, apices obtuse, subacute, or broadly 
acute; bases cuneate-attenuate, often somewhat abruptly narrowed; petioles 2-7 cm 
long; petiole base non-auriculate but sheathing the stem for 1/4-1/2 of its circumfer- 
ence; blade margins toothed, the sinuses 1-2.5 mm long from base of cavity to tip of 
tooth mucro, the mucros mostly 0.2-0.4 mm long, blunt; teeth margins ciliate-scabrous 
with recurved-ascending trichomes about 0.1-0.2 mm long; primary veins tending to 


the largest, petiolate, t 


aa. 


LEBLOND, A NEW SPECIES OF SOLIDAGO FROM SE UNITED STATES 293 


Fic. 1 oar i "17 * I DI 1A Si 1 +1 ere ral RP ® 1 i} L L 3 aL ege ao 
Flower head at anthesis (top view). D. Flower head at anthesis (side view). E. Pappus and achene. F. Lower stem leaf. 
Drawn from type collection by Margret Mueller. 


SIDA 19(2) 


Fig. 2. Solidago villosicarpa LeBlond. Habitat showing example of paniculately branched thyrse. 


LEBLOND, A NEW SPECIES OF SOLIDAGO FROM SE UNITED STATES 295 


Cali 


L. 


RI d Fic 2 (+t Inft\ <j I 


Fics. 3 g icarp (top lett) simp 
elongate thyrse inflorescence form. Fic. 4 (top right) example 
Lek £ 1 1 L L Fic. C ihka#t. Lnaf+\ 1 


up of the inflorescence. Fic. 6 (bottom right) a well-developed 
basal rosette. 


296 SIDA 19(2) 


be raised on both surfaces, the mid-nerve flattish and squarred at the edges, pale 
stramineous; ultimate nerves distinctly reticulate on both surfaces; adaxial surface gla- 
brous to sparsely pubescent with short stiff trichomes mostly along the mid-nerve and 
larger veins; glabrous to glabrate abaxially; texture thin, papery, brittle when dried; color 
drab green to olive-green, the lower surface barely if at all lighter than the upper. Middle 
and upper cauline leaves gradually reduced upwards, sessile, becoming entire; cauline 
leaves 15-50 below inflorescence. Inflorescence a simple or paniculately branched thyrse; 
when simple (elongate terminal thyrse), the terminal axis bracteate, straight, narrow, and 
cylindric, 7-22 cm long by 3-6 cm wide; when paniculiform, producing straight, elon- 
gate, thyrsoid axillary branches up to 20 cm long; bracteal leaves at the base of the 
branches similar to the cauline leaves, gradually reduced in size upwards. Short second- 
ary branches and peduncles moderately densely to densely invested with stiff, soread- 
ing and ascending, straight and recurved trichomes 0.1-0.4 mm long, which appear res- 
inous and segmented; peduncles 0.5-9 mm long. Heads at anthesis 1.4-1.7 cm wide 
measured from ray tip to ray tip. Involucre 5-8 mm long by 3-5 mm wide at anthesis, the 
summit 6-8 mm wide at maturity. Phyllaries strongly imbricate, the outer shorter, ovate, 
somewhat cucullate, the inner broadly linear;outer phyllaries 1.0-2.0 mm wide, appressed; 
inner phyllaries 0.8-1.5 mm wide, with rounded to subacute apices becoming some- 
what squarrose in age; all phyllaries with a narrow but distinct pale reddish-brown mid- 
rib; surface glabrous, glandular centrally and near the apex, greenish near the apex, cream- 
colored centrally and laterally, the margins hyaline, often lacerate, long-ciliate or 
ciliate-fimbriate (at least near the apex) with cilia 0.1-0.3 mm long. Ray florets 4-8 per 
head, limb in vivo 5~7.5 mm long, 1-2 mm wide, bright lemon-yellow. Disk florets 10-18 
per head, the corolla lobes 1.5—2.2 mm long, the entire disk corolla 4.9-6.8 mm long; 
stigmatic lobes 0.9-1.1 mm long, anthers 2.1-2.2 mm long. Pappus (4.2-)4.7-6.1 mm 
long, occasionally some bristles clavate. Achenes villous with ascending hairs 0.3-0.5(- 
0.7) mm long, the achene body 2.6-2.9 mm long when mature. 


Additional specimens examined: NORTH CAROLINA. Brunswick Co.: in live oak scrub on the sand 
dunes, Long Beach, 22 Oct 1949, R.K. Godfrey 50132 (NCSC); in live-oak scrub thickets on sand dunes, 
Long Beach, 28 Oct 1950, R.K. Godfrey 50963 and S.G. Boyce (NCSC). New Hanover Co.: sandy road- 
side, Pembroke Jones Park, Wrightsville Sound, 29 Nov 1963, A.McCrary 1813 (NCU). Onslow Co.: in 
pine-oak forest with open understory, Camp Lejeune Marine Corps Base near mouth of Frenchs 
Creek, 22 Oct 1991,RJ LeBlond 2622 (pers. herb.);same locality, 18 Oct 1992, RJ. LeBlond 3127 (NCU); 
in pine forest with open understory, Camp Lejeune Marine Corps Base near Salliers Bay, 08 Oct 
1995, RJ. LeBlond 4440 (pers. herb.); same locality, 11 Oct 1998, RJ. LeBlond 5074, A.S. Weakley, and K. 
Patterson (RJL pers. herb.); same locality, 18 Oct 1998, RJ. LeBlond 5082 (DUKE, NCSC, NCU); Camp 
Lejeune Marine Corps Base, SW of Mock-up Road, 0.9 mi SE of NC 172,05 Nov 1998,RJ.LeBlond 5124 
and E. Davis (NCU). Pender Co.: in pine-hardwood forest 0.65 mi WNW of Clarks Landing on Long 
Creek, 30 Sep 1998, RJ. LeBlond 5051 and B.A. Sorrie (NCU). 


DISCUSSION 


The most striking feature of Solidago villosicarpa is the width of the floral heads at anthe- 
sis, which measure 1.4—1.7 cm wide from ray tip to ray tip, and with ray limbs in vivo 5-7.5 


LEBLOND, A NEW SPECIES OF SOLIDAGO FROM SE UNITED STATES 297 


mm long. The size combined with the bright lemon-yellow color of the ray limbs make 
this one of the showier goldenrods. The density of the achene pilosity completely ob- 
scures the body surface with hairs 0.3-0.5(-0.7) mm long. The combination of thyrsoid 
inflorescence, persistent and glabrous to glabrate basal leaves, upwardly reduced cauline 
leaves, and pubescent achenes place the new species in Solidago sect. Solidago subsect. 
Solidago in Nesom’'s 1993 overview of infrageneric goldenrod taxonomy. North Ameri- 
can members of subsect. Solidago are S. calcicola (Fernald) Fernald, S.glomerata Michx.,, S. 
multiradiata Ait.(including =S. cutleri Fernald) ,S.nana A.Gray, S.plumosa Small, S. sciaphila, 
S. simplex Kunth, S. spathulata DC.,and S. spithamaea M.A. Curtis. Solidago villosicarpa dif- 
fers from other members of the subsection by a combination of stem pubescence, floral 
head width at anthesis, involucre length (5-8 mm), pappus length (4.2-6.1 mm), length 
of disk corolla and lobes (4.9-6.8 mm), and nature of achene pubescence (villous, the 
hairs 0.3-0.7 mm long). Also, it flowers late September to early November, one to two 
months later than the others. It superficially resembles S. bicolor L., S. hispida Muhl., and S. 
squarrosa Muhl., members of sect. Solidago subsect. Albigula in Nesom's treatment. It is 
readily distinguished from all three by its pubescent achenes, from S. bicolor and S. hispida 
by its glabrous (-glabrate) leaves, and from S. squarrosa by its appressed outer phyllaries. 
The following key distinguishes southeastern U.S. Solidago taxa with thyrsoid 
inflorescences, basally disposed leaves conspicuously larger than middle and upper 
cauline leaves, and pubescent stems. 


— 


1. Inner phyllaries very narrow, <0.5-0.75 mm wide at mid-length, tapering to slender 
tip 


2. Stems and leaves a pubescent with minute, stiffly spreading viscidulous hairs; 
(ey florets mosuy 9-1 S. puberula 


nave leak Vea Fal bea ge | 
\ D 


“eaves alabrous: ray florets mioedy 6- 9 S. roanensis 
1. Inner phyllaries broader, 0.75-1.5 mm wide at mid-length, the margins usually par- 
and Me ip ial to Han acute 


| Leas | c 
(-sparsely f 


8mm long; ray florets 
te 8; plc (4.2-)4.7-6.1 mm iene aehetes villous S. villosicarpa 
3. Leaves pubescent on both surfaces; involucre 3-6 mm long; ray florets 7-14; pa- 
ppus 2.5-4 mm long; achenes strigose-puberulent or glabrous 
4. Achenes strigose-puberulent; disk flowers 14-27; larger leaves 3.5-12 cm long; 
Ale ile werner in spring S.verna 
4. A ture): disk fl i 
i larger leas 8-20 cm long; plants Honea late summer-fall 
5, Ray limbs white (rarely yellow), often turning yellowish in pe phyllaries 


whitish ie stramineous, usually wit a well- cee green tip S. bicolor 
5. R Ray | to orange I lI is the tip weak 
or not at all a S. hispida var. hispida 


Although similar to Solidago sciaphila in overall habit, S. villosicarpa is distinguished by 
several characters, particularly within the inflorescence (Table 1).The stem of S. villosicarpa 
is pubescent throughout, while that of S. sciaphila is normally glabrous below the 


298 SIDA 19(2) 


otf 


losicarpa from S. sciaphila. 


Taste 1.A comparison of floral characters used to distinguish Solidago vil 


Character Solidago villosicarpa Solidago sciaphila 

Involucre length 5-8 mm 3-6 mm 

Phyllary cilia se 0.1-0.3 mm <0.05-0.1(-0.2) mm 
flower limb length 5-7.5mm 2-4mm 

Disk corolla engin (limb and lobes) 49-68 mm 3.2-4.9 mm 

Disk corolla lobe length 1.5-2.2 mm 0.8-1.4mm 

Pappus length (4.2-)4.7-6.1 mm 2.3-3.8 mm 

Density of achene pubescence, villous, 0.3-0.5(—0.7) mm cea ' oe 
length of hairs hairy, 0.1 


inflorescence (Fernald 1950;Gleason 1952;Gleason & Cronquist 1991), though S. sciaphila 
occasionally can have sparsely to moderately pubescent stems (frequently so just below 
the inflorescence). The basal and lower leaves of S. villosicarpa tend to be sparsely pubes- 
cent to glabrous adaxially and glabrous abaxially, while those of S. sciaphila tend to be 
either glabrous on both surfaces or, less frequently, sparsely pubescent on both surfaces. 

With so few collections and known populations, the ecology of Solidago villosicarpa 
is only partially understood. The 1949 and 1950 Brunswick County collections are from 
“live-oak scrub" o vee scrub thickets” on the Long Beach coastal barrier island. This 
likely is either Maritime Evergreen Forest or Maritime Shrub (Schafale and Weakley 1990). 
Maritime Evergreen Forest typically has a canopy dominated by Quercus virginiana, usu- 
ally with Pinus taeda and Q. hemisphaerica. Characteristic understory species include 
Juniperus virginiana var. silicicola, llex vomitoria, Persea borbonia, P. palustris, and Osmanthus 
americana. Wind-borne sand and salt spray often produce dense thickets along the 
ocean-facing side of such forests.|n areas closer to the ocean or more exposed, the Mari- 
time Shrub community forms. It is characterized by a dense growth of such shrubs as 
C (=Myrica cerifera var.cerifera), lex vomitoria, Baccharis halimifolia, and 
stunted Juniperus virginiana var. silicicola and Quercus virginiana. 

The three Solidago villosicarpa sites in Camp Lejeune Marine Corps Base in Onslow 
County have been altered by past logging. Canopies at all three sites are dominated by 
pine (Pinus taeda), with hickory (Carya glabra var.megacarpa or C.alba) and/or oak (Quercus 
falcata, Q. nigra, and Q. stellata the most common). Frequent understory species are Q. 
margarettiae, Liquidambar styraciflua, and Ilex opaca. This composition suggests the Dry 
or Dry-Mesic Oak-Hickory Forest natural community of Schafale and Weakley (1990), 
perhaps transitional to Coastal Fringe Evergreen Forest (ibid.) at the site near Salliers Bay, 
where Q. virginiana is a subcanopy component. Two of the sites occur on excessively 
drained Wando fine sand entisol, and the third site occurs on well-drained Marvyn and 
Norfolk loamy fine sand ultisol. 

The site in Pender County near Clarks Landing occurs on slopes above a drain; it 
also has been disturbed by past logging. Tree species include Pinus taeda, Acer rubrum 
var. trilobum, Gordonia lasianthus, Quercus nigra, and seedling Pinus palustris. Understory 


LEBLOND, A NEW SPECIES OF SOLIDAGO FROM SE UNITED STATES 299 


= 


and shrub species include /lex opaca, Lyonia lucida, and Vaccinium arboreum. This mix 
suggests a former longleaf pine community grading downslope to a pocosin streamhead, 
and likely influenced by proximity to tidal freshwater swamp 300 feet downstream (S. 
villosicarpa itself appears to be an indication of that influence). The upland soil at this site 
is classified as well-drained Baymeade fine sand ultisol. 

Two of the four currently known sites—in Camp Lejeune near Salliers Bay and near 
Mock-up Road—are within one mile of the ocean, and each was impacted by the hurri- 
canes of 1996 and 1998, with considerable canopy blow-down. Increased seedling es- 
tablishment and reproductive maturation was observed in 1998, suggesting a positive 
response to the hurricane impacts. At the Salliers Bay site, 300-400 flowering individuals 
and 1000+ vegetative rosettes were estimated in 1998, compared with 50 flowering and 
100 vegetative rosettes under a closed canopy in 1995. The Mock-up Road site, first dis- 
covered in 1998, had 500+ flowering/fruiting individuals. The other two sites are consid- 
erably inland, with the Frenchs Creek site in Camp Lejeune seven air miles from the coast 
and the Clarks Landing site in Pender County 16 air miles from the coast. Each site, how- 
ever, is located on low uplands or upland slopes adjacent to fresh or slightly brackish 
tidal creeks or floodplains. Due to their more inland locations, these sites were much less 
impacted by the 1996 and 1998 hurricanes. Comparative population data are available 
only for the Frenchs Creek site: 25 flowering with 75 vegetative rosettes in 1998, and 40 
flowering with 150-200 vegetative rosettes in 1992. Another observed habit of Solidago 
villosicarpa that may be associated with hurricane impacts is the tendency of plants in 
canopy openings to produce more robust inflorescences with paniculate thyrsoid 
branches, while plants in shaded areas tend to produce a simple elongate terminal thyrse. 

These variable conditions and evident adaptation to disturbance suggest that So/- 
idago villosicarpa, while likely restricted in range, may be more frequent along the Caro- 
lina coast than is currently known. 


ACKNOWLEDGMENTS 


| am indebted to Alan Weakley for his recognition of the resemblance of the Camp Lejeune 
specimens to those of the 1950 collection at NCSC, and to Guy Nesom for his review of 
this paper in draft. Material assistance was provided in the field by John Hammond and Eric 
Davis of Camp Lejeune Marine Corps Base Division of Fish and Wildlife,and by Bruce Sorrie, 
Karen Patterson,and Alan Weakley. Fieldwork that resulted in the discovery of the Onslow 
and Pender county occurrences was conducted under the auspices of the N.C. Natural 
Heritage Program. Jeff Nekola and Bruce Sorrie provided critical help in the examination 
of habitat and character states of Solidago sciaphila populations in northeastern lowa. | 
am grateful for the assistance provided by the herbarium curatorial staffs at the University 
of North Carolina—Chapel Hill (NCU) and N.C. State University (NCSC). The illustrations 
were prepared by Margret Mueller and photographs were taken by Bruce Sorrie, both 
using living specimens from the type locality on the date of the type collection. Current 
U.S. Geological Survey topographic quadrangle place names are used when available. 


300 SIDA 19(2) 


REFERENCES 


Cronauist, A. 1980. Vascular flora of the southeastern U.S. Vol. 1: Asteraceae. University of 
North Carolina Press, Chapel Hill, NC. 

FERNALD, M.L. 1950. Gray's manual of botany. 8th ed. D.Van Nostrand, New York, NY. 

Fox, W.B., R.K. Goprrey, and H.L. Blomquist. 1952. Notes on distribution of North Carolina 
plants—lll. Rhodora 54:165-182. 

Gteason, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United 
States and adjacent Canada. 3 vols. New York Botanical Garden. 

Gieason, H.A.and A. Cronauisr. 1991.Manual of the vascular plants of northeastern United 
States and adjacent Canada. New York Botanical Garden, New York, NY. 

KALLUNKI, J. 1998. Personal communication (e-mail) to RJ. LeBlond, August 13. New York 
Botanical Garden, New York, NY. 

Nesom, G.L. 1993. Taxonomic infrastructure of Solidago and Oligoneuron (Asteraceae: 
Astereae) and observations on their phylogenetic position. Phytologia 75:1-44. 

Raprorp, A.E., H.E. Ales, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill. 

ScHarAte, M.P.and A.S. Weaktey. 1990. Classification of the natural communities of North 
Carolina (Third Approximation). N.C. Natural Heritage Program, DPR, DENR, Raleigh. 


STRUCTURE OF POPULATIONS OF OTATE 
(OTATEA ACUMINATA SUBSP. AZTECORUM: POACEAE) 
IN HARVESTED STANDS 

Jose Martin Vazquez Lopez', Bruce F.Benz! 
Miquel Olvera Vargas', Sergio Graf Montero? 


'Universidad de Guadalajara, Centro Universitario de la Costa Sur 
~*~ + Ps oo BU a | Pg D Aid | 


FA * 

Apdo. Postal No. 108, Autlan de Navarro 
Jalisco, CP 48900 MEXICO 
?Department of Biology, Texas Wesleyan University 

1201 Wesleyan, Fort Worth, TX 76105, U. 


3Direccion de la Reserva de la Biosfera Si de Manantlan 
ot rs f dol NApv A bj. tr R A] | y Pesca 


Autlan, Jalisco, 48900 MEXICO 


ABSTRACT 


We describe the structure oF ce pes sae of otate (Otatea acuminata (Munro) Cald. & Sod. 
subsp. ¢ 1an, at are subject to traditional extraction. These popu- 
lations: are eatnparedl ee on pa diameter and height using growth stage eaISaOnies, site 
characteristics and history of management.The point quarter method was at to sample popula 


tions to obtain estimates of stem densities. Analysis of varianc t 
and density to compare stands; all Soller ely showed significant SMEs between cians 
Results suggest that the population structure of ot 1 by harvest intensity, which 


appears to be a function of accessibility of the stands by basket makers and Gat history of use. 
Results are described in relation to harvesting otate for craft production. 


Key Woros: Structure, otate, stem category, harvesting, artisan 


RESUMEN 


Se describe la estructura de ocho rodales naturales de otate (Otatea acuminata (Munro) Cald. & 
Sod. subsp. caacial euzinan mneye y eon) que estan sujetos a extraccidn tradicional. Se 
tallos, caracteristicas del sitio e historial de 
ance Se utiliz6 e| método punto cuadrante para estimar la densidad de tallos.Se hizo un andlisis 
de varianza de la Geneidad y categorlas cle eulge para eompala’ > rodales entre sf; todas las 
ee gerd acas 
“ Saito: indican que la estructura ‘de las poblaciones de otate estd siend a por la 
intensidad del aprovechamiento, el cual esta determinado a cee por la cercania y 
accesibilidad de los rodales a im artesanos. Los resultados se discuten en relacién al 


aprovechamient pafd fines artesanale 


Pacapras CLave: Estructura, otate, categoria de tallo, aprovechamiento, artesano 


SIDA 19(2): 301 — 310. 2000 


302 SIDA 19(2) 


INTRODUCTION 


Numerous genera and species of the Bambusoideae play an important role in forests 
because they occupy a wide diversity of habitats and exhibit extremes of morphologi- 
cal diversity (Soderstrom & Calderon 1974). The genera of bamboos in Mexico include: 
Otatea, Olmeca, Guadua, Chusquea and Olyra (Judziewicz et al. 1999). The antiquity of 
Bambusoideae species’ use in Mexico is not known with certainty though we suspect its 
most ancient inhabitants used them. The bamboo otate (Otatea spp.) has been used in 
Mexico since prehispanic times for a wide variety of purposes including house con- 
struction, walking sticks, stakes, and for basket making, among others (Torres 1985; Anaya 
1989; Benz et al. 1994; Bye 1995). Despite its long history of use and considerable study of 
its distribution and taxonomy, it is clear that information about the species’ ecology and 
management is sorely lacking. 

Otatea is a genus indigenous to Mexico and Central America, occurring in Pacific 
watersheds from Sonora to Chiapas and Central America, besides the Mexican states of 
Veracruz, Puebla, Queretaro and Mexico (Guzman et al. 1984). There is a disjunct popula- 
tion in northeastern Colombia where it is called cana brava Judziewicz et al. 1999). There 
are two species in the genus, O. fimbriata and O. acuminata, the last one with two sub- 
species, O. acuminata subsp. acuminata and O. acuminata subsp. aztecorum. These spe- 
cies are used for basket making, walk sticks, broom sticks, corral construction, canes, fur- 
niture, crop supports, and house rafters (Guzman et al. 1984; Judziewicz et al. 1999). 

This research was carried out in the ejido of Platanarillo, in the Municipality of 
Minatitlan in the Mexican state of Colima where otate (Otatea acuminata (Munro) 
Calderon & Soderstrom subsp. aztec Guzman, Anaya & Santana) is an important natu- 
ral resource. This subspecies occurs in the Mexican states of Sonora, Chihuahua, Sinaloa, 
Durango, Nayarit, Jalisco, Queretaro, Mexico, Guerrero y Puebla (Guzman et al. 1984). 

Otate forms dense thickets of erect, two to eight meter tall individual shoots whose 
apex often overarches surrounding vegetation. This species occurs mainly in gorges and 


on pronounced slopes, on thin sc stony see or aclele soils usually derived thee cal- 


careous rocks (Guzman et al.1984).Otate 5 y by rhi h.Young 
individuals are totally covered with culm leaves. Stems (culms) emerge Sail in the 
humid season and reach their maximum height within three to four months. |n Platanarillo, 
otate is used in house and corral construction but its principal utility and value is tied to 
its suitability for making handicrafts, principally baskets. There has been resurgence in 
local interest to promote large-scale extraction because of its market potential as a source 
of stakes for cultivation of tomatoes, chayotes and other vegetable crops. 

Craft production in the ejido Platanarillo is of great socioeconomic importance 
because 32 families, or about 40 percent of the residents, engage in basket manufacture 
as a principal means of generating household income. Otate harvesting for this purpose 
has been carried out over the years under a traditional management scheme, that con- 
sists of the selective cutting of young stems. This traditional form of management is of 
considerable interest because these practices could have negative impacts on the re- 


VAZQUEZ L. ET AL., STRUCTURE OF NATURAL STANDS OF OTATE 303 


source because extraction frequently occurs repeatedly in the same areas and stands. 
The objective of this work is to describe how the structure of otate populations has 
been affected by traditional extraction. 


STUDY AREA 


The Ejido of Platanarillo is located between geographic coordinates 19°21 and 19°29'N 
latitude and 103°56' and 104°00' W longitude in the state of Colima. This ejido encom- 
passes 3,028 hectares at an altitude ranging from 900 to 1800 meters above sea level. 
Nearly one-half of the ejido’s land area is located on the southwest flanks of Cerro Grande, 
a calcareous formation of sedimentary origin with 10 to 45 degree slopes. The remain- 
der of the ejido is located on the north slope of the Sierra Perote, a volcanic formation 
with 25 to 45 degree slopes. Both form part of the northern-most extent of the Sierra 
Madre del Sur. Soils on the slopes of Cerro Grande are litosols and andosols, and on the 
Sierra Perote, regosols and cambisols (INEGI-SPP 1981). 

Lithosols are generally shallow, rocky and infertile soils located in areas with pro- 
nounced slope. These soils are not particularly apt for agricultural purposes. Cambisols 
are superficial soils rich in organic material with weakly developed horizons found on 
moderate slopes. Nevertheless, cambisols are more appropriate for agriculture than litho- 
sols. Andosols form from volcanic ash. They are very light textured and have a high ca- 
pacity for retaining water and nutrients. This type of soil scarcely occurs in the study area, 
generally in areas with high slopes. Regosols are similar to cambisols in being rich in 
organic matter and having weakly developed horizons found on moderate slopes. They 
differ by regosols having almost no horizon development. Regosols support agricultural 
activities. 

The climate in the area surrounding the ejido is mild, mid-latitude humid subtropi- 
cal with dry winters and hot summers according to the modified Koeppen classification. 
Average annual precipitation is 1,350 mm. It presents a marked seasonality with a dry 
season from October to May and a wet season from June to September (Martinez et al. 
1991). The predominant vegetation types in the ejido are the tropical deciduous forest, 
tropical sub-deciduous forest,and deciduous oak forest (Vazquez et al.1995).Otate popu- 
lations are an important component of the tropical deciduous forest that occupies the 
greatest extension in the ejido. These populations of otate occur mainly in communal 
land in the ejido, but also in areas with assigned rights, where basket makers must get 
permission for extracting the resource. 


SAMPLING METHOD 


Sites dominated by otate stands were delimited with aerial photos and a satellite image 
SPOT1 HRV2;band 321,scale 1:50,000 taken on March 30th of 1987 and ground truthing 
in the ejido. Otate populations occur over 340 ha in the ejido. An inventory of the otate 
populations was conducted during the months of May, June and November of 1993 to 
document the presence and evaluate the quality of stands. 


304 SIDA 19(2) 


A total of 25 individuals with an age ranging from 18 to 70 years helped in the 
inventory and were interviewed about the levels of extraction and harvesting intensity. 
Seven populations were selected on the calcareous soils of the flanks of Cerro Grande, 
and one was selected on the volcanic soils of Sierra Perote. These varied according with 
the harvesting intensity. Site characteristics were recorded for each stand in order to 
describe basic aspects of the sites where otate grows. These populations cover about 100 
ha, almost a third of the total area with otate in the ejido. All of these populations have 
existed in the area during the last 30 years at least and remain today in spite of harvesting. 

Estimation of stand density was calculated as the number of stems per hectare 
based on the point-quarter method of Cottam and Curtis (1956). Eight stands subject to 
different extraction levels were sampled systematically. Fifteen to twenty points were 
sampled in each stand. Points were placed on linear transects distributed systematically 
every 10 meters across each stand. The distance between points was established ac- 
cording to observed spatial pattern of the stand. Estimates of density were calculated 
using the equation proposed by Cottam and Curtis (1956): D=dm where: D is equal to 
density and dm is equal to the mean distance of stems from the center point in each 
quarter. Analysis of variance with multiple post-hoc comparisons using Duncan‘s method 
was used to compare population densities. 

Culm diameter at breast height (dbh) was measured using a Caliper. Height was 
measured using marked stakes. As stems reached their maximum height, they were clas- 
sified into four growth-stage categories:new, young, adult and dead, New refers to stems 
that emerged during the last rainy season and were easily recognized by having com- 
plete spiculate culm leaves; young refers to those stems that had emerged two to three 
years ago, losing some culm leaves in the intervening time; adult stems refer to those 
with few or no culm leaves. Dead stems were easily recognized since they had neither 
foliage nor culm leaves and had brittle stems. Average age of shoot death was unknown. 
The average density of stumps (cut stems) was considered an indicator of the level of 
extraction and the average density of new stems was considered an indicator of regen- 
eration. 

Vigor was characterized in the following manner: good refers to green stems that 
are notably healthy, complete foliage; regular, yellowish stems and incomplete foliage 
with some damage; bad, refers to gray and cracked stems with irregular internodes with 
few, notably damaged leaves. 


RESULTS 


Population vigor varied from good to regular for new and young stems. Adult stems 
generally had regular vigor. Site characteristics were similar for stands located on the 
slopes of Cerro Grande while the single population on Sierra Perote was notably more 
favorable (Table 1, stand 8). 

Stand eight exhibited the greatest regeneration and the highest density of stumps 
of sites with evidence of extraction. Stand six showed no evidence of extraction and the 


VAZQUEZ L. ET AL., STRUCTURE OF NATURAL STANDS OF OTATE 305 


Tas.e 1.Site characteristics of eight otate stands in the ejido of Platanarillo. 


Stand _— Accessibility’ Area Aspect Slope in Rockiness? 
(ha) degrees 

] Easy 10 Southwest 10-40 Medium 

2 Easy 1] West 15-40 Medium 

3 Moderate 19 West 15-45 High 

4 Moderate 9 West 15-40 High 

5 Difficult 8 Southwest 20-45 High 

6 Difficult 4 West 20-35 High 

7 Difficult 29 Southwest 20-45 High 

8 Moderate 10 Northeast 25-45 Low 


‘Accessibility S distance, ease of transport and land tenure (communal or assigned rights). 
sy access (w 0 minutes walk from the community, path with moderate incline and not 
Se _ 7 Moderate access (between 30 and 60 minutes walk from the community, 
path rocky with moderate incline, assigned land rights limited to certain individuals), difficult ac- 
cess (greater than 60 minutes walk from the community, path rocky with steep incline, communal 
land) 
2Rockiness refers to abundance and proximity of rocks (sediment > 1m in diameter) (Olvera et al. 
1996). Low (absent to one rock every 20 meters), Medium (a rock every three to 10 meters), High 
(rocks predominate, soil barely visible). 


lowest level of regeneration. Sites three and four exhibited intermediate levels of extrac- 
tion and of regeneration (Table 2). Stands five and eight had the largest diameter of 
young, mature, and dead stems. Stands one, four and five had relatively large-diameter 
stems. Stands six and seven had small-diameter stems (Table 3). The tallest stems oc- 
curred in stands one and five; the shortest stems in stands two, three and seven (Table 
4).Stand seven also had mostly small, short stems while stand eight had the tallest stems 
(Table 3 and 4). 

In summary, stands number six and eight presented significant differences from 
the rest with respect to density, degree of extraction, and stem diameter. Stand eight 
exhibited the highest level of extraction, the greatest density of regeneration, and the 
largest—diameter and height—stems compared to the rest of the stands. In contrast, 
stand six exhibited no evidence of extraction, had the largest density of dead stems, 
showed the lowest density of regeneration, and had some of the smallest diameter stems 
(Tables 2-4). 

According to our interviews with local basket makers, stands one and four were 
intensively extracted in the past. Stems are extracted from stands one and four by the 
greatest number of basket makers, with greatest regularity, each collecting between 
40-60 stems per fortnight during September through November. Stands two, three and 
eight are subject to an intermediate level of extraction, at most ca. 40-60 stems per 
month.Stands five, six,and seven currently have the lowest levels of harvesting, the equiva- 


lent or less of 10 stems per month. 


Taste 2. Density (mean and standard deviation) of stems ha”! in eight stands of Otate in the ejido of Platanarillo, Municipio of Minatitlan, Colima, Mexico. 


Stands with the same letters are not significantly different at p<0.05. 


Stands 

1 2 3 5 6 7 8 

New 2309+ 2556 2582+ 1391 3140+1193 2671 +2646 261843283 1206+ 1266 966+999 313141845 
bcd bcd cd bed bcd abc ab cd 

Young 1412+1302 241142463 355642599 291142922 4810+2603 487143210 355743789 336541416 
abc abcd bcde abcde cde cde bcde bcde 

Adult 4025+6057 208942141 4550+6686 3081+3306 6480+ 3623 8227+6393 5472+6352 6993+8984 
abcde abc abcde abcd abcde cde abcde bcde 

Dead 1304+1065 4402+ 6397 52/70+4653 3106+3463 761449690 9556+9355 8000+ 9999 4144427443 
abc abcde abcde abcd bcde cde cde abcd 

Stumps 2424+ 3238 155142177 2898+ 3981 425744789 968+ 1894 a 414+688 4685+ 3987 
abcde abc abcde acde abc abcd cde 


*No stumps 


90€ 


(2)6L Vals 


Wi 3. Diameter at breast height of stems (in cm, mean and standard deviation) in eight stands of otate in the ejido of Platanarillo, Municipio of Minatitlan, 


olima, Mexico. Stands with the same letters are not significantly different at p<0.05. 


Stands 

1 2 3 4 5 6 7 8 

New 2.54056 24+0.61 2.£0.67 2.9+0.64 2.7+£0.69 24+0.64 2.00.70 2.8+0.76 
bcd bc bc de cde be a de 

Young 2.6+0.60 2.3+0.59 2.3+0.62 2.4+0.63 2.6+0.79 2.00.79 1.80.77 3.00.84 
def bcd bcd pcdef def a a g 

Adult 2.2+0.39 2.0+0.42 2.00.54 2.14048 2.3+0.63 1.8+0.64 1.4£0.60 2.8+0.78 
cd bc bed cd cd be a e 

Dead 2.3+0.47 1.94043 1.80.43 2.10.62 2.0+0.61 1.7+0.42 iesya=l Oho ys 2.6+0.76 
de abc abc cde bcd abc ab f 


Taste 4. Height (in meters, mean and standard deviation) of otate stems in eight stands in the ejido of Platanarillo, Municipio of Minatitlan, Colima, Mexico. 


Stands with the same letters are not significantly different at p<0O. 


Stands 

1 2 3 4 5 6 7 8 

New 4.09+1.14 4.23b41.2 3.90+1.14 492+ 1.03 5.50+1.64 5.08+ 1.66 3.55+1.45 452+1.55 
bcd cd abc def ef ef ab cde 

Young 3.71+0.97 2.97+0,91 3.02+0.89 3.31+£0.79 4.13+1.09 3.59 1.19 242+1.03 3.25+0.84 
cd be bec bc e bcd a bc 

Adult 2.92+0.79 2.46+0.71 2.45+0.82 2.82+0.65 3.47+0.95 2.67+0.90 2.01 +0.84 2.88+0.67 
Cc bc C Cc e bcd a G 

Dead 2.85+0.76 2.28+0.73 2.33+0.79 2./3+0.70 3.23+1.08 2./5+0.64 1.88+0.78 2.64+0.63 
c b b Cc d Cc a Cc 


JIVLO 40 SONVIS TVHNLVN JO JUNLINYLS “TW 13°71 ZINOZVA 


LOE 


308 SIDA 19(2) 


DISCUSSION 


Harvesting in the ejido is based on selection of young stems because these make better, 
more flexible, baskets. Over the years, this kind of management has produced a stand 
structure with an abundance of adult stems and appears to promote regeneration. 

Field observations indicated that stands one and four have the greatest prolifera- 
tion of woody species (Acacia spp.and Lysiloma spp.) and these stands have some of the 
lowest densities of young, adult, and dead stems. According to local informants these 
two sites were once occupied by dense stands of otate that were intensely exploited in 
the past, suggesting that sakes and intense extraction impeded regeneration of 
adult stems in these otate p tions and permitted the invasion and establishment of 
trees and shrubs. This foe harvesting of sites one and four during the past was due 
mainly to their proximity to the communities inhabited by a large number of basket 
makers. 

On the other hand, stand eight appeared to respond to high levels of extraction by 
producing a high density of new large-diameter stems. This could be due either to high 
levels of extraction reducing competition or its location on rock-free acidic soils with 
high fertility. Stand six had the lowest level of regeneration—it had the highest density 
of mature and dead stems—perhaps because the population has not been subject to 
thinning by basket makers extracting stems for basketry manufacture. These results lead 
us to hypothesize that the basket makers of Platanarillo have exceeded sustainable lev- 
els in some stands (one and four) while maintaining appropriate levels of extraction in 
others (two, three, five, seven and eight) appropriate to ecological conditions and the 
population's ability to regenerate. 

Numerous hypotheses have been offered to explain massive flowering of bam- 
boos. This phenomenon (gregarious monocarpy) might be determined by physiologi- 
cal changes caused by growing conditions, attempts to satiate seed predators, the in- 
tensity of extraction or by perturbation like wildfire or similar factors Janzen 1976;Keeley 
& Bond 1999). The flowering periodicity of these otate populations is known from local 
folklore (Santana & Lemus 1992). According to local people, the last massive flowering 
event occurred 30 to 35 years ago. Seven informants between 40 and 55 years old report 
having seen the flowering process once before, and two informants, 75 and 80 years old, 
have seen it flower twice before. 

In 1993 we observed a few culms flowering in some otate populations of the ejido, 
but not all individuals in a stand appeared to flower simultaneously. Widely separated 
individual culms flowered while nearby culms did not. Within one year's time from the 
appearance of one nonnae, individual, all individuals will have flowered. Now, in the 
year 2000, most of the poy flowered completely. Nevertheless, the flowering 
process has been sequential and prolonged, not simultaneous; some populations still 
have not flowered. By the time the last population in the ejido flowers, the first popula- 
tion to flower will have new shoots that can be extracted for basket manufacture. The 
flowering process in the ejido requires a period of about seven years to be completed. If 


VAZQUEZ L. ET AL., STRUCTURE OF NATURAL STANDS OF OTATE 309 


our observations about the flowering process are accurate, we suspect that extraction 
for basket manufacture has little or no impact on flowering process. 


CONCLUSIONS 


Continuous harvesting of young stems from stands three, five, six, seven, and eight ap- 
pears to be possible from the information we have obtained thus far. These stands had 
either a high density of regeneration (stand three) or had high densities of young: ane 
adult stems (stands five, six,and eight). Stand seven had the lowest level of reg 

50, the extraction of new stems here must be the lowest. There is an obvious need to 
reduce extraction in stands one, two, and four, all of which have been intensively ex- 
ploited in the past and exhibit densities of young and adult stems that suggests high 
regeneration rates. 

|t appears contradictory that population six located at the greatest distance from 
the largest number of basket makers possesses the poorest quality for basket manufac- 
ture, while population eight located closest to a significant number of basket makers 
exhibits the greatest intensity of extraction, hast the greatest amount of regeneration 
and possesses stems with greatest useful dimensions. We offer the working hypothesis 
that human extraction of otate stems in Platanarillo has actually fostered growth of more 
stems with better qualities. 

Otate is a very important resource for the ejido, especially for the poor people (most 
of the ejido’s inhabitants) because of its utilitarian and economic value. Commercial 
extraction poses management challenges because the intensity of extraction could be 
considerably greater than potential regeneration, while labor investment and the im- 
mediate economic benefits are significantly higher than actual costs. 


Suggestions For Management 

A controlled management program must be implemented in order to regulate harvest- 
ing practices. Basket makers agree that regulation of extraction is necessary and pro- 
pose some actions of control, like protection against fire, excluding livestock, and com- 
mercial harvesting of adult stems. Establishment of permanent research plots is also 
necessary in order to continuing monitoring and evaluating the populations’ responses 
to different harvesting treatments. These measures could lead to improved manage- 
ment by establishing cutting level as well as a systematic registration of the phenology 
of the species. This will be very important since the process of flowering is nearly com- 
plete in all populations of the ejido. The monitoring process should involve local people 
and trained investigators. 


ACKNOWLEDGMENTS 


We acknowledge the support and participation of the basket makers in the Ejido of 
Platanarillo that allowed us to complete our work. This research was supported by a 
grant from the Mexican Secretary of Environment, Natural Resources and Fisheries to 
the Directorship of the Sierra de Manantlan Biosphere Reserve. 


310 SIDA 19(2) 


REFERENCES 


Anaya C., C. 1989. Estudio de la subfamilia Bambusoideae (Poaceae), con revisién 
taxonomica para el estado de Jalisco, México. Tesis de licenciatura. Facultad de 
Agronomia. Universidad de Guadalajara. Guadalajara Jalisco, México. 

Benz, B.F., F.J. SANTANA M., MLR. Pinepa L., J.Cevattos E., L. Rostes H.,and D. De Niz L. 1994. Charac- 
terization of mestizo plant use in the Sierra de Manantlan, Jalisco-Colima, Mexico. J. 
Ethnobiol. 14:23-41. 

Bye, R. 1995. Ethnobotany of the Mexican tropical dry forests. In: Bullock, S.H., H.A. Mooney 
and E. Medina (eds.) Seasonally Dry Tropical Forests. Cambridge University Press. Cam- 
bridge. Pp. 423-438, 

Cortam, G. and J.T. Curtis. 1956. The use of distance measures in phytosociological sam- 
pling. Ecology 37:451-460. 

Guzman M.R., M.C. Anaya, and FJ. Santana M. 1984. El género Otatea (Bambusoideae), en 
México y Centroamérica. Bol. Inst. Bot., Univ. Guadalajara. 5(10):2-20. 

AGuiLerA-Herrera, N. 1989. Tratado de Edafologia de México. Tomo |. Universidad Nacional 
Autonoma de México. Facultad de Ciencias. Departamento de Biologia. Laboratorio 
de Investigaciones de Edafologia. México, D.F.222 Pp. 

INEGI-SPP. 1981. Sintesis Geografica del Estado de Colima. Coordinacién General de los 
Servicios Nacionales de Estadistica, Geografia e Informatica. SPP. México. 

JANZEN, D.H. 1976.Why bamboos wait so long to flower? Ann. Rev. Ecol. Syst. 7:347-391. 

Jupziewicz, E.J.,L.G.Ciark, X. LoNDonio, and M.J. Stern. 1999. American Bamboos. Smithsonian 
Institution Press. New York. 

KeeLey, J.E.and WJ. Bono. 1999. Mast flowering and semelparity in bamboos:The bamboo 
fire cycle hypothesis. Amer. Naturalist 154:383-391. 

Martinez R., L.M., J.J. Sanoovat, and R.D. Guevara G. 1991. Climas de la Reserva de la Bidsfera 
Sierra de Manantlan y su region de influencia. Laboratorio Natural Las Joyas. Univer- 
sidad de Guadalajara. Informe Técnico. 

OwveraV.,M., S. Moreno G. and B. Ficueroa R. 1996. Sitios permanentes para la investigacion 
silvicola. Manual para su establecimiento. Libros Técnicos del Instituto Manantlan. 
Universidad de Guadalajara. Guadalajara, Jalisco. México. 

Soperstrom, T.R. and C.E. CaLberon. 1974. Primitive forest grasses and evolution of the 
Bambusoideae. Biotropica 6:141-153. 

Torres, B. 1985. Las plantas utiles en el México antiguo segun las fuentes del siglo XVI. En: 
Rojas R., 1. y W.T. Sanders (comps.) Historia de la agricultura. Epoca prehispanica, siglo 
XVI.Coleccién Biblioteca del INAH. México, D.F. 

SANTANA M.,F.and S. Lemus J.1992.La floracién de los otates, un hallazgo sorprendente yun 
estado critico de la planta.En:Cartapacios. Suplemento cultural, Ecos de la Costa. Colima. 
Ano 8, 43(406):7-8. México. 

Vazquez G,, J.A.,R.Cuevas G., T.S. Cocurane, H.H. Iutis, FJ. SANTANA M., and L.Guzman H. 1995. Flora 
de Manantlan. Sida, Bot. Misc. 13. Botanical Research Institute of Texas, Fort Worth. 


BLOOMING “BEHAVIOR” IN FIVE SPECIES OF 
BOERHAVIA (NYCTAGINACEAE) 


Richard Spellenberg 


Department of Biology, MSC 3AF 
New Mexico State University 
Las Cruces, NM 88003-8001, U.S.A. 


ABSTRACT 


Observations are reported on anthesis, stamen and style movement, and insect visitation in five 
taxa of Boerhavia, one a pantropical perennial (B. coccinea Mill), and five that are North American 
annuals (B. intermedia M.E. Jones, B. spicata Choisy, B. torreyana S.Wats and B.wrightil A.Gray). Obser- 
vations were made in natural situations in Las Cruces, New Mexico. Individual flowers are open for 
only a portion of one day. Insect visitors were Hymenoptera, Diptera, and Coleoptera. All species 
received insect visitation; the larger-flowered species had more visitors. Autogamy is believed to 
be the usual method of reproduction, either through insect pollination or self-pollination of the flower 
as stamens contact the stigma as the flowers close. There is no evidence of wind pollination even 
in crowded Pepulslons Chromosome numbers are fairly high, especially for annuals. New chro- 

mosome number pepolise re made for B. coccinea (n = 26),B. diffusa L.(n = 27), B.intermedia (n = 26, 
ca.27),B. linearifolia (n = ca. 26), B. mathisiana F.B. Jones (n = ca. 26), B. spicata (n = 26),and B. wrightii 

(n = 27).|t is suggested that high chromosome number, prevalent autogamy, but occasional out- 
crossing or hybridization, produce a population structure of locally uniform populations that differ 
slightly to greatly from other populations, a pattern that can lead to difficulty in classification. 


— 


RESUMEN 


Se informa de las observaciones del movimiento de los estambres y el estilo durante la antesis, asi 
como de la visita de insectos en cinco taxa de Boerhavia, uno pantropical perenne (B. coccinea 
Mill), y cinco anuales norteamericanos (8. intermedia M.E. Jones, B. spicata Choisy, B. torreyana S. 
Wats.,and B. wrightil A. Srey), Las oiservaclaiies se pIEIETOR € en Sed ells naturales en Las Cruces, 
New Mexico. Las fl nsectos visitantes fueron 
Hymenoptera, Diptera, y Coleoptera. Todas las especies recibieron visitas de insectos; las especies 


de flores grandes tuvieron mas visitantes. Se cree que la autogamia es el método normal de 
oe tanto ia epromeganla como por ote ae de i flor mediante contacto de 
tigma cuand | flor se ¢ cierra. Ne aoe SUIGe ne! ia nem caine. ni siquieta 


en sobliciones Aaa | | 


Se citan B inea (n = 26),B. diffusa L.(n = 27), B.intermedia 
(n = 26, ca. 27), B. linearifolia (n = ca. 26), B. mathisiana FB. Jones (n = ca. 26), B. spicata (n = 26), y 
wrightii (n = 27).Se sugiere que el alto numero crore Oiaee, : autogalie piesornianie con 
reproduccién cruzada ocasional o eee 
localmente uniformes que difieren de ligeramente a mucho de otras poblaciones, un ‘patron que 
puede llevar a dificultades en la clasificacion. 


INTRODUCTION 


Nyctaginaceae comprise a small family of approximately 30 genera and 390 species 
(Mabberly 1997) consisting mostly of American genera, several of which are noted for 


SIDA 19(2): 311 - 323. 2000 


312 SIDA 19(2) 


their taxonomic problems. Among those genera is Boerhavia (ca. 20 species) which, in 
addition to being highly developed in North America, contains some rather difficult 
groups, for example, a pan-tropical complex of perennial forms (8. diffusa L., B. coccinea 
Mill, etc.) and at least two North American groups of annuals (8. spicata Choisy complex; 
B. erecta L.complex). As discussed by Ornduff (1969), insights into reproductive aspects 
may help to understand variation seen within and between populations and this, then, 
may be useful in taxonomic interpretations. 

In the case of Boerhavia, populations in the field often appear homogenous within, 
but differ to a greater or lesser extent with neighboring populations. This population 
structure is conspicuous in the perennial 8. coccinea where there are numerous races of 
maroon-flowered populations that differ in general robustness of plants, in nature and 
distribution of pubescence, or in number of fruits in the terminal clusters of the 
inflorescence. In addition, a few populations in this species vary markedly with respect 
to flower color and other characteristics. For example, an isolated consistently white- 
flowered population with bright green, lightly pubescent foliage occurs on a rock out- 
crop on the plains of southern New Mexico. Elsewhere in the region a yellow-flowered 
race with dull green more heavily pubescent foliage has been discovered adjacent to 
maroon-flowered less densely pubescent plants (white: New Mexico, Dofia Ana Co.,ca.3 
mi S of Cambray on Providence Cone, 25 Aug 1985, Spellenberg and Zucker 8244 [NMC, 
NY]; yellow: New Mexico, Dona Ana Co., Dofa Ana Mts.,5 slopes Summerford Mountain, 
14 Sep 1969, Spellenberg 2141 [NMC]).Within the annual species there are also a number 
of examples. Boerhavia alata S.Wats.(in the B. erecta L.complex) grows on the rocky coast 
in and near Guaymas in southern Sonora, Mexico. Without conspicuous habitat differ- 
ences it contacts and intergrades very locally with B. intermedia M.E. Jones, common on 
the hillsides in the immediate vicinity (Mexico, Sonora, Guaymas, 26 Aug 1973, Spellenberg 
and Willson 3627, 3629 [B. alata], 3630, 3631 [intermediates], 3628, 3632 [B. intermedia] [all 
at NMC, variously distributed to CIIDIR, MEXU, IBUG, RSA, NY, UC, etc J) (herbaria acronyms 
from Holmgren et al. 1990). 

Perhaps because of the curiosity of nocturnal flowering in a number of species of 
Nyctaginaceae and/or the presence of chasmogamic and cleistogamic flowers on dif- 
ferent plants within populations of a species or,commonly,on the same plant,a number 
of authors have reported on floral reproductive features and insect visitation in several 
genera. Several papers report that Nyctaginaceae have flowers that attract insects but 
often self-pollinate by anthers contacting the stigma as the flower closes (Boerhavia, 
Chaturvedi 1989; Mirabilis, section Mirabilis, Hernandez 1990) and/or through cleisto- 
gamy in plants that are also chasmogamous (Acleisanthes, Ammocodon, Selinocarpus, 
Spellenberg and Delson 1977; Cyphomeris, Mahrt and Spellenberg 1995; Mirabilis, sec- 
tion Oxybaphus, Cruden 1973). Self-incompatibility is known also in Abronia (Tillet 1967: 
Williamson & Bazeer 1997) and Mirabilis, section Quamoclidion, (Baker 1964; Pilz 1978), 

Identification of specimens of Boerhavia is often equivocal; species are variable and 
often are differentiated by minute and subjective characteristics of the fruit. Differences 


SPELLENBERG, BLOOMING “BEHAVIOR” IN BOERHAVIA 313 


in taxonomic treatments in floras during the past 50 years attests to the difficulty of 
satisfactorily circumscribing species in some groups of Boerhavia. Often in such groups 
of plants (in general and not just in Boerhavia) a combination of biological characteris- 
tics contribute to the source of the difficulties faced by the taxonomist. Here | make a 
comparison of pollination and floral action of five New World taxa, one perennial and 
four annuals, and relate these observations in a general sense to the variation seen in 
the genus. 


METHODS 


Taxa observed were Boerhavia coccinea, B. intermedia, B. spicata, a small-flowered form 
called B. torreyana (S.Wats.) Standl. (considered a synonym of B. spicata), and B. wrightii A. 
Gray. All were observed in Las Cruces, New Mexico, and are vouchered under my collec- 
tion numbers in the New Mexico State University herbarium (NMC). Among these, B. 
coccinea (7867), the only perennial in this study, is a pan-tropical species similar to B. 
diffusa, the former often considered a synonym of the latter (compare, for example, 
Whitehouse [1996], both species recognized, and Wunderlin [1998] or Diggs et al. [1999], 
one species recognized). Boerhavia spicata Choisy (7866) is an annual which, in its inclu- 
sive sense, includes several synonyms referring to phases more or less different from 
one another (e.g., B. torreyana - 7868) but linked by various intermediate forms (Reed 
1969). Boerhavia intermedia (7869), an annual of arid and semi-arid regions in southern 
North America, is sometimes considered as a variant of the widespread, weedy, B. erecta 
L. [B. erecta var. intermedia (M. E. Jones) Kearney & Peebles]. Boerhavia wrightii (7870) is 
part of a small complex of species from North and South America that are fairly distinct 
from one another. 

Boerhavia plants respond to summer rains, flowering primarily in August and Sep- 
tember in southern New Mexico.|n 1984 summer rains in the region were “good,” result- 
ing in ample late season growth for both perennials and annuals. Observations on polli- 
nation mechanisms were made daily during an eight day period (31 Aug — 7 Sep). The 
positions of stamens and stigma during the period of anthesis were observed with a 
10x hand lens. Boerhavia coccinea and B. spicata were studied in a small weedy area on 
the NMSU campus where the species were intermixed. The three other annuals were 
studied ca.5 km E of the NMSU campus in Chihuahuan Desert vegetation dominated by 
Larrea, where they were also intermixed but plants were much more sparsely distrib- 
uted. 

Periods of observation.—Plants were observed daily, with observation of each taxon 
lasting for 10 minutes. The observation periods were rotated from one species to the 
next, with a few minutes allowed in between for relocation. The first period each morn- 
ing began with a different species. Observations were made at each location (on or off 
campus) on alternate days. 

Insect visitation.—Records were kept of insect visitors, the duration of visits, insofar 
as possible where the insect next visited, and general identity. Records were kept for 


314 SIDA 19(2) 


visits to individual flower clusters (terminal spikes in B. spicata, B. torreyana, and B.wrightii, 
individual subumbellate clusters in B. coccinea and B. intermedia) and visits to entire plants 
under observation. The latter data were not corrected for number of visitors relative to 
the number of total terminal inflorescences. In addition to observations, insects were 
also collected either with a net or by aspiration, were killed, and later identified to taxon 
as precisely as possible. Number of insect visits per taxon were analyzed using JMP 3.0 
(SAS Institute, Cary, North Carolina) to determine if insect visitation differed among spe- 
cies. An a@ level of 0.1 was considered significant. 

Wind pollination.—To estimate the role of wind pollination, four glycerin-coated 
microscope slides were oriented horizontally on tops of stakes that were located within 
10 cm of inflorescences of a single plant of each species. Slides, each having a surface 
area of 12.5 cm? were approximately aligned with cardinal directions. Slides were placed 
in populations one day during the entire period from immediately prior to perianth 
opening to closure. 

Exclusion of pollinators. —Cross pollination was prevented by loosely wrapping an 
inflorescence in several layers of fine nylon stocking mesh immediately prior to anthesis. 
This effectively prevents passage of the large and spinulose Boerhavia pollen (pollen 
described in Nowicke 1970). The netting remained on the plant until seed maturation, 
typically 9-10 days. The netting prevented loss if fruits were shed prior to removal of the 
net. At the same time, other inflorescences were marked, exposed to open pollination, 
then bagged in a similar manner the following morning, after the flowers closed. Fruits 
were then collected from both treatments and opened to determine whether each con- 
tained a seed. 

Pollen / ovule ratios.—Since Boerhavia species produce one ovule per fruit, pollen / 
ovule ratios were calculated simply by counting the pollen grains produced by a flower. 
Anthers from a single flower were crushed in cotton blue and lactophenol on a micro- 
scope slide and the pollen grains were counted using a compound microscope at 100x 
magnification. Ten flowers for each species were examined. 

Hybridization.—As a generalization, breeding barriers may be weak within autoga- 
mous species beyond the barrier provided by the breeding system itself (e.g., Lewis 1963; 
Stebbins 1957), which might account for some of the variation patterns with the B. spicata 
or B. erecta complexes. Woodson and Kidd (1961) suggested that hybridization occurs 
within mixed populations of B. diffusa (perennial) and B. erecta (annual), the putative hy- 
brids representing B. coccinea in the sense of Standley (1918). To attempt to gain some 
perspective on the potential for hybridization, an interspecific cross was made in early 
September, 1984. In the mixed population on the New Mexico State University campus, 
B.coccinea was used as the pollen-receiving plant, 8B. spicata was the pollen donor.Eleven 
flowers within a single umbellate cluster were emasculated very early in the onset of 
anther dehiscence with fine forceps under a dissecting microscope. Stigmas were thor- 
oughly inspected with the dissecting microscope and the few pollen grains already on 
the stigmas were removed by “sweeping’ them off with a moistened dissecting needle. 


a 
= 


SPELLENBERG, BLOOMING “BEHAVIOR” IN BOERHAVIA 315 


The stigmas were then saturated with many grains of the donor pollen by bringing an 
inflorescence of B. spicata to B. coccinea and brushing the dehisced anthers against the 
recipient stigmas. The flower cluster was bagged in double layers of nylon mesh and 
mature fruits were collected in 10 days. 


RESULTS AND DISCUSSION 


Floral “behavior.’—In each of the species of Boerhavia an individual flower is open for 
only a portion of a day (Fig. 1). Anthesis begins near dawn, usually for all or most of the 
flowers in a single flower cluster. |n B. spicata and B. coccinea anthesis began 1-2 hours 
before it began in the other three (the perhaps warmer and more mesic campus envi- 
ronment may have influenced timing and duration of anthesis). In all, anthesis begins 
with the opening of the corolla-like perianth, stamens and styles uncoil and, in the larger- 
flowered species, the final stigma position is slightly beyond the anthers. In the smaller- 
flowered species stigma and anthers are not as well separated initially (Fig. 1). As the 
morning progresses, the filaments and style curl and the anthers haphazardly contact 
the stigma. Self-pollination may occur at this time. Perianth closure begins in late morn- 
ing and progresses rapidly, so that by mid-afternoon the perianth is crumpled, the sta- 
mens and style usually contained within. Only rarely do flowers weakly open a second 
day. These observations fully support those of Chaturvedi (1989), who reported that the 
widespread perennial, B. diffusa, is auttogamous (as known from plants studied in the 
botanic garden, Allahabad University, India). 

Insect visitation.—Insect visitors varied in kind, frequency and duration (Table 1), 
with various bees and flies frequent visitors. This is concordant with Bittrich and Kuhn's 
(1993) review that Boerhavia flowers fit the profile of bee and fly pollination. In a previ- 
ous observation, however, ants covered with pollen grains were noted entering flowers 
of B.coccinea.On such occasions, ants may also serve as pollen vectors (observation and 
comment on specimen, Mexico, Colima, 10 km SW of Tecoman, El Real, Spellenberg 2955 
NMC).The total number of insect visits varied considerably for the five taxa of Boerhavia 
studied (Table 2).Small sample size and considerable variation must be considered when 
viewing data in these tables. Raw data of total insect visit per species per day, summa- 
rized in Table 2, is not normally distributed (Shapiro-Wilk W Test, prob < W <0.0001). The 
Wilcoxon / Kruskal-Wallis Ranked Sums Test indicated significant differences in the data 
(y? = 9.196, df = 4, p = 0.0564). The Tukey-Kramer comparison of all pairs indicated that 
Boerhavia spicata differed most from all other species, significantly from from B. wrightil, 
B.intermedia and B. torreyana. 

Based on these preliminary observations, no conspicuously strong floral fidelity was 
observed by visiting insects. An insect initiating visits on one species might fly to several 
inflorescences on the same plant, then move to another plant of the same or different 
species, then return to the original plant, and so forth. Therefore, insect movement would 
allow both for autogamy, xenogamy, and hybridization. 

Wind pollination —Very little Boerhavia pollen was trapped on the glycerin-coated 


316 SIDA 19(2) 


x 
QO 
LL 
O 
of 
sae 
Oo | 
\ 
B. spicata ; 
1300 ------- 
B. intermedia B. torreyana 
1400 —— 
B. wrightii 
Fic. 1."Picture graph’ of stages of anthesis in five species of Boerhavia. Each species is diag fi lumn, anthe- 


slides. Grass and Salsola pollen was frequent. For each species the following number of 
grains were trapped upon the slides and from this data the number of grains per centi- 
meter squared per hour was calculated (number of grains: grains/hr/cm2): B. coccinea (5: 
0.025), B. intermedia (7: 0.035), B. spicata (17: 0.09), B. torreyana (0: 0), B. wrightii (8: 0.053). 
One slide from B. wrightii was removed from scoring because it had a large clump of 


SPELLENBERG, BLOOMING “BEHAVIOR” IN BOERHAVIA 317 


Taste 1.Total number of insect visitors observed in multiple observation periods for indi- 
vidual plants of each of five species of Boerhavia.N is equal to the number of ten minute 
observation periods. The average number of seconds per visit in given in parentheses. 
Small Hymenoptera visitors consisted of Bethylidae, Andrenidae (Perdita), Halictidae 
(Dialictus, Lasioglossum, Halictus). 


B.coccinea B.intermedia B.spicata B.torreyana — B.wrightii 
N=18 N=11 N=23 N=15 N=20 
Total number of visits for all observation periods ( ge duration of visit in 
seconds) 
Insect visitor 
HYMENOPTERA 
LARGE 
Scoliidae (Scolia) 14(5) 4(7) 
Mepium 
Anthophoridae 
(Mellisodes) 8(5) 5(2) 6(4) 4(0.75) 6(4) 
Chrysididae 
(Holopyga rudis 5(1) 8(0.5) 1(3) 5(0.25) 
Kimsey) 
Ichneumonidae — 1(3) 4(2) 
Sphecidae 1(60) 
(Ammophila) 
SMALL (see table 25(10) 21(5) 2(1) 1(5) 
tio 
DIPTERA 
Mepium 
Calliphoridae 2(7) 1(2) 
Syrphidae 
(Pseudodoros 2(6) 
clavatus [Fab.]) 
Tachinidae 1(1) 1(0.25) 1(5) 
MALL 
Syrphi 
(Toxomerus 6(9) 3(2) 56(7) 2(15) 5(17) 
Paragus) 
Muscidae 2(10) 
COLEOPTERA 
Nitulidae or 8(10) 
Byturidae 


pollen, presumably from the landing of a pollen-laden insect. From this data it seems 
unlikely that wind pollination is significant in Boerhavia. Pollen sexine is spinulose, and 
pollen size is rather large, consistent with that reported for these and other species by 
Nowicke (1970), and characteristic of insect pollinated plants. 

Pollination.—Pollen load on stigmas (as determined by direct observation of pol- 
len grains on stigmas with a 10x hand lens) progressively increased during anthesis, the 


318 SIDA 19(2) 


Taste 2. Estimated number of insect visits to individual flower clusters per hour in five species of 
Boerhavia.Estimations are based upon 10 minute observation periods spread throughout the pe- 
riod of anthesis. Number of periods involved in estimations and number of flower clusters indi- 
cated in parentheses. 


Hour of day 07-0800 08-0900 09-1000 10-1100 11-1200 12-1300 13-1400 Total 


B. coccinea (15 ten minute periods; 125 flower clusters) 
0.0 0.3 1.0 0.3 


1.] 0.0 3.7 

B. intermedia (14 ten minute periods; 111 flower clusters) 

0.0 0.0 0.0 0.6 0.1 1.0 17 
B. spicata (21 ten minute periods; 174 flower clusters) 

0.0 0.0 2.3 0.6 4.0 2.2 0.0 9.1 
B. torreyana (14 ten minute periods; 86 flower clusters) 

0.0 0.0 0.0 0.4 0 0.2 0.0 0.6 
B.wrightii (17 ten minute periods; 124 flower clusters) 

0.0 0.0 0.0 14 1.6 0.7 14 5.1 


majority of stigmas having more than 6 grains adhered by early afternoon (Table 3). For 
samples of B. torreyana (Table 3), the species with the smallest flowers, data indicate the 
same trend, though somewhat erratic. This species also received the fewest insect visits dur- 
ing anthesis (Table 2). Four of the five species, B. spicata the exception, still had a low to 
moderate percentage of stigmas without pollen at the time of perianth closure (Table 3). 

Even though some stigmas were unpollinated at time of flower closure, percent- 
age of filled fruits in each species was high for both open pollinated and pollinator- 
excluded inflorescences. Curling of stamens and the style places pollen on the stigma, 
with the assumption that autogamy results. In each case in the following pairs of data, 
the number of filled seeds precedes the number of unfilled seeds for plants protected 
from pollinators and for plants openly pollinated: in plants protected from pollinators — 
B. coccinea 20/0; B. intermedia 5/0; 8. spicata 8/0; B. torreyana 19/0; B. wrightii 5/0;in plants 
openly pollinated—B. coccinea 21/3; B. intermedia destroyed; B. spicata 23/0; B. torreyana 
24/1; B. wrightii 13/0. 

Pollen/ovule ratios —Pollen/ovule ratios for the five Boerhavia species examined 
ranged between 28:1 and 102:1.These figures lie between those proposed by Cruden 
(1977) for obligate and facultative autogamy (Table 4).For each taxon more than 95% of 
the pollen grains stained well in cotton blue and lactophenol, suggesting a high level of 
fertility. 

Hybridization.—Eleven mature fruits were collected from the single head of 
Boerhavia coccinea that had received pollen from B. spicata. All appeared normal. One 
was opened and the seed was normally filled. The other 10 were planted in native soil 
in pots outdoors near the end of September, 1984. Even after 16 years, none have 


SPELLENBERG, BLOOMING “BEHAVIOR” IN BOERHAVIA 319 


Taste 3. Percentage of stigmas with number of pollen grains in Boerhavia at different hours of the 
day. Number of stigmas is the total scored for the entire period. Different stigmas were counted 
during each period. 


Number of grains on stigma 
1-3 6-1 


>10 
Species, hour of day, number of stigmas scored Percent of stigmas with above 
number of pollen grains 
B. coccinea (160 stigmas) 

00-090 86 10 4 - 
0900-1100 69 16 13 2 
1100-1300 17 12 32 39 
1300-1500 Perianth closed 

B. intermedia (150 stigmas) 

00-0900 100 — _ — 
0900-1100 83 10 4 3 
1100-1300 56 19 13 iv 
1300-1500 14 12 12 62 

B. spicata (210 stigmas) 

—0900 64 12 17 7 
0900-1100 8 17 37 38 
1100-1300 ] 6 14 79 
1300-1500 = 10 10 80 

B. torreyana (150 stigmas) 
0600-0900 100 = = = 
0900-1100 66 23, 11 — 
1100-1300 32 30 18 20 
1300-1500 32 34 21 13 
B. wrightii (150 stigmas) 

600-0900 100 — = _ 
0900-1100 46 4 50 as 
1100-1300 33 7 20 40 
1300-1500 22 20 14 44 


germinated. The cross may have been too distant (perennial crossed to very different 
annual). In general, seeds in Boerhavia are difficult to germinate in high frequency (per- 
sonal observation) and dormancy may never have been broken in otherwise “normal” 
seeds. 


GENERAL DISCUSSION 


The flowers of the five different types of Boerhavia each opened for a few hours in the 
morning, and then closed permanently. Movement of stamens and style apparently as- 
sure self-pollination. These observations conform with those of Chaturvedi (1989) for 
Boerhavia. A similar mechanism was noted by Cruden (1973) and Hernandez (1990) for 
some Mirabilis, Mahrt and Spellenberg (1995) for Cyphomeris, and Spellenberg and Delson 
(1977) for Ammocodon. Allowing for the exception in the specialized Asclepiadaceae 


320 SIDA 19(2) 


Taste 4. Pollen/ovule ratios in five species of Boerhavia as related to predicted breeding systems as 
proposed by Cruden (1977, Table 1). 


Breeding system (Cruden) Mean + standard deviation, 
or species of Boerhavia pollen/ovule ratio 
Cleistogamy 28 3 

B. torreyana 51 +16 

B. wrightii 77 +18 

B. intermedia 80 +19 

B. spicata 86 +20 

B. coccinea 102 #39 
Facultative autogamy 162 +22 
Facultative xenogamy 797 +87 

enogamy 5859 +937 


(Wyatt et al. 2000), pollen/ovule ratios suggest high levels of autogamy (Cruden 1977).In 
an independent study to determine various sugar concentrations in nectar (unpublished, 
ratios provided on herbarium vouchers at NMC),| extracted nectar from all species; flowers 
of B. coccinea and B. spicata individually produce much more nectar than flowers of the 
other three; these two species had the most insect visitors, B. spicata significantly so. 
Bittrich and KUhn (1993) review that Boerhavia flowers produce nectar in the narrow 
tube, are melittophilous, and are suited for head pollination by small bees, all observa- 
tions supported by this study. Insect visitation between plants within a species would 
allow for occasional outcrossing as noted for autogamous plants by Lewis (1963). This 
would also allow for the potential of hybridization, which is supported by seed pro- 
duced (but not germinated) in a very small trial of artificial interspecific pollination. Hy- 
bridization may take place occasionally in the field, as suggested by Woodson and Kidd 
(1961).This is especially likely in closely related, little-differentiated populations, as might 
be the case between B. alata and B. intermedia in the Guaymas region in Sonora. Chro- 
mosome numbers known in Boerhavia are fairly high (n = 13, 20, 26, 27,58,see Appendix 
!), especially for annuals, which for those known n = 26 or 27.A high n number would be 
an important contributor to a high recombination index, promoting a higher number of 
new gene combinations through segregation and recombination in a limited number 
of generations (Stebbins 1951) than would a low n number. Hybridization followed by 
recombination and repetitive inbreeding would be expected to produce a rather fine- 
grained patchwork of populations homogeneous within and more or less different be- 
tween. This kind of population structure, reviewed by Lewis (1963) and Stebbins (1957) 
in discussions of the relationship between autogamy and problems of classification, 
applies in Boerhavia. As noted by Lewis, autogamy per se creates no taxonomic problem, 
and a number of species of Boerhavia seem fairly trouble free. When combined with 
outcrossing, many local phenotypically differentiated populations may be temporarily 
stabilized by autogamy. This may apply in other Boerhavia groups, such as B. coccinea 


SPELLENBERG, BLOOMING “BEHAVIOR” IN BOERHAVIA 321 


and B. spicata (both sensu lato),and such complexes may be treated in the same manner 
as those where complex patterns of variation result from outcrossing and/or hybridiza- 
tion without notable inbreeding (Lewis 1963). In such situations, the taxonomist exer- 
cises considerable personal judgment, attempting to communicate in a classification a 
useful taxonomy that corresponds to broad limits on gene flow and/or fidelity to certain 
ecological situations. 


ACKNOWLEDGMENTS 


A am grateful to Naida Zucker for critically reading the manuscript, and to James 
Zimmerman for helping with insect identification. Two anonymous reviewers provided 
valuable comments and suggestions. 


APPENDIX 


Known chromosome numbers in Boerhavia.My original counts are indicated by voucher 
citations, unless otherwise indicated, collection number is mine; voucher specimens are 
at NMC, with many widely distributed. Other counts were compiled from the literature, 
the citation provided; for counts reported in the literature, somatic numbers were con- 
verted to expected gametic numbers for ease of comparison within the following listing. 
Within a species, all are organized alphabetically by country.Original counts were obtained 
from buds fixed in cold modified Carnoy’s solution (4:3:1 - chloroform: ethanol:glacial 
acetic acid), and stained and squashed in hydrochloric acid-carmine (Snow 1963). 


Boerhavia coccinea Mill.(perennial).n = 26. Arizona, Maricopa Co., 12 mi.N of Phoenix. 2527; 
New Mexico, Dofia Ana Co, Dona Ana Mts.N of Las Cruces, 1943; Bolivia (Fernandez Casas & Fernandez 
Piqueras, 1981); Mexico, Vera Cruz, Paso de Ovejas, Pilz & Strother_683;n = ca. 26. New Mexico, Dona 
Ana Co., Las Cruces, 7867. 
Boerhavia diffusa L.(perennial).n = 13.India, (Srivistava & Misra 1966); Tanzania (Gill & Abubakar 
1975): n = 26. Hawaii (Carr 1978); n = 27. Hawaii, Oahu, SW part of island, roadside weed along 
ney H-93, s end of Waianu Range, 6396; Haleiwa at Waialu Bay, 6406; n = 58, India (Thombre 
Do), 


havia intermedia M. E. Jones (annual).n = 26. New Mexico, Doha Ana Co., Las Cruces, 
2080; Texas, Presidio Co., 24.9 mi.E of Redford, 3431;n =ca.27.New Mexico, Dona Ana Co., Las Cruces, 
7869 


se linearifolia A. Gray (perennial). = ca. 26. New Mexico, Chaves Co., 9.2 mi. NW of 
ee 343 
ee ae F.B.J (p ial).n = ca. 26. Texas, San Patricio Co.,ca.2 mi. NW of 
eee Turner 80-8 
oerhavia nae Willd. (perennial). n = 20. India (Gajapathy 1962); n = 21.India (Tandon 
and Rao 1963). 
Boerhavia spicata Choisy (annual). = 26. New Mexico, Doha Ana Co., Las Cruces, 2080; n = 
ca. 26. Mexico, Puebla, 1 mi.W of Acatlan, Pilz & Strother 671;n = 27. New Mexico, Dofia Ana Co., Las 
Cruces, 7866; 8291 [latter B. torreyana (S.Wats.) Standl. phase]. 
Boerhavia wrightii A. Gray (annual).n = 27. New Mexico, Dofa Ana Co., Las Cruces, 7870. 


REFERENCES 


Baker, H.G. 1964. Variation in style length in relation to out-breeding in Mirabilis 
(Nyctaginaceae). Evolution 18:507-509 


322 SIDA 19(2) 


Birtrich, V.and U. KUHN. 1993. Nyctaginaceae. In: Kubitzki, K., J.G. Rohwer, and V. Bittrich, vol. 
eds. The families and genera of vascular plants (ed. by K. Kubitzki). Springer-Verlag, 
Berlin. Po. 473-486. 

Carr, G.D.1978.Chromosome numbers of Hawaiian flowering plants and the significance 
of cytology in selected taxa. Amer. J. Bot. 65:236-242, 

CHATURVEDI, S.K. 1989. A new device of self pollination in Boerhavia diffusa L.(Nyctaginaceae). 
Beitr. Biol. Pflanzen 64:55-58. 

Cruben, R.W.1973.Reproductive biology of weedy and cultivated Mirabilis (Nyctaginaceae) 
Amer. J. Bot. 60:802-809. 

. 1977, Pollen-ovule rations: A conservative indicator of breeding systems in 
flowering plants. Evolution 31:32-46. 

Diccs, G.M., B.L. Lirscomp, and R.J. O’Kennon. 1999, Shinners & Mahler's illustrated flora of 
North Central Texas. Sida, Bot. Misc. no. 16. 

FERNANDEZ Casas, J. and J. FERNANDEZ PiquerAas. 1981. Estudio carioldgico de algunas plantas 
bolivianas. Anal. Jard. Bot. Madrid 38:149-151. 

GaJAPATHY, C. 1962. Chromosome numbers of some south Indian plants. Current Sci. 31: 
115-117. 

Git, L.S.and A.M. Asusakar. 1975. In: A. Love, ed. |OPB chromosome number reports XLVIII. 
Taxon 24:367-372. 

HerNAnpez, H.M. 1990. Autopolinizacion en Mirabilis longiflora L.(Nyctaginaceae). Acta Bot. 
Mex. 12:25-30. 

HOumareN, P.K., N.H. Houmaren, and L.C. Barnett. 1990. Index herbariorum, Part |: The herbaria 
of the World. Regn. Vegetabile 120:1-693. 

Lewis, H.1963.The taxonomic problem of inbreeders or how to solve any taxonomic prob- 
lem. Regnum Veg. 27:37-44. 

Masserty, DJ. 1997.The plant-book, 2" ed. Cambridge Univ. Press, Cambridge. 

Manet, M. and R. Spettenserc. 1995. Taxonomy of Cyphomeris (Nyctaginaceae) based on 
multivariate analyses of geographic variation. Sida 16:679-697. 

Nowicke, J.W. 1970. Pollen morphology in the Nyctaginaceae |. Nyctagineae (Mirabileae). 
Grana 10:79-88. 

Ornourr, R. 1969. Reproductive biology in relation to systematics. Taxon 18:121-133. 

Pi.z,G.E. 1978. Systematics of Mirabilis subgenus Quamoclidion (Nyctaginaceae). Madrono 
25511.3=132. 

Reep, C.F. 1969. Nyctaginaceae. In: C.L. Lundell, ed. Flora of Texas 2(1):151-220, 

Snow,R.1963.Alcoholic hydrochloric acid—carmine as a stain for chromosomes in squash 
preparations. Stain. Tech. 38:9-13. 

SPELLENBERG, R. and R.K. Detson. 1977. Aspects of reproduction in Chihuahuan Desert 
Nyctaginaceae, pp. 273-287 in R.H.Wauer and D.H. Riskind, eds. Transactions of the 
symposium on the biological resources of the Chihuahuan Desert region, United States 
and Mexico. U.S. Dept. Interior, National Park Service Transactions and Proceedings 
series, #3. 


SPELLENBERG, BLOOMING “BEHAVIOR” IN BOERHAVIA 323 


Srivistava, A.K. and K.C. Misra. 1966. Chromosome number in Boerhavia diffusa Linn. Sci. 
Cult,32%315:, 

STANDLEY, P.-C. 1918. Allioniaceae. North American flora 21(3):171-254. 

Stesains, G.L. 1951.Variation and evolution in plants. Columbia University Press, N.Y. 

. 1957. Self-fertilization and population variability in the higher plants. Amer. 

Naturalist 91:337-354. 

TANDON, S.L. and G.K. Rao. 1963. Meiotic studies in Boerhavia repanda Willd. Current Sci. 
32:234-235. 

THromere, M.V.1959. Chromosome numbers in some common flowering plants. Sci. Cult. 
$2:315 

Tutett, 5.S. 1967. The maritime species of Abronia (Nyctaginaceae). Brittonia 19:299-327. 

WuireHouse, C. 1996. Nyctaginaceae. In R.M. Polhill, ed. Flora of tropical East Africa. A.A. 
Balkema, Rotterdam, Brookfield 

WILLIAMSON, PS. and S.K. Bazeer. 1997. Self-incompatibility in Abronia macrocarpa 
(Nyctaginaceae). Southw. Naturalist 42:409-415. 

Woooson, R.E.and H.J.Kiop. 1961. Nyctaginaceae. In:R.E. Woodson and R.W. Schery, Flora of 
Panama, pt. lV, fasc. 4. Pp. 51-63. 

WUNDERLIN, R.P. 1998. A guide to the vascular plants of Florida. University Press of Florida. 

Wyatt, R.,S.B. Brovies, and S.R. Lipow. 2000. Pollen-ovule rations in milkweeds (Asclepiadaceae): 
an exception that probes the rule. Syst. Bot. 25:171-180. 


TAXONOMIC ADDENDUM 


This paper was written and submitted so that some of its conclusions could be referenced in an 
upcoming Flora of North America treatment of Boerhavia.The taxonomy followed was that of Reed 
(1969). As the paper was in galley and virtually “out the door” for publication, the taxonomy of the 
Boerhavia spicata group in the United States yleiere to study. This requires name changes from 


those given in the paper that may be referenced by way of voucher given in the 
body of the paper.N hang ur only for those collections originally called B. spicata and B. 
torreyana. 


Boerhavia coulteri (Hook.f. S.Wats., 7868, fourth column of Fig. 1;8297, chromosome number, n = 
26.\t has been introduced in the Las Cruces area. 


Boerhavia spicata Choisy, Pilz & Strother 671, chromosome number,n = ca. 26 (no Sica name). 
Boerhavia torreyana (S.Wats.) Standl., 7866, middle col f Fig. 1 2080, 7866, ct SOr 
ber,n = 


In tables 1,2,3 and 4 Boerhavia spicata now becomes B. torreyana, and B. torreyana now 
becomes B.coulteri. 


324 SIDA 19(2) 


BOOK REVIEW 


Detena Tutt. 1999. Edible and Useful Plants of Texas and the Southwest. (ISBN 0-292- 
78164-4, pbk.). University of Texas Press, RO. Box 7819, Austin, TX 78713-7819, U.S.A. 
(512-471-4032, 512-320-0668 fax; orders: 800-252-3206 or 800-687-6046 fax; 
www.utexas.edu/utpress/). $24.95 pbk, xxiv + 518 pp. Line drawings, 57 color photos. 


Edible and Useful Plants of Texas and the Southwest is just as its subtitle says: a practical guide. In its 
second edition (first in paperback) this book, by Delena Tull, offers information on plant character- 
istics, habitat, and range in Texas, to be used with a supplemental and more detailed flora of Texas. 
Despite the preface in which the author goes on about the joys of her new home, Alaska, the book 
is undoubtedly written by a lover of Texas biota. Subsequently, other lovers of the Texas outdoors 
and its biological diversity will find it valuable and enjoyable 

aks down in seven major parts followed by a glossary, bibliography, and index. 
Part One, titled “Edible and Useful Wild Plants” discusses native plants and tips on grazing from 
wild plants. What follows is a list of over 30 plant families who have species that are edible and/or 
useful. This is roughly a third of the book and offers encyclopedic reference to native Texas plants 
and their uses, Part Two, “Teas and Spices" covers 40 plants used as such. Part Three, “Edible and 
Poisonous Berries and Other Fleshy Fruits” is ey valuable for the camper or naturalists who 
wants to distinguish a poisonous fruit from an edible one. Part Four, "Poisonous and Harmful Plants” 
discusses similar issues and covers first aid methods for toxic plant ingestion, dispels some myths 
about poisonous plants, livestock poisoning, plant toxins, and then lists toxic wild plants by family. 
Part Five, “Colorful Dyes with Texas Plants” is definitely a strong point of this book. First discussing 
vegetable dyes historically and dyes today, and going on to discuss fibers for dyeing, factors that 
influence dye colors, dyeing techniques and dye recipes. Part Six, “Fibers from Texas Plants” covers 
plants for basketmaking, textile fibers, and papermaking. Part Seven, “Rubber, Wax, Oil, and Soap: 
Industrial eer from Texas Plants" closes the boo 

This would be a great book for any Texas cavern: level economic botany class. This is a 
great ae to take out in the field to identify useful and edible plants and taking them back to a lab 
to do a number of projects, whether, papermaking, basket weaving, or dyeing textiles. This book, 
along with Simpson and Ogorzaly’s Economic Botany, Murphey’s Indian Uses of Native Plants, and 
perhaps a selected ethnography of local group, would be suitable for an Economic Botany class 
that could be cross listed for both anthropology and botany credit.With 50 plus color plates and 
twice as many black and white illustrations this edition is field ready and easily accessible. 

While the possibilities of using this book in the academic level are exciting, it should find 
wide readership among naturalists and/or Texas lovers. As a field guide for identification and an 
encyclopedic reference of edible and economic uses of Texas plants, this book is must have for 
anyone waiting in anticipation for the spring and summer camping and hiking seasons.—Kevin D. 
Janni, Botanical Research Institute of Texas, kianni@brit.org 


SIDA 19(2): 324. 2000 


CYPERUS SANGUINOLENTUS (CYPERACEAE) NEW TO THE 
SOUTHEASTERN UNITED STATES, AND ITS RELATION TO 
THE SUPPOSED ENDEMIC CYPERUS LOUISIANENSIS 


Richard Carter 
Herbarium, Biology Department 
Valdosta State Waneniey 
Valdosta, GA 31698-0015, U.S.A. 


Charles T. Bryson 


Southern Weed Science Research Unit 
USDA, ARS, Jamie Whitten Delta States Research Center 


PO. Box 350 
Stoneville MS 38776, U.S.A. 
ABSTRACT 


ae sudie® show eee lOUane nth is aaa me) and weedy in eastern HOuIslane and 
Georgia. Morphom 

studies show C. sanguinolentus and ‘a louisianensis are paride se ae and C. 

louisianensis is treated as a synonym of C. sanguinolentus. The widespread weed, C. sanguinolentus, 

formerly thought to be restricted to the Eastern Hemisphere, is reported new to North America, 

and data on its frequency, distribution, and ecology in the southeastern United States are 

presented. 


RESUMEN 


/ j | la hierha 


Los estudios de campo muestran que 
en el este de HOUslate . el sur de Mp ae pf que se extiende hasta Alabama y Gears Estudios 

q sanguinolentus y C.louisianensis son taxondmicamente 
indistinguibles, y C. louisianensis se trata como un sindnimo de C. sanguinolentus. La mala hierba 
extendida, C. sanguinolentus, que pas se wee restringida al hemisferio este, se cita aqui 
como nueva para Norte América, y se presentan datos de su frecuencia, distribucion y ecologia en 
el sureste de los Estados. 


INTRODUCTION 


Cyperus sanguinolentus Vahl is widely distributed in the Eastern Hemisphere, where it 
has been cited as a weed (Holm et al. 1991; Mingyuan & Dehu 1970; Reed 1977; Kuhn 
1982). It is known from northeastern Africa, the Middle East, India, Sri Lanka, central Asia, 
southeastern Asia, China, Taiwan, Japan, Korea, the Philippines, Indonesia, Malaysia, and 
Australia (Clarke 1894; Holm et al. 1991; Kukenthal 1935-1936; Ohwi 1965; Mingyuan & 
Dehu 1970; Kern 1974; Reed 1977; KUhn 1982; Haines & Lye 1983;Wilson 1993) but has 
not been previously reported from the Western Hemisphere. 


SIDA 19(2): 325 - 343. 2000 


326 SIDA 19(2) 


Cyperus sanguinolentus is highly variable. Kukenthal (1935-1936) segregated five 
varieties and named seven forms, including six under the typical variety. Kukenthal’s 
(1935-1936) infraspecific taxonomy of C. sanguinolentus is difficult to use, since he pro- 
vided neither keys nor parallel descriptions of the taxa. Kern (1974) treated four subspe- 
cies, including the typical one, for Malaysia, and others (e.g., Ohwi 1965; Haines & Lye 
1977) have treated additional infraspecific taxa. Table 1 compares infraspecific taxono- 
mies of Kukenthal (1935-1936) and Kern (1974). Further research on this widespread 
and variable species throughout its range is needed for a more complete understand- 
ing of its infraspecific variation; however, such is beyond the scope of our study to deter- 
mine the range, distribution, taxonomic relationships, and status of C. sanguinolentus in 
North America. 

Its bifid style and lenticular achene with achene angle adjacent to rachilla clearly 
place C. sanguinolentus into subgenus Pycreus. Clarke (1894, 1908) segregated Pycreus as 
a genus and treated the taxon as Pycreus sanguinolentus Nees in subgenus Reticulatae 
section Vestitae. Kukenthal (1935-1936) adopted a broader definition of Cyperus, incor- 
porating this taxon into subgenus Pycreus section Sulcati of that genus. The floral scales 
of C. sanguinolentus are distinctive, being characterized by lateral grooves (or sulci) typi- 
cal of section Sulcati and, as its specific epithet implies, blood-red floral scale pigmenta- 
tion. Although some recent authors (e.g., Koyama 1985; Goetghebeur 1986, 1989: Adams 
1994; Bruhl 1995) fragment Cyperus and segregate Pycreus at the rank of genus, we have 
followed the more conservative generic taxonomy of Kikenthal (1935-1936) and 
Corcoran (1941), which, with some modification, is still widely used (e.g., Kern 1974; Haines 
& Lye 1977; Tucker 1983, 1987, 1994). 

In 1977, Thieret described a new species, C. louisianensis, from specimens he col- 
lected at two close sites in Tangipahoa Parish, Louisiana. Thieret (1977) placed C. 
louisianensis in subgenus Pycreus, noted its similarity with C. sanguinolentus, and pro- 
vided several contrasting characteristics separating it from C. sanguinolentus (Table 2) 
and a dichotomous key distinguishing it from related North American species in subge- 
nus Pycreus. Cyperus louisianensis was listed by the Department of Interior, United States 
Fish & Wildlife Service, in category two among endangered or threatened species (Anony- 
mous 1993). Until Bryson and Carter (1994) showed it was widespread and weedy in 
southern Mississippi, C. louisianensis was thought to be a narrow endemic restricted to 
two sites in southeastern Louisiana. 

In 1993, the first author was contracted by the United States Fish & Wildlife Service 
to prepare a status survey on C. /ouisianensis, which provided the initial financial support 
for this study. The major objectives of the survey were to seek additional populations of 
C.louisianensis and additional collections in herbaria, to review its status as a potentially 
rare plant, and to examine its taxonomic relationship with the Old World weed C 
sanguinolentus.|n this report, we provide a complete record of our Hel and herbarium 
investigations into the distribution, ecology and taxonomic rel 
with C. sanguinolentus. 


of C. louisianensis 


Taste 1. Infraspecific taxonomy of Cyperus sanguinolentus. 


Kiikenthal (1935-1936) - worldwide treatment Kern (1974) - Flora Malesiana 
C. sanguinolentus Vahl C. sanguinolentus Vahl 
var. sanguinolentus ssp. sanguinolentus 
f. rubro- eee (Schrenk) KUk. not treated 
f. neurotropis (Steud.) KUk. not treated 
f. flaccidulus (Boeck.) KUk. not treated 
f.cyrtostachys (Miq.) KUk. ai eer: (Miq.) Kern 
f. et a (Miq.) Kuk. ssp. melanocephalus (Miq) Kern 
f, humilis Kuk. [=ssp. ence rats 
var. micronux (C.B. Clarke) Ktk. not tr 
var. teysmannii (Boeck.) Kuk ssp. teysmannii (Boeck.) Kern 
var. korshinskii (Meinsh.) Kuk n 
var. pratorum (Korotky) KUk not treated 
var. areolatus (R. Br.) Kuk not treated 
f. setaceus Ktik. not treated 


Taste 2. Comparison of Cyperus louisianensis and C. sanguinolentus (fide Thieret 1977). 


C. louisianensis C. sanguinolentus 

Achenes elliptic to elliptic-obovate Achenes orbicular-obovate 

Achenes ge flattened (thickness-length Achenes turgid (thickness-length ratio: 0.40) 
ratio: 0.25-0.30) 

Floral scales well imbricated and flat Floral scales often barely imbricate, at least 
along the margin proximally, permitting some see through 


frequently involute along the margin 


MATERIALS AND METHODS 


Field studies —Systematic intensive field surveys for C.louisi is populations in south- 
eastern Louisiana and southern Mississippi were made by the authors during the peri- 
ods 15-21 September 1993 and 14-18 October 1993. Subsequently, the authors have 
continued to search sporadically for this taxon when time and circumstances have al- 
lowed. Thieret’s (1977) published account and more recent collections from the type 
locality with additional documentation, kindly provided by Nelwyn Gilmore, Louisiana 
Natural Heritage Program, were used to relocate the holotype locality in Tangipahoa 
Parish, Louisiana. Attempts to re-locate Thieret's paratype Gea Ele unsuccessful, which 
is not surprising since habitat in the vicinity of this site was lly altered by high- 
way and commercial construction activities. Observations at the holotype locality showed 
the species to be locally abundant along the margin of a shallow artificial pond and 
nearby ditches in the flatwoods, habitat greatly altered by humans. Searches of potential 


328 SIDA 19(2) 


habitat began outward from the holotype locality. Habitat descriptions, estimates of 
population size, and voucher specimens were made when populations were found. 

Greenhouse studies. —Transplants and plants of C. /ouisianensis started from seeds 
were maintained under controlled conditions in a greenhouse at the United States De- 
partment of Agriculture, Agricultural Research Service, Jamie Whitten Delta States Re- 
search Center, at Stoneville, Mississippi. Observations on these plants by the second au- 
thor were made in order to understand better the life history and phenology of C. 
louisianensis, especially to determine whether it is annual or perennial. 

In greenhouse experiments, seeds of C. louisianensis were sewn in flats on top of a6 
cm- dee mixture of a Basket silt a soil (Mollic Hapludaf) and sphagnum (50% v/v) in 
the first week of March, June, September, and December in 1994, 1995, and 1996. Trays 
were watered from beneath to prevent seed and soil disturbance. Individual C. 
louisianensis seedlings (> 5 cm tall) were transplanted into 15 cm-diameter pots and 
watered from beneath. The greenhouse was maintained at 30 to 35°C day and 25 to 30° 
C night, at 60 to 75% relative humidity, and without supplemental lighting. In 1994 and 
1995, flowering and fruiting plants of C. louisianensis were transplanted from several sites 
in Hancock County, Mississippi, and St. Tammany Parish, Louisiana, into 15 cm-diameter 
pots and maintained under the same greenhouse conditions as plants grown from seeds. 
Data were taken on date of seedling emergence, date of flowering and fruiting, and 
plant longevity. 

Herbarium and morphometric studies —Specimens of C. louisianensis and C. 
sanguinolentus were borrowed from selected herbaria (BRIT,GH, MO, NY and US) in order 
to examine the variation in C. louisianensis and its taxonomic relationship with C. 
sanguinolentus. Quantitative data on the achene characteristics used by Thieret (1977) 
to distinguish C. louisianensis from C. sanguinolentus were taken from these specimens 
and from our own collections. Five achenes from each of 13 specimens of C. louisianensis 
and 20 specimens of C. sanguinolentus were examined. A Bausch & Lomb stereozoom 
(6X-30X) dissecting microscope with ocular micrometer was used to measure various 
achene dimensions (Table 3, Fig. 1), and ratios (Table 3) derived from these measure- 
ments were used to analyze differences in the plane shapes of achenes used by Thieret 
(Table 2) to distinguish C.louisianensis and C. sanguinolentus. Minitab release 11.21 (Anony- 
mous 1996) was used to analyze these data statistically and to construct scatter dia- 
grams for comparing variation in C. louisianensis and C. sanguinolentus. Representative 
spikelets from selected specimens were photographed using an Olympus™ SZ40 
stereozoom dissecting microscope equipped with a Kodak™ DC120 zoom digital cam- 
era. Our more numerous recent collections were used with Thieret’s (1977) original de- 
scription and type materials to prepare a revised description and an illustration. 


= 


RESULTS 


Distribution and ecology of Cyperus louisianensis.—Since 1993, our field surveys have 
revealed numerous populations (>40) of C. louisianensis in southeastern Louisiana and 


329 


90 degrees 
ACHL 
frontal view lateral view 
Fic. 1. Frontal and lateral aspects of achene showing dimens - paring Caieislousianeadvand 
sanguinolentus. 


| > 


Taste 3.Quantitative characters and alysis of specimens of Cyperus louisianensis 
and C. sanguinolentus. Ratios defining plane shapes pee from Stearn (1992). 


ACHL Achene length (mm) 

ACHW Achene width (mm) 

ACHMX Achene distance from base to widest point (mm) 
Achene thickness (mm) 

ACHTH/ACHL Ratio used by Thieret (1977 

ACHL/ACHW Ratio approximates achene outline, e.g. elliptic (2:1), ovate & obovate (3:2), 
orbicular (1:1). 

ACHL/ACHMX Ratio approximates achene outline, e.g., elliptic & orbicular (2:1), ovate (<2:1), 
obovate (>2:1). 


southern Mississippi and isolated populations in southern Alabama and southeastern 
Georgia. Cyperus louisianensis appears to be most abundant in Hancock County, Missis- 
sippi, and adjacent St. Tammany Parish, Louisiana. Field observations show that C. 
louisianensis is weedy, often locally abundant, and restricted to periodically disturbed 
habitats, especially those maintained by mowing, such as ditches and edges of artificial 
ponds, where it appears to be aggressive and frequently forms dense stands often in 
association with other introduced and native weeds (Table 4). 


330 SIDA 19(2) 


Greenhouse observations.—In greenhouse experiments, C. louisianensis plants 
emerged from mid- and late May until mid-September each year; plants flowered from 
late August until mid-December, with peak flowering in early September to early Octo- 
ber; and plants subsequently fruited. Cyperus louisianensis plants emerging later in the 
year were shorter at flower initiation than those emerging earlier, suggesting that C. 
louisianensis is photoperiodic. Most of our field collections were made from September 
16 through December 9 (see specimen citations for the U.S.A), with peak fruiting ob- 
served and mature plants collected in mid-October; however, a few fruiting plants were 
observed and/or collected in April and May following a very mild winter (1993-1994). If 
indeed C. louisianensis is photoperiodic, this may account for the few collections of it 
from May until mid-September by us and by other botanists. All C. louisianensis plants 
transferred into the greenhouse from the field and those grown from seeds in green- 
house experiments died shortly after fruiting, showed no evidence of perennation and, 
thus, consistently exhibited an annual habit. 

Relationship with Cyperus sanguinolentus.—Thieret (1977) was limited by a small 
number of specimens collected from only two sites located less than 10 miles apart. Our 
larger sample from a wider geographical area shows that C. louisianensis is more variable 
than previously thought in the diagnostic achene characters used by Thieret (1 977) to 
separate it from C. sanguinolentus. Figure 2 shows some of the spikelet variation ob- 
served in herbarium specimens of C. sanguinolentus and C. louisianensis. Of the numer- 
ous specimens of C. sanguinolentus we have examined from throughout the Old World, 
certain ones Het Japan [e.g., M. Furuse s.n., 30 Sep 1959 (GH); M. Furuse s.n., 11 Oct 1960 
(GH); Hutoh 11517 (NY); Okamoto NSM 584 (BRIT, NY)] are indistinguishable from C. 
louisianensis in een and in spikelet (Fig.2) and achene characteristics used by Thieret 
(1977).Our field and herbarium observations show considerable variation in the degree 
of development of pigmentation and sulcus in floral scales. At least some of the variation 
in floral scale pigmentation appears related to photoperiod and temperature. Field ob- 
servations in the southeastern United States show that in mid-September the floral scales 
are typically pale with only faint pigmentation, and by mid-October they are deeply 
pigmented (cf, Figs. 2A and 2B). Opposite sides of an individual spikelet may also vary 
greatly in pigmentation os noe 26 ane nee ek ec ae (Figs. 3-6) show C. 
louisianensis and C ble with regard to the critical achene 
characteristics used By Thieret (1977) to ee them. 


DISCUSSION 


Cyperus louisianensis is widely distributed in the coastal plain of the southeastern United 
States and is often locally abundant in habitats subject to periodic artificial disturbance 
and maintenance. Distribution, frequency, and habitat of C louisianensis are indicative of 
a semi-agaressive weed, not of a narrow endemic species. Furthermore, it appears that 
the range of this taxon is apparently expanding in the southeastern United States and 
that its dispersal has possibly resulted in part from road construction and maintenance 


331 


Taste 4. Composite list of taxa associated with Cyperus louisianensis in the southeastern United States. 


Acer rubrum L. 

Acmella opposi itifolia as ) Jansen 
UTE virginic 

Anellena: nudiflora (L (L ) Brenan 

Aster tenuifolius L. 

Axonopus fissifolius (Raddi) Kuhlm. 

Bacopa caroliniana (Walt.) Robins. 

ica ( 


ck. 
Centella asiatica (L.) Urb 
oe cane (L .) Schott 


ae compressus L. 
| 


CG. ee fan 
C. elegans L 

C. esculentus L. 

C. flicinus Vahl 

C. flavescens L 

C. flavicomus Michx. 


C. polystachyos Rottb. 
C. pseudovegetus Steud. 
C. retrorsus Cha 


a0 
ge 
a 


Erigeron vernus (L. 


acq.) Macbr. 


Fimbristylis annua (All) R.& S. 
F.autumnalis (LJ R&S. 


F. tomentosa Vahl 
Conocli lest (L.) DC. 
Fuirena breviseta (Cov.) Cov. 
Gratiola sp. 

Hedyotis uniflora (L.) Lam. 
Hydrocotyle umbellata L. 


Ipomoea s 
iva annua L. 
Jacquemontia tamnifolia (L.) Griseb. 
eUies spp. 
llinga bi a Rottb. 
K. bdoraia Van 
K. pumila 
Leersia neon Swartz 
Lilaeo 
Lippia nodior L Greene 
Ludwig! tovalvis (Jacq.) Raven 
ii pens R. Forst. 
Lycopus 


eer virginiana ie 
Mikania scandens (L.) Willd. 
Mitreola sessilifolia (amel.) G.Don 
Myrica cerifera 
Oxypolis filiformis (Walter) Britt. 
Panicum repens 

Paspalum notatum Fluggé 
P.urvillei Steud. 
Phyllanthus urinaria L. 
Polygonum hydropiperoides Michx. 
Polypremum procumbens 
Rhynchospora St et (Lam) Gray 
See indica (L.) Cha 
Sagittaria sp. 
Salix ae Marshall 
Scleria reticularis Michx. 
Setaria s a 


Solidag 

So ane epee (L.) Pers. 

Piseee indicus (L.) R. Br. 

Stenotaphrum secundatum (Walt.) Kuntze 
ie platyphylla (Munro) Webst 


332 SIDA 19(2) 


Fic. 2. Spikel iation i inol C. louisianensis.—A.U.S.A., 16 September 1993, Carter 11342.-B. 
U.S.A., 18 October 1993, Carer 11502 -C.U. . A., vale’ 11579. =D sel 11 La Ms es —E. Japan, Hotoh 
517.-F. China Tsang 2 apan cale bar= 1mm 


06 | . i | 
= ze ot sang 
e e @ m 7 | 
z 05 — e ee * ° e 
rs) g° ¢ sar ° 
<< "er 33 ° 
at e had 2 % ro 7 
Tr 04 2 go eo 8 Ma 
28 0 8 oOo i] ) 
- O = O a rer) on % bd ~ 
= O . e 8 2 oe 0° a e fe) 
9 DG : | 
0.3 : e% 28 | 
| . 
Vile rn | | | —————— 
1:05 415. 1.25: 35) 45enroo: 1.60: 13/5 1,05° 1.95 
ACHL / ACHW 
Fic.3.Two g g 4 9 
A+ 2 -_ T. ] 
1.95 = eA 
oo & 
1.85 — e e e 
| e e o. 
1S c 
= e ° ox ° e 
r 1.65 | s . 2 . s 
O 1.55 ste oe : 
< 0 5 OOM a0 cen 
~ : oS 8 oo eee bs ee e | 
> 1.45 | v : » - P ‘'~ Bors 7 . | 
a e 
©O 40 5 @ Ul - ~ ee e 4 ; e | 
e # O 
- lize oP ee “#3 
1.15 are - is (> louis | 
. e | 
1.05 : Ta pe ene 
| eee —T 
15 20 25 3.0 
ACHL / ACHMX 
Fic. 4. Two-di g g Cy guinol 
fining ach keyed in Table 2. 


SIDA 19(2) 


334 
06 . _ © louis 
e san 
e e = : g 
®e e 
e e e 
‘nied 
7 Ua es ° r 
O got = 4 . 2,9 ® 
< e a a *% . se 
I 04 a se A 
F- . oO "a8 0° ),@ : e Qo ® 
rc a e .° S @9 oe ao = 
om) ow cn) e 
7 e, FO C 00 Oe ey © e 
03 oe e | 
. O . “0 e . 
e Co 
e 
ve - ime il —T - 
1 2.0 2.0 3.0 
ACHL / ACHMX 
Fic.5. Two-di liag I grel I G 9 1C 
g dA + HH A Be ode g t t L I y T.1 ») 
_ = | o louis | 
—  |{ + sang | 
” | L Se | 
ACHTH / ACHL 04- | 
= 3.0 
25 
ACHL / ACHMX 


1 C. louisianensis 


335 


activities, as suggested for C. entrerianus Boeck. (Carter 1990). At Kings Bay Submarine 
Base, where extensive populations were found in Camden County, Georgia, road rights- 
of-way are maintained by work crews traveling from Alabama under contractual service 
agreements with the Department of Defense (pers. comm., R. Wilkerson). Thus, it is pos- 
sible that achenes of C. louisianensis were accidentally dispersed into Georgia from Ala- 
bama, or elsewhere, with the transport of mowing equipment. 

Thieret (1977) distinguished C. louisianensis from C. sanguinolentus based on its more 
overlapping floral scales and its more elliptical and more flattened achenes (Table 2). 
Our results indicate the New World populations called C. /ouisianensis are encompassed 
within the total range of variation exhibited by the more variable widespread Old World 
weed, C. sanguinolentus, and are most similar to certain specimens from Japan. Further- 
more, as shown in Figures 3-6, the United States specimens are less variable than those 
from the Old World, which is consistent with the “founder principle” (Mayr 1942; Davis & 
Heywood 1973) and would be expected in a case of accidental long-distance dispersal. 
Although typification of the plethora of synonyms and accepted infraspecific names 
under C. sanguinolentus (Table 1) is beyond the scope of this study, it appears the United 
States specimens are closest to C. sanguinolentus var.sanguinolentus (fide Kukenthal 1935- 
1936) or C. sanguinolentus ssp. sanguinolentus (fide Kern 1974). 

The presence of C. sanguinolentus in the southeastern United States is not unex- 
pected for the following reasons. (1) It has been cited as an agricultural weed in the 
Eastern Hemisphere (Mingyuan & Dehu 1970; Kern 1974; Kuhn 1982;Holm et al. 1991). (2) 
Reed (1977) listed it among foreign weeds posing “potential problems in the United 
States.” (3) There are numerous other examples of weedy Cyperus spp.and other sedges 
in the southeastern United States introduced from Asia or elsewhere (Carter 1990; Bryson 
& Carter 1992; Bryson & Carter 1994; Carter et al. 1996; Carter & Bryson 1996; Bryson et al. 
1996; Bryson et al. 1997; McKenzie et al. 1998). (4) Kral (1971) reported Fimbristylis spp. 
(Cyperaceae), common in current and former rice-growing areas of the United States, 
that were likely introduced from Asia with rice (Oryza sativa L.) agriculture. (5) Histori- 
cally, rice was grown in Hancock County, Mississippi, where C. louisianensis is most abun- 
dant (Anonymous 1959, 1982). 


CONCLUSIONS 


Cyperus louisianensis is much more widespread than previously thought (Thieret 1977; 
Bryson & Carter 1994). It is locally common in southern Mississippi and southeastern 
Louisiana, and satellite populations have been found in southern Alabama and south- 
eastern Georgia. |ts habitat and frequency are characteristic of a weed, and its range and 
frequency are likely to increase, particularly in the outer Coastal Plain of the southeast- 
ern United States. 

Specimens of C. louisianensis from the United States are morphologically indistin- 
guishable from certain Old World specimens of C. sanguinolentus. Thus, we think the two 
are conspecific and treat C. louisianensis as a synonym of C. sanguinolentus. Herein, we 


336 SIDA 19(2) 


Fic. 7. Cyperus sanguinolentus Vahl.—A. Habit, pee otplant weenan paemrnennien. -B. Inflorescences (lett Bryson 
14610 & MacDonald; right, Bryson 13276).-C sty 13276).-D 
—E. Spikelet (Bryson 13276).—F. Stem section (Bryson 13276). 


= 


Floral scale (Bryson 13276). 


report C. sanguinolentus (including C. louisianensis) as new to the United States and North 
America and provide revised synonymy, description, illustration (Fig. 7), distribution map 
(Fig. 8), and specimen citations. Additional duplicates collected by the authors will be 
distributed later. 


lImitad Ctat. 


Fic. 8. Distributi by 


DICHOTOMOUS KEY TO CYPERUS SANGUINOLENTUS 
AND RELATED NORTH AMERICAN SPECIES 


1a. Floral scales membranous, bilaterally sulcate (with two narrowly elliptic translu- 
t lateral ); floral scales usually at least marginally suffused with reddish 


~~ 


livided base; style branches conspicuously exserted 

beyond floral scale, eeseced Shige about as long as floral scale; plants 

cespitose; plants of northeastern and upper mid-western United States. C. diandrus 
Torr. 


pigmentation 
2a. Stamen 


2b. Stamens 3; style pices aa than ol its enon eye een BOE sO ele 


spicuously 
decumbent, appearing stoloniferous by development of series of lateral 
branches from lower nodes; plants of southeastern United States. C. sanguinolentus 
ahl 
Floral scales fi s, not laterally sulcate; floral scales usually brown- 


“ish, blackish or reddish with Sareea more-or- ne cay distributed. 


338 SIDA 19(2) 


33. a] 1] ne +,,/ ere eee | | K If its length; 
style branches eo ecuaie: “bracts divine to reflexed; plants rhizoma- 
tous; plants ranging from southwestern United States southward oes 


sen lies America. eae! Ruiz & Pav. 
ually with 1 or more peduncul tyle divided than 

“half its its ere style PIADCHes eonapicuous mostly pro aout beyond floral 

scale; bracts mostly divaricate ually in spread- 

ing cas Sane widespread in United States, C. bipartitus Torr. 


ae pede ete al Enum. PI.2:351.1805.Pycreus sanguinolentus (Vahl) Nees, Linnaea 
.1835. Type: INDIA, Uttar PrabesH: NW Himalaya, Distr. Tehri-Garhwal, 3000 ft, Oct 1894, Gamble 
7 (L) [typ. cons. prop., Kukkonen 1995]. 

Cyperus louisianensis J W.Thieret, Proc. Louisiana Acad. Sci. 40:23. 1977. Tyee: U.S.A. Louisi- 
ANA. Tangipahoa Parish:ca. 7 mi E of Ponchatoula, along road to Lee's Landing, 1 Oct 
1972, Thieret 33585 (Holotype: GH!; lsotypes: DUKE, KNK, LAF, NC, OS). Paratypes: U.S.A. 
Louisiana. Tangipahoa Parish:junction of Ponchatoula-Madisonville highway and road 
to Lee’s Landing, ca. 7 mi E of Ponchatoula, 18 Oct 1970, Thieret 32609 (DUKE, GH!, 
KNK). 


Annual herb, appearing stoloniferous by repeated development of decumbent vegeta- 
tive lateral branches from lower nodes. Stems (6.5—)12—38(-60) cm long, 0.7-2.0 mm 
wide, trigonous. Leaves (1—)3-7; bases sheathing; blades linear, (3-)5-11(-16.5) cm long, 
(1-)2-3.5 mm wide. Primary inflorescence bracts 2—3(-4), linear; longest (1-)3-12(-19) 
cm long, 1-3 mm wide. Inflorescence terminal, usually appearing capitate, or with 1-3 
pedunculate rays to 4 cm long. Spikelets narrowly ovate-elliptic, 4-10(-15) mm long, 
2.2-3.0 mm wide, with 8-32 floral scales. Floral scales mostly closely imbricate, membra- 
nous, ovate, 1.8-2.7 mm long, carinate; keel green, 3-5 nerved; sides variable in color, 
usually variegated whitish, reddish brown to sanguineus, each with a narrowly elliptic 
translucent sulcus devoid of pigment; reddish pigment and sulcus generally becoming 
more conspicuous late season. Stamens 3; anthers 0.3-0.6 mm long. Style bifid one-third 
to one-half its length; stigmas exserted. Achene lenticular, biconvex, 1.0-1.4 mm long, 
0.6-0.9 mm wide, 0.3-0.5 mm thick, elliptic to obovate, usually asymmetrical near apex 
along side adjacent to rachilla; surface reticulate, grayish brown to brown. Fig. 7. 

Distribution and habitat—Widely distributed weed in tropical and subtropical re- 
gions of the Eastern Hemisphere; central and eastern Asia, Japan, southeastern Asia, In- 
donesia, Malaysia, Philippines, Australia, and eastern Africa. Introduced into the outer 
coastal plain of the southeastern United States of North America, ranging from south- 
eastern Louisiana into southwestern Alabama with an isolated station in southeastern 
Georgia (Fig. 8). In the southeastern United States often locally common and weedy in 
periodically disturbed sites with high hydro-period soils (e.g., road ditches, margins of 
artificial ponds, etc.). 

Phenology.—I|n the southeastern United States, flowering and fruiting from Sep- 
tember until frost and sporadically earlier. 


a 


339 


Specimens examined. North America. U.S.A. ALABAMA. Baldwin Co.:Foley, 17 Sep 1994, Burkhalter 
14368 (VSC). Mobile Co.: Mobile, Battleship Park, Hwy. US 90, 30 May 1994, Mears 94-25 (ctb,VSC); 12 
Sep 1995, Carter 12705 (VSC). GEORGIA. Camden Co.: Kings Bay Submarine Base, 0.2 mi E jct.U.S.S. 
Henry L. Stimson Dr.and James Madison Rd., U.S.S. Henry L. Stimson Dr, 11 Oct 1996, Carter 13873 
(VSC);just N jct.U.S.S. Benjamin Franklin Rd.and U.S.S. Georgia Ave., U.S.S. Georgia Ave., 25 Oct 1996, 
Carter 13939 (VSC); ca. 100 mN ject. U.S.S. Benjamin Franklin Rd. and i S.S. James Madison Rd., U.S.S. 
mes Madison Rd., 25 Oct 1996, Carter 13940 (VSC); ca. 200 m S$ jet. U.S.S. Henry L. Stimson Dr. and 
ee Kamehameha Ave., U.S.S. Kamehameha Ave., 25 Oct 1996, aoe 13941 (VSC); 0.09 mi N ject. 
U.S.S. Daniel Webster Rd. and U.S.S. Benjamin Franklin Rd., U.S.S. Daniel Webster Rd., 25 Oct 1996, 
Carter 13954 (VSC); ca. 300 m E Franklin Gate, S side U.S.S. Benjamin Franklin Rd.,4 Dec 1996, Carter 
13962 (VSC). LOUISIANA. St. Tammany Parish: Goodbee, 12 Oct 1960, Hebert 377 (MISS); Hwy. US 
190,0.35 mi E jct. Hwy.US 190 and LA 1077 in Goodbee, 18 Sep 1993, Carter 11367 (VSC); Slidell, Hwy. 
US 190E, 0.84 mi W jct. Hwy. US 190E and I-10, 15 Oct 1993, Carter 11490 (VSC); Slidell, ICG Railroad 
right-of-way, 0.14 mi N jct. Hwys. LA 433 and US 11,15 Oct 1993, Carter 11505 (VSC); Slidell, Hwy. US 
190, 0.10 mi W jct. Hwys. US 190 and US 11, 16 Oct 1993, Carter 11539 (VSC); Slidell, Hwy. US 19 
mi W jct. Hwys. US 190 and US 11, 16 Oct 1993, Carter 11540 (VSC); Lacombe, Hwy. US 190 at Tran- 
quility Road, 16 Oct 1993, Carter 11541 (VSC); Slidell, Hwy. US 11,250 ft.S jct. Hwy. US 11 and Carollo 
Avenue, 17 Oct 1993, Carter 11558 & Bryson (VSC), Bryson 13218 & Carter (ctb,VSC); Slidell, Hwy.|-10,S 
jct. Hwys. I-10 and US 190, 17 Oct 1993, Carter 11561 & Bryson (VSC); Slidell, SW jct. Hwys.!-10 and LA 
533,17 Oct. 1994, Bryson 14565 & MacDonald (ctb,VSC). Tangipahoa Parish: ca. 7 mi SE Ponchatoula, 
ca.1 mi N Lees Landing, 16 Nov 1989, Gilmore 3977 (VSO), Gilmore 3978 (ctb, VSC); 19 Sep 1993, Carter 
11374 (VSC); Ponchatoula, Hwy. LA 22,0.5 mi E jct.LA 22 and North First Street, 17 Sep 1993, Carter 
11355 (VSC); Ponchatoula, Hwy.LA 22,W jct. Hwy.LA 22 (E. Pine Street) and West Street, 18 Sep 1993, 
Carter 11372 (VSC). MISSISSIPPI. Hancock Co.: Hwy. MS 43, 0.44 mi N jct. Hwy. US 90 and MS 43 in 
Waveland, 16 Sep 1993, Carter 11342 (VSC);0.8 mi N jct.US 90 and MS 43 in Waveland, 16 Oct 1993, 
Bryson 13166 & Carter (ctb, SWSL,VSC), 17 May 1994, Bryson 13535 (ctb,VSC); Hwy. MS 43,5.25 mi E jct. 
Hwys. MS 43 and I-59 in Picayune, 18 Oct 1993, Carter 11567 & Bryson (VSC), Bryson 13265 & Carter 
(ctb, SWSL, VSC); 9.0 mi NW jct. Hwys.43 and 603 in Kiln, Hwy. MS 43, pipeline crossing, 18 Oct 1993, 
Carter 11568 & Bryson (VSC); 17 May 1994, Bryson 13542 (ctb, VSC); 21 Oct 1997, Bryson 16217 (ctb, 
SWSL,VSC);N of Kiln, 0.6 mi.S jct.of Hwys.MS 43 and MS 603, 18 Oct 1993, Bryson 13267 & Carter (ctb, 
SWSL); Kiln, Hwy. MS 43, 0.6 mi S jct. Hwys. MS 43 and MS 603, 18 Oct 1993, Carter 11569 & Bryson 
(VSC); Kiln, Hwy. MS 43, vicinity Shifalo Baptist Church and Kiln Post Office, 18 Oct 1993, Carter 11570 
& Bryson (VSC), Bryson & Carter 13268 (ctb, SWSL, VSQ); 21 Oct 1997, Bryson 16216 (ctb, VSC); jct. of 
Hwys. MS 43 and I-10 between Kiln and Waveland, 18 Oct 1993, Carter 11571 & Bryson (VSC); SE jct. 
Hwys.|-10 and MS 43, 18 Oct 1993, Bryson & Carter 13271 (ctb, SWSL,VSC), 17 May 1994, Bryson 13534 
(ctb, VSC); E of Picayune, 5.8 mi E jct. Hwys. |-59 and MS 43, 16 Oct 1994, Bryson 14537 & MacDonald 
(ctb, SWSL, VSC); Mississippi Welcome Center, SE jct. Hwys. I-10 and MS 607, 17 Oct 1994, eas 
14567 & MacDonald (ctb, SWSL, VSC); Kiln, NW jct. Hwy. MS 43 and Kiln-Delisle Road, 17 Oct 
Bryson 14597 & MacDonald (ctb, SWSL, VSC); Waveland, Nicholson Avenue, 0.2 mi S jct. Hwys. if Ss 
and MS 43, 18 Oct 1994, Bryson 14608 & MacDonald (ctb, SWSL, VSC); Waveland, Central Avenue, 
between Central Avenue and RR just W Washington Street, 18 Oct 1994, Bryson 14610 & MacDonald 
(ctb, SWSL, VSC); Waveland, NW jct. Hwys. US 90 and MS 43, 7 Dec 1994, Bryson 14709 (ctb, SWSL, 
VSC); Waveland, NW jct. Hwys. US 90 and MS 43,21 Oct 1997, Bryson 16214 (ctb,VSC), 20 Nov 1998, 
Bryson 16939 & Sudbrink (ctb, SWSL,VSQ); Waveland, 1.2 mi S Hwy. US 90 on Nicholson Avenue, then 
1.6 mi E on Central Avenue, 21 Oct 1997, Bryson 16215 (ctb, SWSL,VSC). Harrison Co.: Orange Grove 
Community Center Park, W Hwy. US 49, 0.3 mi N jct. Hwys. US 49 and I-10, 16 Oct 1993, Bryson 13164 
& Carter (ctb, SWSL, VSC), Carter 11544 & Bryson (VSC); Orange Grove, Hwy. US 49, 1.1 mi S jct. Hwy. US 
d O'Neal Road, 18 Oct 1993, Bryson 13276 (ctb, SWSL,VSC); N Gulfport, Harrison Drive, 0.3 mi W 
jct. Harrison Drive and MLL. King Jr. Blvd., 18 Oct 1993, Carter 11574 (VSC); Popps Ferry Road, 3.32 mi 


340 SIDA 19(2) 


W jct. Popps Ferry Road and D'lberville Boulevard (Hwy.MS 67), 18 Oct 1993, Carter 11577 (VSC); NW 
jct. Hwys. |-10 and US 49, 18 Oct 1993, Bryson 13279 (ctb, SWSL, VSC); Long Beach, SE jct. Klondyke 
and Commission Road, 18 Oct 1994, Bryson 14606 & MacDonald (ctb, SWSL, VSC). Jackson Co.: 
Pascagoula, SE jct. Washington Ave. and Louise St., vic. Bayou Casotte, T8S RSW S17, 16 Sep 1991, 
Bryson 11032 (ctb, NY, NYS, VSC); 16 Sep 1993, Carter 11337 (VSO); St. Martin, Rosefarm Road, 0.2 mi N 
jct. Old Fort Bayou Road and Rosefarm Road, 18 Oct 1993, Carter 11579 (VSC); vicinity St. Martin, 1.13 
mi W jct. Old Fort Bayou Road and Yellow Jacket Boulevard, 18 Oct 1993, Carter 11580 (VSC); vicinity 
St. Martin, Old Fort Bayou Road, 0.19 mi E jct. Old Fort Bayou Road and Yellow Jacket Drive, 18 Oct 
1993, Carter 11581 (VSC); Pascagoula, 16 Oct 1994, Bryson 14547 & MacDonald (ctb, VSC); Moss Point, 
SE jct. Hwys. 1-10 and MS 63, 16 Oct 1994, Bryson 14550 & MacDonald (ctb, SWSL, VSC); Latimer, 1.2 mi 
N jct. Hwys. I-10 and MS 609, 1.2 mi N Tucker and Cook Roads, along Tucker Road, 16 Oct 1994, 
Bryson 14559 & MacDonald (ctb, SWSL, VSC); Pascagoula, on Hwy. US 90, 1.5 mi.W jct.of Hwys. US 90 
and MS 63, 6 Nov 1994, MacDonald 8195 & Leidolf (ctb, SWSL); N Ocean Springs, Hwy. |-10, E mi 
marker 50,19 Oct 1994, Bryson 14636 (ctb, SWSL,VSC). Pearl River Co.: Picayune, Hwy. MS 43,0.50 mi 
W jct. Hwys. MS 43 and I-59, 18 Oct 1993, Carter 11562 (VSC), 28 Oct 1998, Bryson 16874 & Sudbrink 
(ctb, VSC); Picayune, frontage road W Hwy. I-59, ca. 250 m N jct. Hwys. MS 43 and I-59, 18 Oct 1993 
Bryson 13222 & Carter (ctb, SWSL, VSC), Carter 11565 & Bryson (VSC), 29 Oct 1998, Bryson 16900 & 
roe (ctb, SWSL,VSC), 9 Nov 1999, Bryson 17730 & Sudbrink (ctb, SWSL, VSC); Picayune, N side of 
43,0. . mi. i ct.of Hwys.MS 43 and I-59, 18 Oct 1993, Bryson 13257 & Carter (ctb, SWSL,VSC); 
ie SW jct. Hwys. I-59 and MS 43 S, 19 Oct 1994, Bryson 14634 (ctb, SWSL, VSC); Picayune, 21 
Oct 1997, ae ae (ctb, VSC); Picayune, 21 Oct 1997, Bryson 16213 (ctb, VSC). Stone Co.: Wiggins, 
NW jct. Hwys. US 49 and MS 26, 22 Oct 1997, Bryson 16219 (ct 
EURASIA. RUSSIA. Primorski, Distr. Michaelowsky, Bepeks Valley, 1 2 Sep 1929, Baianova 647 
(NY). BHARAT (INDIA). Bengal, Griffith s.n. (NY); Hassan District, Mysore, tank near Dandiganahalli, 
11 Nov 1971, Hooper & Gandhi HFP 2401 (MO); Sikkim, J.D. Hooker s.n. (NY); Maharashtra, Nagpur, 
Ambala, 2-10-1962, Donde D28 (NY); Chamba, Khajiar, 11.7.1936, Koelz 8833 (NY); Pahlgam, 16 Aug 
1920, Stewart & Stewart 5709 (NY); Srinagar, Dal Lake, 13 Jul 1917, Stewart 3274 (NY); Kulu, Rotang 
Pass, Sep 1930, Koelz 1359 (NY).PAKISTAN. Baltistan, ca. 1.5 mi E Skardu, alt.Ca. 7500 ft, 26 i 1955, 
Webster 6585 (GH); Skardu to Shigar, 8 Jul 1940, Stewart 20472 (NY); Kishenganga Valley and r 
Nanga Parbat, below Wangat, Sind Vy.,7 Aug 18080, Stewart & Stewart I Ht NEPAL. Kali end 
Stainton, Sykes & Williams 9238 (GH, NY); Kali Gandaki, Tatopani, S of Dana, 30 Aug 1954, Stainton, 
Sykes & Williams 7591 (NY); Bongakhani, 22 Aug 1954, Stainton, Sykes & Williams 3954 (BRIT, NY); Aruna 
Valley, Sedua, NW of Num, 31 May 1956, Stainton 485A (NY); Samri Khola, 7 Apr 1953, Gardner 141 
(NY); Argam, near Pokhara, 11.9.1954. Stainton, Sykes & Williams 7178 (NY); Jajarkot District, Maina 
gaon, 12 Aug 1979, Rajbhandara & Roy 4585 (NY);Rukum District, Gija gaon, 18 Sep 1982, Rajbhandara 
& Malla 6535 (NY); Sindjuli District, Patlebhaniyang, 2 22 Dec 1975, lees & A ae 3343 (NY); 
Dolakha District, Lamabagar to Hum, 16 Jul 1977 bhandara & 8 (NY). CEYLON. Amparia 
ate Senanaike Samudra, Padagoda, 6 Feb 1 1971, Kovane 13983 fa read, a Dec 1967, 
Comanor 690 (NY); Central Province, Kandy District, ca. 5 mi SE Gampola, 24 Oct 1974, Davidse & 
Sumithraarachchi 7924 (NY); Sabaragamuwa Prov. Ratnapura District, 11 miE aa Oct 1974, 
Davidse 7886 (MO, NY); Northwestern Prov., Wilpattu National Park, Manikepola Uttu, 24 Mar 1968, 
Heart & Cooray 13460 (NY); Amparai District: Senanaike Samudra, Padagoda, 6 Feb 1971, Koyama & 
Balakrishnan 13983 (NY). CHINA. Prov. Hainan, Janfengling, Chow 78471 (GH); Prov. Hunan Sheng, 
Xinning Xian, Ziyun Shan, 13 Sep 1984, Li Zhen-yu et al, 1826 (MO); Prov. Hupeh, 1885-88, Henry 2907 
(GH); Prov. Kiangsi, Dagangshan, Fenyi City, 24 Aug 1985, Yao 9193 (GH, NY); Prov. Sikang, 1935- 36, 
Wang 66442 (GH); Shanxi, Kolan Hsien, 31 Aug 1929, lang 173 (NY); Sichuan, Dujiangyan, E of Longxi 
6 Sep 1988, Boufford & Bartholomew 24675 (MO, NY); Stee Aug 1930, Tsiang 6512 (NY), Nov 
1930, Tsiang 9542 (NY); Manchuria, Prov. Kininensis, 18 Jul 1896, V Komarov 219 (NY); Kwangtung 
Prov. Chong Uen Shan near Kau Fung, 2-30 Nov 1932, Tsang 20665 (NY); Yunnan, Ducloux 252 (NY): 


— 


341 


Hong Kong, Lantao Island, Tungchung, Taishui-hang, 29 Sep 1940, Taam 1819 (NY). TAIWAN. 
Niitagagun, Hosya, Kusunokizinzya, 14 Oct 1935, S. Suzuki s.n. (MO); Little Quemoy, 16 Sep 1961, 
Chuang 4469 (GH); Botel Tobago, 16 Aug 1972, Chang 7110 (NY). JAPAN. Hondo, Yamanakako in Kai, 
3 on 1952, K. Okamoto NSM 584 RIT, NY); Prov. Tootoomi, Pref. Shidzuoka, Hondo, 30 Sep 1959, 
fiyoshi Furuse s.n. (GH), 11 Oct 1960, Miyoshi Furuse s.n. (GH); Prov. Kadzusa, Pref. Chiba, Hondo, 5 Sep 
1962, Miyoshi Furuse s.n. (GH); Prov. Samashire: Mt. Hieizan OR 9 Sep 1931, Tagawa 617 
(NY); Pref. Mie, Ogurusu, Kiwa-cho, Minamimuro-gun, 17 Nov 1977, Mimoro, Tsugaru & Deguchi 4289 
(MO); Pref. Settsu, Ashiya, 10 Sep 1954, Hutoh 17517 (NY). 

SOUTHEASTERN ASIA. VIETNAM. Tonkin, Chapa, Petelot 6099 (NY). THAILAND, Phetchabun, 
Phu Miang, 2.10.1967, Schimizu et al. 7.11391 (NY); Chiang Mai, Doi Intanoid, 13 Sep 1974, Larsen & 
Larsen 34512 (NY); Maehongson, Khun Yuam, 7 Sep 1974, Larsen & Larsen 34254 (NY). INDONESIA. 
Lesser Sunda Islands, Flores, near Keli Moetoe, Kampong Ndoearia, 8-6-1938, Jaag 1801 (GH); Java, 
Mt Gedeh, Tjibodas, 30 Apr 1950, Kern he (GH). PHILIPPINES. Northern Luzon, Prov. Benguet, 
ae . se 1904, ‘alae 1972 (GH,N 
A. QUEENSLAND. i District, Brisbane, 5 Aug 1937, Blake 12965 (GH). 

pe ae Cusai, 16 Sep 1902, Pappi 1799 (NY) 


ACKNOWLEDGMENTS 


We gratefully acknowledge the following individuals and agencies: Cary Norquist, United 
States Fish & Wildlife Service, for providing financial support for the first author's field 
work in Louisiana and Mississippi during 1993; Nelwyn Gilmore, Louisiana Natural Heri- 
tage Program, for providing data and specimens she collected at the holotype locality; 
Georgia Natural Heritage Program for funding the first author's field work during 1996 at 
Kings Bay Submarine Base,Camden County, Georgia; Ron Wilkerson for assistance at Kings 
Bay Submarine Base and providing information on road maintenance practices there; 
J.R. Burkhalter and Randy L.Mears for sending Alabama specimens to the first author for 
determination; John R. MacDonald, Don Sudbrink, and Shaharra Usnick for assistance 
with fees collections; Serial RENE) for providing rice production statistics for Hancock 
County, Mississippi; Russ Goddard | for assistance with and lending of photomicrographic 

quipment; eGorden C. ei for providing helpful comments on the manuscript; and 
the following herbaria for lending specimens without which completion of this project 
would not have been possible: BRIT, GH, MO, NY and US 


REFERENCES 


Apams, C.D. 1994. Cyperaceae. In: G. Davidse, M.S. Sousa and A.O. Chater, eds. Flora 
Mesoamericana, Vol. 6: Alismataceae a Cyperaceae. Universidad Nacional Aut6noma 
de México, Cuidad Universitaria.Pp. 262-485 

Anonymous. 1959. United States Census of Agriculture, Vol. 1, Part 33, Mississippi. United 
States Department of Agriculture. 

Anonymous. 1982. United States Census of Agriculture, Part 24, Mississippi. United States 
Department of Agriculture. 

Anonymous. 1993.Endangered and threatened wildlife and plants; review of plant taxa for 
listing as endangered or threatened species; Department of Interior, Fish and Wildlife 
Service, 50 CFR Part 17. Federal Register 58 (188):51144-51190. 


342 SIDA 19(2) 


Anonymous. 1996, Minitab reference manual: release 11 for Windows™. Minitab Inc., State 
College, PA. 

Bru, J. 1995. Sedge genera of the world: relationships and a new classification of the 
Cyperaceae. Aust. Syst. Bot. 8:125-305. 

Bryson, C.T. and R. Carter. 1992. Notes on Cyperus and Kyllinga (Cyperaceae) in Mississippi 
with records of six species new to the state. Sida 15:119-124. 

Bryson, C.T. and R. Carter. 1994. Notes on Carex, Cyperus, and Kyllinga (Cyperaceae) in Mis- 
sissippi with records of eight species previously unreported to the state. Sida 16:171- 
182. 

Bryson, C.T., R. Carter, L.B. McCarty, and F.H. Yetverton. 1997. Kyllinga, a genus of neglected 
weeds in the continental United States. Weed Technology 11:838-842. 

Bryson, C.T., J.R. MacDonato, R. Carter, and S.D. Jones. 1996. Noteworthy Carex, Cyperus, 
Eleocharis, Kyllinga, and Oxycaryum (Cyperaceae) from Alabama, Arkansas, Georgia, 
Louisiana, Mississippi, North Carolina, Tennessee, and Texas. Sida 17:501-518. 

Carter, R. 1990.Cyperus entrerianus (Cyperaceae),an overlooked species in temperate North 
America. Sida 69-77. 

Carter, R.and C.T. Bryson. 1996. Cyperus entrerianus:A little known aggressive sedge in the 
southeastern United States. Weed Technology 10:232-235. 

Carter, R., R.L. Mears, K.C. Burks, and C.T. Bryson. 1996. A report of four exotic Cyperus 
(Cyperaceae) species new to Florida, U.S.A. Sida 17:275-281. 

Crarke, C.B. 1894. Cyperaceae. In: J.D. Hooker, ed. The flora of British India, Vol. VI, Part XIX. L. 
Reeve & Co., London. Pp. 585-672. 

Crarke, C.B. 1908. New genera and species of Cyperaceae. Kew Bull. Add. Ser. 8:1-196. 

Corcoran, M.L.1941.A revision of the subgenus Pycreus in North and South America.Catho- 
lic Univ. Amer. Biol. Ser. 37. 


GoetGHeBeuR, P. 1986.Genera Cyperacearum:een bijdrage tot de kennis van de morfologie, 
systematiek en fylogenese van de Cyperaceae—genera. Doctoral Thesis, Rijksuniversiteit 
Gent. 

GoeTcHeBeuR, P. 1989. Studies in Cyperaceae 9. Problems in the lectotypification and 
infrageneric taxonomy of Cyperus L. Bull. Soc. Roy. Bot. Belg. 122:103-114. 

Haines, R.W. and K.A. Lye. 1983. The sedges and rushes of east Africa. East African Natural 
History Society, Nairobi. 

Hoi, L.G., J.V. PANCHO, J.P. Hersercer, and D.L. PLucknett. 1991.A geographical atlas of world 
weeds. Krieger Publishing Company, Malabar, Florida. 

Kern, J.H. 1974. Cyperaceae. In: C.G.GJ. van Steenis, ed. Flora Malesiana, Series |, Vol. 7(3). 
Noordhoff, Leyden. Pp. 435-753. 

Koyama, T.1985.Cyperaceae. In: M.D. Dassanayake, and F.R. Fosberg, eds. Flora of Ceylon, Vol. 
5.Amerind Publishing Co. Pvt.Ltd., New Delhi. Pp. 125-405. 

Kral, R.1971.A treatment of Abildgaardia, Bulbostylis and Fimbristylis (Cyperaceae) of North 
America. Sida 4:57-227. 

KUHN, U. 1982. Cyperaceae. In: Hafliger, E., et al. Monocot Weeds 3. Ciba-Geigy Ltd., Basil. 


343 


KUKENTHAL, G. 1935-1936. Cyperus. In:A.Engler and L. Diel, eds. Das Pflanzenreich IV. 20 (Heft 
101). Pp. 1-671. 
KUKKONEN, |. 1995. Two proposals to conserve species names in Cyperaceae. Taxon 44:625- 


Mayr, E. 1942. Systematics and the origin of species. Columbia University Press, New York. 

McKenzie, PM.,B. JAcoss, C.T. Bryson, G.C. Tucker,and R. Carter. 1998. Cyp (Cyperaceae) 
new to Missouri and Nevada, with comments on its occurrence in North America. 
Sida 18:325-333. 

Mincyuan, X. and M. Denu. 1970. Farmland weeds in China: a collection of coloured illus- 
trated plates. Agricultural Publishing House, Beijing, China. 

Oxwi, J. 1965. Flora of Japan. Smithsonian Institution, Washington, D.C 

Reep, C.F. 1977. Economically important foreign weeds. Agriculture Handbook No. 498. 
United States Department of Agriculture, Washington, D.C 

Stearn, W.T. 1992. Botanical Latin.4th Ed. David and Charles Publishers. Brunel House. New- 
ton Abbot, Devon, England 

Tuieret, J.W. 1977. Cyperus louisianensis (Cyperaceae),a new species from southern Louisi- 
ana. Proc. Louisiana Acad. Sci. 40:23-26. 

Tucker, G.C. 1983. The taxonomy of Cyperus (Cyperaceae) in Costa Rica and Panama. Syst. 
Bot. Monog. 2:1-85. 

Tucker, G.C. 1987.The genera of Cyperaceae in the southeastern United States. J. Arnold 
Arbor. 68:361-445. 

Tucker, G.C. 1994.Revision of the Mexican species of Cyperus (Cyperaceae). Syst. Bot. Monog. 
43:1-213. 

Witson, K.L. 1993. Cyperaceae. In: G.J. Harden, ed. Flora of New South Wales, Vol. 4. New 
South Wales University Press, Kensington. Pp. 293-396. 


344 SIDA 19(2) 


BOOK REVIEW 


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both a sense of humor and general enthusiasm about trees. Peter Thomas, lecturer in environmen- 


tal sciences at Keele University, UK, brings together information 1 through myriads of jour- 
nals and books from all parts of the world into one concise text, Trees: Their Natural History. Not only 
vital to healthy ecosystems, but also unparalleled in the range of materials they provide for human 


use, trees are a escinating group of plants. This book—written for a non-technical audience— 
answers such questions as: How do trees work? How are they designed? And how do they grow 
and reproduce? And in fewer than 300 pages, Thomas provides a comprehensive introduction to 
the natural history of trees covering their biology and ecology 

Beginning with a preface and an overview of trees, then followed by eight chapters that 
more specifically address the parts of trees and how they live and die. Chapter Two covers the 
leaves, Chapter Three the trunk and branches, Chapter Four the roots, Chapter Five flowers, fruits, 
and seeds, Chapter Six,"The growing tree," Chapter Seven, the shape of trees; Chapter Eight,"New 
trees from old,”and Chapter Nine,"Health, damage, and death."The book is then concluded by a list 
of further reading and an index. Boxes, black and white photographs, and black and white line 
drawings are abundant throughout and used appropriately. Another pleasing feature is the list of 
further reading that follows each chapter. These lists are chapter specific and contain books and 
journal references not Iie in the list of further reading at the end of the boo 

The author succeeds in his goal to “draw together strands of information to create a readable 
book that would answer common questions about trees, set right a number of mytl 
the remarkable world of how trees work, grow, reproduce and die."While written for lay audiences, 
the book is substantially scientific and the references at the end of each chapter offer the reader 
more specific academic follow-up information as they desire. Trees: Their Natural History would be 
suitable for introductory botany classes studying trees as well as lay readers looking for answers to 
questions they've wondered about for years.—Kevin D. Janni, Botanical Research Institute of Texas, 
Kjanni@brit.org. 


SIDA 19(2): 344. 2000 


CYPERUS (SUBG. QUEENSLANDIELLA) HYALINUS 
(CYPERACEAE) NEW TO THE UNITED STATES AND THE 
WESTERN HEMISPHERE 


Richard Carter 


Herbarium, Biology Department 
Valdosta State University 
Valdosta, GA 31698-0015, U.S.A. 


Randy L. Mears 


11399 SW 66th Street 
Miami, FL 33173, U.S.A. 


ABSTRACT 


Cyperus hyalinus Vahl is reported from Dade County, Florida, U.S.A. This paleotropical species has 
not been previously reported from the Western Hemisphere. Its taxonomy and pest potential are 
discussed. A dichotomous key to the subgenera of Cyperus in the United States, technical descrip- 
tion, and photographs are provided, and collection data and voucher specimens are cited. 


RESUMEN 
Se cita Cyperus hyalinus Vahl del condado de Dade, Florida, U.S.A. Esta expecie paleotripical no ha 
sido citada previamente del hemisferio occidental. Se discute su taxonomia y su potencial como 
mala hierba. Se ofrece una clave dicotomica de los subgéneros de Cyperus en los Estados Unidos, 
descripcidn técnica, y fotografias, y se citan los datos de la recolecci6n y los pliegos testigo. 


INTRODUCTION 


In late 1999, the second author discovered a population of an unknown Cyperus species 
in Dade County, Florida. Specimens were sent to the first author for determination and 
were identified by him as C. hyalinus Vahl. Cyperus hyalinus has a wide paleotropical dis- 
tribution, ranging from eastern Africa, Madagascar, Mauritius, India, Sri Lanka, tropical 
Australia (Queensland),and Malaysia (KUkenthal 1935—1936;Kern 1974; Haines & Lye 1983; 
Koyama 1985). This remarkable little sedge has not been previously reported from the 
Western Hemisphere. 

The taxonomic relationships of C hyalinus are obscure,and its nomenclature is com- 
plex.In addition to Cyperus, the species has been treated in the segregate genera Pycreus, 
Kyllinga,and Queenslandiella.|t has also been placed in various subgenera of Cyperus, i.e, 
subg. Kyllinga (Kern 1974), subg. Mariscus (KUkenthal 1935-1936), subg. Pycreus (Clarke 
1884), and subg. Queenslandiella (Govindarajalu 1975; Haines & Lye 1983). Its lenticular 
achene, bifid style, compressed spikelets with multiple flowers and fruits, and open 
anthelate inflorescence suggest a relationship with subgenus Pycreus. However, persis- 
tent scales and disarticulating spikelets defy placement there and indicate an affinity 
with Kyllinga or Mariscus. Because its treatment as a Pycreus, Kyllinga, or Mariscus is prob- 


SIDA 19(2): 345 — 350. 2000 


346 SIDA 19(2) 
lematical, one could segregate it in the monotypic genus Queenslandiella; however, con- 
sistency would require the segregation of other genera from Cyperus, which would up- 
set current nomenclature. Based upon a study of its anatomy, Govindarajalu (1975) placed 
C. hyalinus in monotypic subg. Queenslandiella of Cyperus. Until there is unequivocal 
molecular evidence to the contrary, we think a broadly defined Cyperus similar to the 
concept of Haines and Lye (1983) is of value in conserving current nomenclature. Thus, 
we treat this species in Cyperus subg. Queenslandiella. 

Our objectives herein are to report C. hyalinus new to Florida, U.S.A.,and the West- 
ern Hemisphere and to provide a dichotomous key, technical description, notes, and 
photographs to facilitate its identification in the United States. 


TAXONOMY 
KEY TO THE SUBGENERA OF CYPERUS IN THE UNITED STATES 
1. Spikelet rachilla remaining attached to rachis, not basally articulated; floral scales 
and achenes disarticulating from base to apex of rachilla. 
2. Style branches 3; achenes trigono 
3. eae las eas ie notin digitate clusters; plants of various habi- 
y present. subg. Cyperus L. 
3; ee in arene hee ales of hydric to mesic habitats; kranz anatomy 


absent. ubg. Anosporum CB. Clarke [=subg. Pycnostachys C.B. Clarke] 
2. style praneves 2; peut eneculet rarely turgid is ao 
4. C 


gle adj achilla. subg. Pycreus 

A. Gray 

4. Spikelets cylindrical: achene face adjacent to rachilla. subg. Juncellus (Griseb.) 
C.B. Clarke 


1. Spikelet, floral scale,and achene articulation not as above. 

5. Style branches 3;achenes trigonous; spikelet basally articulated and deciduous 
as a unit with floral scales and achenes still attached or spikelet breaking apart 
transversely into 1-fruited segments 
6. Spikelet basally articulated decide lous as a unit with floral scales and achen 

still attached to rachilla. subg. Mariscus (ahi C.B. Clarke 
6. Spikelet breaking apart transversely into 1-fruited segments. subg. Diclidium 
(Schrad. ex Nees) C.B. Clarke [=subg. Torulinium (Desv.) Kk} 

5. Style branches 2; achenes lenticular to plano-compressed; spikelet basally ar- 
ticulated, deciduous as a unit with floral scales and achenes still attached to 
rachilla. 

7. Floral scales more than 2 per spikelet;flowers and fruits more than 1 per cau 
let; inflorescence an open anthelus of mostly ae spikes; achene 
plano-compressed. g. Queenslandiella oan Govind. 
Te iG scales 2 per spikelet; flowers and fruits 1 per ee inflorescence capi- 
ate, of 1-several dense sessile spikes; achenes lenticular. subg. Kyllinga (Rottb. 
JV. Suringa 


a 


Cyperus subg. Queenslandiella (Domin) Govind. Reinwardtia 9:194. 1975. 


Inflorescence an open anthelus of mostly pedunculate spikes. Spikelets with 2 or more 
flowers and achenes, basally articulated, falling intact. Floral scales and achenes persis- 


347 


tent. Style bifid. Achene plano-compressed, angle adjacent to rachilla. Kranz 
(chlorocyperoid) anatomy. Subgenus monotypic. 


Cyperus hyalinus Vahl, Enum. PI.2:329. 1805. Tyee: INDIA (C).Queenslandiella mira Domin, Biblioth. 
Bot.85:416.1915.Mariscopsis suaveolens Cherm. Bull.Mus. Hist. Nat. (Paris) 25:60. 1919. Pycreus 
hyalinus (Vahl) Domin, Biblioth. Bot.85:417.1915.Mariscopsis hyalinus (Vahl) F.Ballard, Bull. Misc. 
Inform. Kew 9:458. 1932. Queenslandiella hyalina (Vahl) F. Ballard in Hook. Icon. PI. 33:t. 3208. 
1933. Kyllinga hyalina (Vahl) T. Koyama, J. Jap. Bot. 51:313. 1976. 


Loosely cespitose tic annual herb. Roots fibrous, brown. Stems glabrous, trigonous, 
3-14 cm x 1-2 mm. Leaves 3-7, basal; bases sheathing; blades 4-15 cm x 2-5 mm. Pri- 
mary inflorescence bracts 4-8, mostly exceeding rays, longest to 12 cm long, 2-4 mm 
wide. Inflorescence anthelate; rays 3-8, longest 2.5-4 cm long; spikes simple (rarely with 
short basal branch), mostly pedunculate, oblong-ovate, (7—)12-20 mm x 8-15 mm, with 
(5-)12-17 mostly divaricate spikelets; rachis grooved, winged. Bracteoles narrowly trian- 
gular to aristate, 0.4-2.2 mm long, membranous. Spikelet prophylls rounded to acute, 
0.7-1.4 mm long, membranous. Spikelets laterally compressed, narrowly ovate to elliptic, 
4.1-5.7 x 1.9-2.2 mm, deciduous; rachilla wing ca. 0.5 mm wide, membranous. Floral 
scales 4-7 [3-4 fertile], imbricate, broadly ovate, 2.1-2.4 mm long, mucronate, membra- 
nous; keel green, scabrid; wings yellowish to whitish to pale green; lateral nerves 6-8. 
Stamens 2; anthers narrowly oblong, 0.4-0.5 mm long. Style bifid, divided ca. 3/4 of its 
length. Achene brown, broadly oblong to suborbicular, 1.0-1.4 x 1.0-1.1 mm, plano- 
compressed, 0.3 mm thick, gibbous, truncate-retuse, minutely puncticulate. Fig. 1. 

Phenology.— |n the United States, flowering late July through November. 

Distribution.—In the Old World, ranging from eastern Africa, Madagascar, Mauritius, 
India, Sri Lanka, tropical Australia (Queensland), and Malaysia (KUkenthal ibid.; Kern 1974; 
Haines & Lye 1983; Koyama 1985).Herein reported new to the Western Hemisphere, where 
so far it is restricted to southern Florida, U.S.A. Fig. 2. 


pean 


Voucher specimens. U.S.A. FLORIDA. Dade Co.: E side of Miami International Airport, just N of 
Perimeter Rd, road shoulder, sandy soil, 26 Oct 1999, Randy Mears s.n. (EIU, FLAS, MICH, MO, US, USF, 
VDB,VSC). 


DISCUSSION 


Cyperus hyalinus (Fig. 1) is readily distinguished from all other congeners by the follow- 
ing combination of characteristics: broadly oblong, truncate-retuse, plano-compressed 
achene; bifid style; 3-4-flowered, deciduous, flattened spikelets; membranous, yellowish 
to pale greenish, 6-nerved, mucronate floral scales with scabrid keel;and open anthelate 
inflorescence. Morever, dried specimens exhibit the distinctive odor of fenugreek 
(Trigonella foenum-graecum L.), previously noted by various authors (e.g., Kern 1974; 
Govindarajalu 1975; Bruhl 1995) and also characteristic of C. fuscus L., C. setigerus Torr. & 
Hook, and C. squarrosus L. (see McKenzie et al. 1998). 

Herein, the range of C. hyalinus is extended to North America (Fig. 2), where it is the 
most recent in a series of exotic Cyperus spp. reported new to the United States (Carter 


SIDA 19(2) 


1 and dried i (R. Mears s.n., 26 Oct 1999), scale bar=1 


Fic. 1. Cyperus hyalinus Vahl.— 
m.—B. Photograph sa scale bar= in mm. Jc Photograph of achenes, scale bar=1 mm 


— ee ee 
64 = 
ms svt 
ee e, Ee 
¢ ~ . 
y eg 
‘So 
. 2} 
3 3) > a> 
; 4) e 
ay ‘ + XY AN 
LS N \ 
: \ 
eat \ x 4 
; 
e > 
6 
JA 
#e 
yy 
Ke (s we, 
yy is pes 
+ 3 a 
\ 
ve 
_ = SN > 
ea = 
Fic y) Id rr | Dee es) eee Wehkil I 4 re i 4\ Ji 


citations (Ballard 1932, 1933; Kiikenthal 1935— 1936; Kern 1974; Haines & Lye 1983; Koyama 1985). 


349 


1990; Carter et al. 1996; Carter & Bryson 2000). The broad dispersal of such species is not 
surprising given their weedy nature, their copious production of small fruits, and the 
current ease and frequency of rapid, long-distance transportation of humans and cargo. 
Its rarity and proximity to the Miami International Airport suggest a recent introduction 
of C.hyalinus via shipment of air-freight. 

In addition to the original population growing along an open, sandy road shoulder, 
the second author has discovered another one about one-half mile away along a rail- 
road right-of-way. Plants were observed again at these sites in July 2000 associated with 
Bidens alba (L.) DC., Cenchrus incertus M.A.Curtis, Chamaesyce hirta (L.) Millsp.,C. hyssopifolia 
(L.) Small, C. maculata (L.) Small, Dactyloctenium aegyptium (L.) Willd.ex Asch. & Schweinf, 
Polypremum procumbens L., Setaria parviflora (Poir.) Kerguélen, Sida elliotti/ Torr. & A. Gray, 


LS 


and Tridax procumbens bs 

The occurrence of C. hyalinus in Australia, Madagascar, Mauritius, and Zanzibar is 
sporadic (Ballard 1932, 1933), and it does not appear to exhibit aggressive or invasive 
properties in southern Florida. Thus, currently C. hyalinus would not seem to threaten 
native biota in the United States, and its tropical distribution in the Old World suggests 
establishment is unlikely in more temperate regions of North America. However, it should 
be monitored and additional populations sought in southern Florida, especially in light 
of its description as “a weed of sandy soils, near sea level” in eastern Africa (Haines & Lye 
1983). 


REFERENCES 


BaLtarb, F. 1932. LXIll.-The genus Mariscopsis. Bull. Misc. Inform. Kew 9:457-458. 

Battarb, F. 1933. Queenslandiella hyalina (Vahl) Ballard. Hooker's Icon. PI. 33:t. 3208. 

BruHL, J. 1995. Sedge genera of the world: relationships and a new classification of the 
Cyperaceae. Aust. Syst. Bot. 8:125-305. 

Caater, R. 1990. Cyperus entrerianus (Cyperaceae),an overlooked species in temperate North 
America. Sida 69-77. 

Carter, R. and C.T. Bryson. 2000. Cyperus sanguinolentus (Cyperaceae) new to the south- 
eastern United States, and its relation to the supposed endemic Cyperus louisianensis 
Sida 19:325-343. 

Carter, R., R.L. Mears, K.C. Burks, and C.T. Bryson. 1996. A report of four exotic Cyperus 
(Cyperaceae) species new to Florida, U.S.A. Sida 17:275-281. 

Crarke, C.B. 1884.On the Indian species of Cyperus. J. Linn. Soc., Bot. 21:1-202. 
GovinparaJatu, E. 1975. Studies in Cyperaceae. XIV. Endomorphic evidences for placing 
Cyperus hyalinus under the new subgenus Queenslandiella. Reinwardtia 9:187-195. 
Haines, R.W. and K.A. Lye. 1983. The sedges and rushes of east Africa. East African Natural 

History Society, Nairobi. 

Kern, J.H. 1974. Cyperaceae 1.In:C.G.GJ. van Steenis, ed. Flora Malesiana, Vol. 7. Noordhoff, 
Leyden. Pp.435-753. 

Koyama, T.1985.Cyperaceae. In: M.D. Dassanayake and F.R.Fosberg, eds. Flora of Ceylon, Vol. 
5.Amerind Publishing Co. Pvt. Ltd., New Delhi. Pp. 125-405. 


350 SIDA 19(2) 


KUKENTHAL, G. 1935-1936. Cyperus.|n: A.Engler and L. Diel, eds. Das Pflanzenreich IV. 20 (Heft 
101). Pp. 1-671, 

McKenzie, P.M.,B. Jacoss, C.T. Bryson, G.C. Tucker,and R. Carter. 1998. Cyperus fuscus (Cyperaceae), 
new to Missouri and Nevada, with comments on its occurrence in North America. 
Sida 18:325-333. 


LEGUMES FROM THE CENTRAL PART OF THE 
STATE OF CHIHUAHUA, MEXICO 


A.Eduardo Estrada C.and Alfonso Martinez M. 


Facultad ae Cienci ias HOIRSTCIES 


| Iniy ersidad A 


A.P.41,67700, Linares NI MEXICO 


ABSTRACT 


Our study of the legumes from the central part of the state of Chihuahua records 42 genera and 
117 species. The number of genera/number of species for each of the three leguminous subfami- 
lies are: Lotoideae 27/77; ule eolenl ee Caesalpiniodeae 7/12.Oak-pine est had the high- 
est number of species. Ger number of Dalea (19),Acacia (1 1),Phaseo- 
lus (7), Desmodium (6), and both Senna and Mimosa (5) Dalea Desmodium and Phaseolus are most 
common on oak-pine forest communities, while Acacia, Senna, and Mimosa are most common on 
shrublands and grasslands. 


RESUMEN 


Nuest tudio del | tral del estado de Chihuahua registré 42 géneros 
y 117 especies. El numero de géneros/ntimero de especies para cada una de las tres subfamilias es: 
Lotoideae 27/77; Mimosoideae 8/28; Caesalpinioideae 7/12. Los géneros con mayor numero de 
especies son Dalea (19), Acacia (11), Phaseolus (7), Desmodium (6) y Senna y Mimosa con (5). Dalea, 
Desmodium y Phaseolus son mas comunes en los bosques de encino-pino, mientras que Acacia, 
Mimosa y Senna son mas comunes en matorrales y pastizales. 


INTRODUCTION 


Legumes are constituted by three subfamilies of plants Mimosoideae, Caesalpinioideae 
and Lotoideae, and they are the second most diversified group of plants in Mexico after 
Asteraceae (Sousa & Delgado 1993). Legumes are found in all plant communities of the 
country, and are one of the dominant groups of plants in north Mexico. They are found 
on several types of shrublands of Nuevo Leon and Chihuahua (pers. obs). 

The study area covers 7500 km?,is located at 28° 15'-29° 05' N, 105° 07'-107° 35' W, 
and includes the eight municipios, Aldama, Aquiles Serdan, Chinuahua, Coyame, General 
Trias, Julimes, Meoqui and Riva Palacio on a surface of 7500 km? (Fig. 1). The study area 
includes two physiographic provinces (Anénimo 1987), the eastern part of the Sierra 
Madre Occidental and Sierras y Llanuras del Norte, which has two subprovinces, Bolson 
de Mapimi and the Sierras Plegadas del Norte. Most rocks are sedimentary or volcanic 
origin from the Quaternary Period. Most mountains in the area are acidic intrusive igne- 
ous rocks from Tertiary Period and metamorphic rocks (limestone) outcrops from the 
Mesozoic Era, and from the Lower Cretaceous Period (Anénimo 1981). The three main 
climate types for the area are: (1) very dry semi-warm, characterized by summer rains, 
the winter rainfall less that 5% of total, the winter cool—characteristic of western part of 


SIDA 19(2): 351 - 360. 2000 


352 SIDA 19(2) 


To Juarez City 


alata’ -1) 


To Ojinaga City 
/ 


To Cuauhtemoc City a 
28°15' 


107°35' To Camargo City 


Fic. 1. Map showing study area. 


study area, (2) dry, temperate, with summer rains, winter rainfall less than 5% of total, 
summer warm—occurring as a N-S-oriented 30-40 km wide strip in central part of study 
area; (3) Semi-warm, semidry temperate, with summer rains, the winter ranges 5—10.2% 
of total, summer warm—found on high plains and small mountains ranges at 1600- 
2400 m elev. These climatic types are mapped as BWhw(w), BSokw(w) and BS, kw, re- 
spectively, in Garcia (1973). 

There are three main vegetative communities, grasslands, shrublands and oak-pine 
forest (Rzedowski 1978). Bouteloua, Sporobolus, and Hilaria are the dominant genera in 
the grassland landscape, Bouteloua gracilis Lag, B. curtipendula (Michx.) Torr, B. eriopoda 
(Torr) Torr, B. hirsuta Lag, Sporobolus airoides (Torr. Torr, Hilaria mutica (Buckl.) Benth, 
Eragrostis intermedia Hitchc, E.lehmaniana Nees, Enneapogon desvauxii B and Lycurus 
phleoiodes H.B.K,. are the dominant grassland species found. Shrubland communities 
have mostly low elements, seldom over 2.5 m tall, forming subtt hrubl (Andonim 
1981), where Larrea tridentata (Sess. & Moc. ex DC.) Cav., Flourensia cernua DC., Acacia 
neovernicosa |sely, A. constricta Benth, A. schaffneri (Wats.) Herm., Lycium berlandieri Dun, 
Koeberlinia spinosa Zucc., Condalia ericoides (A. Gray) M.C. Johnst,, Yucca elata (Engelm.) 
Engelm,, Opuntia spp., Mimosa spp,, Celtis pallida Torr, Parthenium argentatum Gray and P 


ESTRADA C. AND MARTINEZ M., LEGUMES OF CENTRAL CHIHUAHUA, MEXICO 353 


incanum Kunth are the predominant species (Andénimo 1978). Oak-Pine forests are lo- 
cated in mountain areas, north of Cd. Chihuahua, especially Sierra El Nido and Parque 
Nacional Cumbres de Majalca, and 30-40 km west of Cd. Chihuahua in direction to Cd. 
Cuauhtemoc. Quercus-Juniperus and Pinus-Quercus associations are present; most com- 
mon oak species are Quercus chihuahuensis Trel,, Q. grisea Liebm. and Q. emoryi Torr, and 
in the lower hills and plains, where the woodlands contact the grasslands, Juniperus 
monosperma (Engelm.) Sarg.is more frequent, often forming savanaah. Higher and cooler 
areas of the mountains have Quercus hypoleucoides A.Camus, Q. arizonica Sarg,,Q. rugosa 
Nee, Q. grisea Liebm,, Q. aeplessipes Trel,, Pinus cembroides Zucc., P engelmanii Carr, P. 
chihuahuana Engelm.and J Steud. Southeast of Cd. Chihuahua Prosopis 
glandulosa Torr. var. onevana (L. Bence) IM. Johnst., forms dense shrubby areas 
(mezquital), especially along roadsides and disturbed areas from Cd. Chihuahua to 
Estacién Horcasitas, 25 km SW form Cd. Chihuahua (Andnimo 1981). Pressed and dried 
vouchers are stored in the herbarium CFNL; incomplete sets are deposited at ANSM, 
BRIT, MEXU, NMC and TEX (Holmgren & Holmgren 1990). 


METHODS 


During 1994-1998 numerous routes through all plant communities of the central part 
of the state of Chihuahua were sampled. On each site where legumes were collected, 
main vegetation type, geographic coordinates, nearest towns, altitude and slope were 
recorded. Nine hundred and fifty samples of legumes were collected in this study. 


DISCUSSION 


Only a partial geographical representation of legumes exist for Mexico. Regional studies 
have in particular contributed to the knowledge of these plants. Of particular merit are, 
for southern Mexico, Standley and Steyermark (1946),and Woodson et al. (1980); for cen- 
tral, Matuda (1980), and Rzedowski and Rzedowski (1979);for southwest, McVaugh (1987); 
for northeastern, Correll and Johnston (1970), Isely (1981), Estrada and Marroquin (1991), 
Carranza and Villarreal (1997), and Ramos (1999); for northwestern, Munz (1959), Kearney 
and Peebles (1960), Shreve and Wiggins (1964), Isely (1981), Spellenberg et al. (1996), and 
Estrada et al.(1997). These studies show distribution, diversity, and ecology of legumes in 
this country, while this research focuses on additional knowledge on presence, distribu- 
tion and ecological aspects of the legumes in northern Mexico. 

In this study, 42 genera, 117 species and 36 varieties of legumes from the central 
part of the state of Chihuahua were recorded. Lotoideae has the highest number of 
genera and species with 27 and 77 respectively, followed by Mimosoideae with 8 and 28 
and Caesalpinioideae, with 7 and 12. Appendix 1 shows the taxa by family, and vegeta- 
tion type(s) where they were more frequently found. Dalea, Acacia, Desmodium and 
Phaseolus are the most diverse genera, each with 5 or more species. These genera have 
species in the three main plant communities. Dalea is the genus with more species (19), 
nine of which occur in pine-oak forest, eight are dominant in grasslands, while six are 


354 SIDA 19(2) 


typical of matorral. The eleven species of Acacia mainly occur in plains and low sierras 
under 1700 m elevation, except for A. angustissima that grows in mountainous areas 
mainly associated to Bouteloua grasslands. Acacia constricta and A. noevernicosa are the 
dominant species in matorral communities. Five out of seven Phaseolus species are ex- 
clusive to pine oak-forest in the cooler parts of the region (Sierra El Nido and Majalca), 
above 1800 m elevation. All of Desmodium species occur in oak or pine forest, only D. 
neomexicanum occurs in the three dominant communities. None of the Astragalus spe- 
cies occurs in forest areas, while all of them occur in grassland communities. No species 
of Caesalpinoideae occur in pine or oak forest, the five species of Senna were seldom 
found in low numbers in matorral and grassland. Most Mimosa species were often asso- 
ciated with oak forests, in particular those formed by Quercus emoryi and Q. grisea, and 
matorral. In both communities, M. aculeaticarpa is the most abundant. 

Highest number of legume species were recorded on oak-pine forests (33) and 
Bouteloua grasslands (31). Oak Forest and Pine Forest, each have a similar number of 
legume species, 21 and 20 respectively.In shrublands where Acacia is dominant, 25 spe- 
cies of legumes were recorded, three times the number of species recorded in Larrea 
and Flourensia shrubland.Within different grassland association, the Sf | Hilaria 
types are areas with lowest legumes, with 9 and 6 species respectively, although they 
shelter some characteristic species from these plant communities such as Hoffmanseggia 
spp, Astragalus spp. and Peteria scoparia. Several species are found mainly in disturbed 
and overgrazed areas, they are Acacia farnesiana, Mimosa aculeaticarpa var. biuncifera, 
Prosopis glandulosa var. torreyana, Acacia schaffneri var. bravoensis, Crotalaria pumila, 
Chamaecrista nictitans and Senna lindheimeriana; the last three are found as roadside 
weeds. Prosopis glandulosa var. torreyana is one of the main species associated with me- 
dian grassland (Bouteloua spp.) in central plains, especially in the area between Chihua- 
hua-El Sueco and Chihuahua-Estacién Horcasitas. Acacia farnesiana occurs throughout 
Mexico (Clarke at al. 1989), except for Chihuahua. In this study we found it occasionally 
in abandoned agricultural lands, close to human settlements and next to roads, around 
Cd.Aldama. 

Eight legume species (7.5%) are introduced, Caesalpinia gilliesii, Parkinsonia aculeata, 
Alhizin jialihriccin di, fi | If LABIAL Pahin} -l Hy 


f | 


+; AA 


and Wisteria sineneis, the latter three are cultivated as ornamental plants. Eighty eight species 
are native of northern Mexico, some of them occur in southern U.S.From the approximately 
135 genera and 1724 legume species present in Mexico (Sousa & Delgado 1993), 31.1% and 
7.1% of them respectively are present in the study area. Grasslands and shrublands cover 
95% of the studied area, but they have lower legume diversity than oak-pine forest, which 
represent only 5% of the surface. Total taxa recorded are distributed in a 7500 km? surface 
(lower than a tenth part of the state territory) on three main vegetal communities from the 
seven recognized for the state of Chihuahua (Andnimo 1978).A very different and heteroge- 
neous relief, altitude, climate and vegetation association on the western part of the state of 
Chihuahua, reveal a constant legume species diversity, especially on cooler 


ESTRADA C. AND MARTINEZ M., LEGUMES OF CENTRAL CHIHUAHUA, MEXICO 355 


oak, pine, oak-pine forests and subtropical shrublands. Spellenberg et al. (1996) list 30 
genera and 65 legume species from Parque Nacional Cascada de Basaseachi; Estrada et 
al. (1997) list 21 genera and 53 species in Babicora Lake. Table 1 shows the affinities of 
legumes in the three studied areas. The most oustanding contrast in the areas is the 
presence of 14 genera in central Chinuhahua, absent from Basaseachi and Babicora 
(Desmanthus, Peteria, Painteria, Albizia, Prosopis, Zapoteca, Hoffmanseggia, Pomaria, 
Parkinsonia, Lablab, Melilotus, Nissolia, Pediomelum, Robinia, and Wisteria),and seven gen- 
era in Basaseachi, absent from Babicora and Central Chihuahua, these are Conzattia, 
Pithecellobium, Lysiloma, typical of moist warm environments and Erythrina, Lathyrus, 
Marina and Minkelersia. In Babicora no Caesalpinioideae occurs, while in Basaseachi, three 
species were found, all of which had tropical affinity. Dalea is conspicuously more abun- 
dant in the center of the state than in the other two areas, fourteen of the central species 
are absent in Babicora and Basaseachi.All of these are from semiarid climates. Desmodium 
is more diverse in Basaseachi, seven of its species, that are absent in the other two areas, 
are more common in moist environments. Perhaps the greater species diversity of le- 
gumes found for the central part of the state reflects the variation on climate, soil and 
topography of this region. These three factors are in sharp contrast in plains, sierras and 
mountains, and thus affect the distribution and abundance of different taxa. 


Taste 1.Subfamilies, genera and species of legumes in three regions of the state of Chihuahua. 


Basaseachi Babicora Central 
spellenbera etal.(1996) | Estradaetal.(1997) Chihuahua 
Subfamilies 3 2 3 
Total genera 30 22 42 
Total epee 65 53 117 
Genera/spe 
Ca S ininiodeae 3/3 0 7/12 


Genera/species 
Mimosoideae 4/6 3/4 8/28 


Genera/species 
otoideae 
Genera exclusive to 


the region 9 0 15 
Genera with m 


Phaseolus species 


356 


SIDA 19(2) 


APPENDIX 1 


Subfamilies, genera and species of legumes are recorded for the central part of the state 
of Chihuahua and plant communities where most frequently found. HG, Hilaria Grass- 
land; SG, Sporobolus grassland; BG, Bouteloua grassland; SL, Shrubland (Larrea as domi- 
nant); SF, Shrubland (Flourensia as dominant); $A, Shrubland (Acacia as dominant); SP, 
Shrubland (Parthenium as dominant); OF, Oak forest; OPF, Oak-Pine forest; PF, Pine Forest: 


D, Disturbed Areas; C, Cultivated. 


MIMOSOIDEAE 


Acacia angustissima (P. eer Kuntze. var. 
chisosiana |sely [SA,$ 
Acacia angustissima (P. me Ktze. var. texensis 
(T.& G,) Isely [SA, 
Acacia berlandieri Benth, [ SA] 
Acacia biaciculata S.Wats. [BG, HG, SG, SA] 
SA 


Acacia glandulifera S.Wats. [SA, SF 
a greggii A. Gray var. arizonica Gray [BG, SA, 


Pees greggii Gray var. greggii [SA, SP] 

Acacia neovernicosa lsely [SA, SF, SP, SL] 

Acacia roemeriana Scheele [SA] 

Acacia schaffneri (Wats.) Herm. var. bravoensis 
lsely [D, SA 

Acacia a wrightif B enth. [SA] 

Albizia julibrissin Durazz. 

Calliandra conferta Gray [SA, SL, SP, BG] 

Calliandra eriophylla Benth. var. oak [SA, 


ga 


Calliandra humilis Benth. var. reticulata (A. Gray) 
.Benson [OF, OPF 
Calliandra humilis Benth. var. humilis [OF, OPF] 
Desmanthus cooleyi (Eat.) Trel. (BG, SA, SL, OF] 
Desmanthus. virgatus (L.) Willd. [BG, SA, SF] 
Mimosa aculeaticarpa oe var. biuncifera (Benth. 
Bar men [D, SA,O 
Mj pa ea [BG, OF, OPF] 
Mimosa emoryana Benth. var. chihuahuana (B. & 
R.) Barneby [OF, SA] 
Mimosa emoryana Benth. var.emoryana [OF, SA] 
Mimosa pringlei Wats. var. pringlei (SA, OF] 
Mimosa texana (Gray) Small var. texana [SA, SF] 
Painteria elachistonhvila (W. Rr 2 PR 


ae 


[SA, SL] 
DC) Britton &R (BG, OPF 
Prosopis glandulosa Tor var. torreyana (L.Benson) 
t.[BG, SG, SA SL, SF] 
Prosopis Peece (H. & B. ex Willd.) M.C. Johnst. 


Prosopis pubescens Benth. [SL] 
Zapoteca media (Mart. & Gal.) H.M. Hern. [OF] 


CAESALPINIOIDEAE 


Chamaecrista nictitans (L.) Moench ssp. nictitans 
var. mensalis (Greenm.) Irwin & Barneby [BG, 


Caesalpinia gilliesi (Hook.) Benth. [SG, C] 

Hoffmanseggia glauca (Ort.) Eifert. [BG, HG, SG] 

Parkinsonia aculeata L.[D 

Pomaria jamesii (T.& G.) Walp. [HG, GS, BG] 

Pomaria multijuga (S. naa, B.B. Simpson [SA] 
(cited by Simpson 

Senna bauhinioides co Irwin & Barneby (HG, 
SG, B BG, SL, SF, SA ) 


le) | in ®& Rarn by 
} 


D, BG] 
Senna pilosior (Macbride) Irwin & Barneby [SP 
SL] 


Senna ripleyana (Irwin & Barneby) Irwin & 
Barneby [SL, SF, BG] 

Senna wislizeni (Gray) Irwin & Barneby var. 
wislizeni [SL, SF, SA] 


LOTOIDEAE 


Rescnymomene fasciculans een (SA, BG] 
ar. playanus (Jones) 


Isely [BG, D] 

Astragalus mollissimus Torr. Ma earlei (Greene ex 
Rydb.) Tidest. [SG, HG 

Astragalus mollissimus Le var. irolanus (Jones) 
Barneby [BG. SG] 

— nuttalianus A.DC. var.austrinus (Small) 
Barneby [BG, HG, SA 

Astragalus pringlei Wats. [BG, D, SA] 

Astragalus quinqueflorus S.Wats. [BG, HG] 

Astragalus wootonii Sheld. [D, BG, SA, SF] 

Cologania angustifolia H. : K. [OPF, OPF] 

Cologania Schlecht. [PF, PF 

oT caribaea (Jacq.) an var. sericea ( Gray) 


is 


Lavin [OF, B 
en caribaea (Jacq.) Lavin var. caribaea [OF] 
couse sas a (Gray) Lavin [OPF, PF] 


talaria pumila Ort. [D, SA] 
. brach aie Gray [D, SL, SA] 
Dalea filiformis Gray [PF, OPF 


ESTRADA C. AND MARTINEZ M., LEGUMES OF CENTRAL CHIHUAHUA, MEXICO 357 


Dalea foliolosa (Ait.) Barneby var. foliolosa [OPF, 


PF] 
Dalea formosa Torr. [SL, SA, SF, SP] 
Dalea grayi (Vail) LO Williams [OPF, OF, PF] 
Dalea humilis G. Don. (PF, OPF 
Dalea jamesii (Torrey) T. & G. [BG] 
Dalea lachnostachya Gray [SL, SA] 
Dalea leporina (Aiton) Bullock [BG, OPF, OF] 
Dalea leucostachya Wats. var. leucostachya [OF, 
OPF] 


Dalea lutea (Cav.) a var. lutea [SA, OPF] 

Dalea mollis Benth. [BG] 

Dalea nanaTorr.var.carnescens (Rydb.) K.& P.[BG 
SA 


Dalea neomexicana (Gray) Cory var.neomexicana 
[BG] 

Dalea pogonathera Gray var. pogonathera [BG, 
SA, SF, SP, SG, HG] 

Dalea prostrata Ortega [BG] 

Dalea versico Hs ans var. glabrescens (Rydb.) 
Barneby [O 

Dalea hee ee var. sessilis (Rydb. 


YS 


Barneby 


Dalea viridiflora Wats. [OPF] 
Bae Hg Gley BG 


lium DC. [OF 
Desmodium batocaulon Gray [OPF, PF] 
Desmodium hartwegianum Hemsl. var. 


hartwegianum [PF, OPF 
Desmodium neomexicanum OE [SA, OF, BG] 
cee G 
modium retinens Schlecht. TOPF 
Ssenhrdia, aan asl [SA, SP] 
lactia macreii M.A. Curtis [OF] 


al 


actia wrightii Gray ish | 

ladigeera montana Rose [OPF, PF] 

ees ornithopodioides Schlecht. & Cham. 
, OF 


[SA 
Lablab purpureus (L.) Sweet [C] 


Lot nei (Woot.& Stand.) Ottley ex Kearn 
‘g Peebles [OPF, PF] 

Lotus oroboides (H.B.K.) Ottley ex Kearney & 
Peebles [OF, OPF] 

Lotus plebeius (Brandeg.) Barneby [PF] 

Lupinus aff. delicatulus sprag ue & Riley [PF] 

upinus t i Schlecht. [OF, OPF] 
Macroptilium gibbosifolium (Ort.) A. Delgado 
G, D, 

Medicago sativa L. [D, BG] 

Medicago lupulina L. [BG] 

Melilotus indicus (L.) All. [D] 

Melilotus officinalis L. [D] 

Nissolia pringlei Rose (SF, SA, SL, D] 

Nissolia wislizeni (Gray) Gray [SF, BG] 

Pediomelum palmeri (Ock.) Grimes [SG, BG] 

Peteria scoparia Gray [BG, SG, HG] 


Phaseolus grayanus Woot.& Standl.[OF, OPF, PF] 

Phaseolus leptostachyus Benth. [OF, OPF, PF] 

Phaseolus parvulus Greene [OF, OPF, PF] 

Phaseolus ritensis ae [OPF, OF] 

Phaseolus vulgaris L. [D] 

Rhynchosia Benth. [SA, OF, OPF] 

Rhynchosia senna Gill ex Hook. var. angustifolia 
(Gray) Grear [D, SA] 

Robinia pseudoacacia C) 

Sophora gypsophila B.L. Turner & Powel [SG, SA, 
SF 


ieoiesian menENG ny [SA] 

e H.B.K. var. amabile [OPF, PF] 

m Michx. [OPF, PF] 

Trifolium wormskioldii Lehm. var. ortegae 
(Greene) ee [OPF, PF] 

Vicia ludoviciana Nutt. ssp. fudoviciana [OPF, PF] 

Vicia pulchella H.B.K. ssp. pulchella [OPF, PF] 

Wisteria sinensis (Sims.) Sweet [D, C] 

Zornia gemella (Willd.) Vog. [SA, OF, OPF] 

Zornia reticulata Sm. (OF, OPF] 


Trifoali linin 


ACKNOWLEDGMENTS 


We thank Alfonso Delgado and Fortunato Garza for comments on the manuscript and 
Richard Spellenberg, Emily Lott, Barney Lipscomb and Enrique Jurado for reviewing the 
English translation and critical reviews. Miguel Angel Gonzalez provided the map. 


REFERENCES 


ANONIMO. 1978. ComisiOn técnico consultiva para la determinacidn de los coeficientes de 
agostadero. Gufas para la determinaci6én de condicién. Secretaria de Agricultura y 


Recursos Hidrdulicos. 


358 SIDA 19(2) 


ANONIMO, 1981. Carta de uso de suelo y vegetacidn, escala 1: 1 000 O00. Secretaria de 
Programacion y Presupuesto. México, D.F 

ANONIMO, 1981. Carta geoldgica, escala 1: 1 000 O00. Secretaria de Programacién y 
Presupuesto. México, D.F. 

ANONIMO. 1987. Mapa 1.2, Fisiografia, escala 1:2 000 000. Instituto Nacional de Estadistica 
Geografia e Informatica, México, D.F. 

Acosta, J.A.,J.M.Martinez y F. CARDENAS. 1983. Distribucion de las especies del género Phaseo- 
lus en el estado de Durango. Agric. Tec. Méx. 9 (1):13-22. 

Barnesy, R.C. 1964. Atlas of North American Astragalus, part I-ll, Mem. New York. Bot. Gard. 
13:1-1188. 

Barnesy, R.C. 1977. Daleae Imagines. Mem. New York Bot. Gard. 27:1-879. 

Barnesy,R.C. 1989. Intermountain flora. Fabales Vol 3 Part B. New York Bot.Gard.Pp. 11-279. 

Barnesy, R.C. 1991. Sensitivae censitae. Mimosa. Mem. New York Bot. Gard. 65:1-835. 

Barnesy, R. C. and J. Grimes. 1996. Silk tree, Monkey’s earring: A generic system for the 
synandrous Mimosaceae of the Americas. Part |. Abarema, Albizia and allies. Mem. New 
York Bot. Gard. 74:1-292. 

Barnesy, R.C. and J. Grimes. 1997. Silk tree, Monkey’s earring: A generic system for the 
synandrous Mimosaceae of the Americas. Part Il. Pithecellobium, Cojoba and Zygia. Mem 
New York Bot. Gard. 74:1-149. 

Barnesy, R.C. and J. Grimes. 1998. Silk tree, Monkey’s earring: A generic system for the 
synandrous Mimosaceae of the Americas. Part Ill. Calliandra. Mem. New York Bot. Gard. 
74:1-223. 

Burkart, A.1971.El género Galactia (Leguminosae-Phaseoleae) en Sudamérica con espe- 
cial referencia a al Argentina y paises vecinos Darwiniana 16:663-796, 

Burkart.A. 1976.A monograph of the genus Prosopis (Leguminosae, subfam. Mimosoideae) 
J.Arnold Arbor, 57:219-249, 

Carranza, M.A.&J.A.Villarreal.1997.Legumin de Coahuila, México.U.A.A.A.N. Buenavista, 
Saltillo, Coah., México. 

Ciarke, D.and D.S. Seicter, and J.E. Esincer. 1989.Acacia farnesiana (Fabaceae—Mimosoideae) 
and related species from Mexico, the southwestern U.S. and the Caribbean. Syst. Bot 
14:549-564. 

Carke, D., D.S. Seicter, and J.E. Egincer. 1990. Acacia constricta (Fabaceae: Mimosoideae) and 
related species from the southwestern U.S. and Mexico. Amer. J. Bot. 77:305-315. 

Corrett, D.S. and M.C. JoHNsTON. 1979, Manual of the vascular plants of Texas. Univ. Texas, 
Dallas. Pp. 761-889. 

Cronaquist, A. 1988. The evolution and classification of flowering plants, 2nd.ed. New York 
Botanical Garden. Pp. 1-555. 

Euas, 1.S.1974. The genera of Mimosoideae (Leguminosae) in the southern United States. 
J.Arnold Arbor. 55:67-113. 

Garcia, E. 1973. Modificaciones al sistema de clasificacidn climatica de Koeppen. 3a ed. 
Instituto de Geografia. Universidad Nacional Auténoma de México. D.F. 


ESTRADA C. AND MARTINEZ M., LEGUMES OF CENTRAL CHIHUAHUA, MEXICO 359 


Grear, J. W. 1978. A revision of the New World species of Rhynchosia (Leguminosae - 
Faboideae). Mem. New York Bot. Gard. 31:1-168 

Grimes, J. 1990. A reivision of the New World species of Psoraleeae (Leguminosae: 
Papilionoideae). Mem.N.Y. Bot. Gard. 61:1-114. 

Gunn, C.R. 1979. Genus Vicia with notes about tribe Vicieae (Fabaceae) in Mexico and 
Central America. U.S.D.A. Tech. Bull. No. 1601. 

HERNANDEZ, H. 1986. Zapoteca: A new genus of noetropical Mimosoideae. Ann. Missouri. 
Bot. Gard. 73:755-763. 

HERNANDEZ, H. 1989.Systematics of Zapoteca (Legumin ) Ann.Missouri. Bot. Gard. 76:781- 
862. 

Houmoren, PC. and N.H. Houmeren. 1990. Index herbariorum, Ed.8. Part 1.New York Botanical 
Garden. 

Irwin, H.S and R.C. Barneby. 1982. The American Cassiinae. A synoptical revision of 
leguminosae tribe Cassiieae subtribe Cassiinae in the New World.Mem. New York. Bot. 
Gard. 35:1-918. 

Isety, D. 1969. Legumes of the United States: |: Native Acacia. Sida 3:365-386. 

Isecy, D. 1973. Leguminosae of the United States: |. Subfamily Mimosoideae. Mem. New 
York. Bot. Gard. 25:1-152. 

Isety, D. and F.J. Peabody. 1984. Robinia (Leguminosae: Papilionoideae): Castanea 49:187—- 
202. 

Isety,D.1975.Legumi f the United States: Il. Subfamily Caesalpinioideae. Mem. New 
York Bot. Gard. 25:1-228. 

Isety, D. 1981. Leguminosae of the United States. Ill. Subfamiliy Papilionoideae: Tribes 
Sophoreae, Podalyrieae, Loteae. Mem. New York Bot. Gard. 25:1-264. 

JoHNsTON, M.C.1962.The North American mesquites, Prosopis Sect. Algarobia (Leguminosae). 
Brittonia 14:72-90. 

Kearney, T.and R.H. Peebles. 1960. Arizona flora, 2nd. ed. Univ. Calif. Press. 

Lavin, M. 1988. Systematic of Coursetia (Leguminosae-Papilionoideae). Syst. Bot. Monogr. 
21:1-167. 

Lesins, K.A.and |. Lesins. 1979.Genus Medicago (Leguminosae).A taxogenetic study.W. Junk 


The Hague-Boston-London. 

Luckow, M. 1993. Monograph of Desmanthus (Leguminosae—Mimosoideae). Syst. Bot. 
Monogr. 38:1-165. 

McVauch, R. 1987. Flora Novo-Galiciana. Vol. 5. Leguminosae 

MarecHal, R.J.,J.M.MascHerea, and F. Stainier. 1978. Etude taxonomique d’un gruope complexe 
dé espéces des generes Phaseolus et Vigna (Papilionaceae) sur la base de donn’ees 
morphologiques et polliniques, traitées par lanalyse informatique. Boissiera 28:1-273. 

MoHtensrock, R.H. 1961.A monograph of the leguminous genus Zornia.Webbia 16:1-141. 

MoHLENBROCK, R.H. 1962. Tribe Hedysareae, subtribe Stylosanthinae (Leguminosae) of Cen- 
tral America and Mexico. SouthW. Nat. 7:29-40. 

Munz, PA. 1959. A California flora. University of California Press. Pp. 795-897. 


360 SIDA 19(2) 


Orttey, A.M. 1944. The American Loti with special consideration of a proposed new sec- 
tion, Simpeteria. Brittonia 5:81-123. 

Piper, B.C. 1926. Studies in American Phaseolineae. Contr. U.S. Natl. Herb. 22:663-701. 

Porter, C. L. 1956. The genus Peteria (Leguminosae). Rhodora 58:344—354, Robertson, K.R. 
and Yin-Tse Lee. 1975. The genera of Caesalpinioideae (Leguminosae) in the South- 
eastern United States. J. Arnold Arbor. 57:1-53. 

Ramos, M.C. 1999. Las Leguminosas de la Reserva El Cielo, G6mez Farias, Tamaulipas. Tesis. 
Instituto Tecnolégico de Cd. Victoria, Tamaulipas. 

Rupp, V.E.1956.A revision of the genus Nissolia. Contr. U.S. Natl. Herb. 32:173-206. 

Ruop,V.E. 1968. Leg! f Mexico. Faboideae |. Sophoreae and Podalarieae. Rhodora 
70:493-532. 

Ryosera, P.A. 1919. Fabaceae. In: North Amer. Fl. 24(1):1-250. 

Rzepowski, J. 1978. Vegetacidn de México. Ed. Limusa. México, D.F. 

RzeDowsk\, J. y G.C. be Rzepowski. 1979, Flora fanerogamica del Valle de México. CECSA. Pp. 
279-354 


Rzevowski, J. 1988. Analisis de la distribucidn geografica del complejo Prosopis 
(Leguminosae, Mimosoideae) en Norteamérica. Acta Bot. Mex, 3:7-19 

Senn, H.A. 1939. North American species of Crotalaria. Contr. Gray Herb. 125:31 7-370. 

Sureve, F.and |.L.Wiccins. Vegetation and flora of the Sonoran Desert. Pp. 585-731. 

SHuseRt, B.G. 1940, |. Desmodium: Preliminary studies |. Contr. Gray Herb. 129:1-31. 

SHuBERT, B.G. 1941.11. D dium: Preliminary studies Il. Contr. Gray Herb. 136:78-114. 

Simpson, B.B. 1998. A revision of Pomaria (Fabaceae) in North America. Lundellia 1:47-71. 

SMITH, CP. 1938-1953. Species Lupinorum 1-768. en 44 fasciculos, publicado por el autor. 

Sousa, M. and A. Detcapo. 1993. Mexican Leguminosae: Phytogeography, endemism and 
origins In: Biological diversity of Mexico: Origins and distribution. Oxford Univ. Press. 
Pp.459-507 

SPELLENBERG, R., T. Lesaue y R. Corrat. 1996. Listados floristicos de México. XIll. A specimen- 
based annotated checklist of the vascular plants of Parque Nacional “Cascada de 
Basaseachi" and adjacent areas. Instituto de Biolog{fa. Universidad Nacional Aut6énoma 
de México. 

Stanbey, P. C. 1922. Leguminosae. In: Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 


STEVENSON, G.A. 1969. An agronomic and taxonomic review of the genus Melilotus Mill. 
Canad. J. Pl. Sci. 49:1-20. 

Turner, B.L. 1950. Texas species of Desmanthus. Field & Lab. 18:54-65. 

Turner, B.L. 1950.Mexican species of Desmanthus (Leguminosae). Field & Lab. 18:119-130. 

Turner, B.L. and A.M. Powett. 1972. A new gypsophilic Sophora (Leguminosae) from 
Northcentral Mexico and adjacent Texas. Phytologia 22:419-423. 

Turner, B.L. 1992. Taxonomic overview of the genus Cologania (Fabaceae, Phaseoleae). 
Phytologia 73:281-301. 

Zonary, M. and D. Heiter. 1984. The genus Trifolium. The Israel Acad. Sci. Hum., Jerusalem 


A VASCULAR FLORA SURVEY OF 
CALCASIEU PARISH, LOUISIANA 


Ray Neyland and Billie J. Hoffman 


secre ac & Envi | Sciences 
se State University 
i sens LA 70609, U.S.A. 


Mark Mayfield and Lowell E. Urbatsch 


Department of Plant Biology 
Louisiana State University 
Baton Rouge, LA, 70803, U.S.A. 


ABSTRACT 


Located in southwestern Louisiana, Calcasieu Parish is composed of five major vegetation regions. 
These regions are defined by their respective vascular plant composition and physiography.A field 
survey of the vascular flora of the approximately 2,844 square kilometers that make up the Parish 
was conducted from August 1995 to October 2000. Additionally, specimens housed in various Loui- 
siana herbaria d to complete the survey. Names of the 1,147 specific and subspecific 
entries representing 147 families found during this survey have been compiled in an annotated 
catalogue that includes collector name and collection number for one representative specimen 
anda oe as to whether each species is considered native or introduced. Vegetation re- 
gions where each species occurs are included in the catalogue. An additional 134 taxa reported! 
collected in oa Parish that were not discovered during our field research nor located during 
our visits to other herbaria are also listed. Euphorbia texana is reported new for Louisiana. 


Key Worpbs: Vascular Plant Survey, Calcasieu Parish, Louisiana 


RESUMEN 


Localizad | te de Louisi | ia de Calcasieu esta compuesta por cinco regiones 
de vegetacion principales. ie regiones se definen por s Su pill. Ht y su fisiografia. 
Una prospeccién d p flora vascular | 4 kildmetros cuadrados 
| li d to de 1995 a octubre fe 2000. Adicionalmente, 
especimenes albergados en varios herbarios de ‘Louisiana fueron examinados para completar el 
estudio. Los nombres de las 1,147 entradas especificas y subespecificas que representan las 147 
familias encontradas durante esta prospeccién se han compliado en catalogo comentado que 
incluye nombre del colector y nel de la recoleccién de un espécimen respresentativo y una 
esignacién de cada esp idera nativa o introducida. Se incluyen en el catalogo 
las regiones de vegetacién en que vive cada especie. Se listan también 134 taxa adicionales citados 
como colectados en la parroquia de Calcasieu que no se encontraron durante la investigacién de 
campo ni en las visitas o otros herbarios. Euphorbia texana se cita como nueva para Louisiana. 


Located in southwestern Louisiana, Calcasieu Parish lies adjacent to Beauregard Parish 
to the north, Jefferson Davis Parish to the east, Cameron Parish to the south and the 
State of Texas to the west (Fig. 1).Summoers in the Parish are hot and humid; winters are 
warm but are occasionally interrupted by freezing temperatures (Hardner 1960; Roy & 


SIDA 19(2): 361 — 386. 2000 


362 SIDA 19(2) 


Ae AA A AA A wR ATENEE, A A A A a aA A a an A A A A Aw 


[ARKANSAS (. Coastal Marsh 


| 
| ‘ LOUISIANA Bottomland Forests 


SISSIPPI 
vou) ino Coastal Prairie 
BA heel 
NY 
| ae Ge Swampland BRARA 
uur a 
— Longleat Pineland [| 


kK + L om 1 e L +h iT “as Lek om od PR L 
Fic. 1. The major Louisiana p 


State. 


be J J 


Midkiff 1988). Annual precipitation averages 135.1 cm (Roy & Midkiff 1988). The Parish is 
occasionally impacted by hurricanes. 

Elevation in Calcasieu Parish ranges from sea level to 29 meters (Roy & Midkiff 1988). 
The Calcasieu and Houston rivers drain the central and eastern portions of the Parish, 
and the Sabine River (Hardner 1960) drains the western portion. The Sabine Diversion 
Channel runs west to east near the center of the Parish and diverts water from the Sabine 
into the Calcasieu River. The Intracoastal Waterway is situated in the extreme southern 
part of the Parish and runs from Calcasieu Lake to the Sabine River. Barge traffic is exten- 
sive in the Intracoastal and to a point north of Lake Charles on the Calcasieu River Jones 
et al. 1954; Roy & Midkiff 1988).A deep-water port is located in Lake Charles Uiones et al. 
1954). 

Calcasieu Parish lies in the West Gulf Coastal Plain geomorphic province (Fenneman 
1938). Surface sediments are of Recent and Pleistocene es and are underlain by 


NEYLAND ET AL., FLORA OF LOUISIANA 463 


Tertiary rocks (Hardner 1960; Roy & Midkiff 1988). The recent deposits generally consist 
of fine sand, silt and clay and occur along the southern edge of the parish and in the 
Sabine and Calcasieu river valleys (Hardner 1960). The Pleistocene deposits consist of 
stream-transported sediment washed down during the past four glaciation periods 
(Hardner 1960). 

Approximately 46% of the Parish’s land is devoted to either agriculture or range- 
land; approximately 23% is woodland; 11% is marsh, 4% swamp and the remaining area 
is urban (Roy & Midkiff 1988). Land use is primarily devoted to producing timber, raising 
cattle, and growing rice and soybeans.Large petroleum-related industries are present in 
the Parish and are concentrated around the area of West Lake. 

The purpose of this study is to survey Calcasieu Parish for all vascular plants that are 
native or introductions that appear to be naturalized and to define and map the vegeta- 
tion regions the Parish. Cultivars and introductions that do not appear to be naturalized 
are excluded from the survey. 


METHODS 


Approximately 2,000 plant collections from Calcasieu Parish were made between Au- 
gust, 1995 and October 2000. Voucher specimens were prepared using standard her- 
barium practices and are housed in the McNeese State University (MCN) herbarium. Speci- 
mens from the following Louisiana herbaria were examined to complete this survey. 
These herbaria include: University of Louisiana at Lafayette (LAF); Louisiana State Univer- 
sity (LSU), University of Louisiana at Monroe (NLU) and Tulane University (NO). 

Specimens were identified primarily through the use of the following references: 
Allen (1992); Correll & Correll (1941); Correll & Jonnston (1970); Cronquist (1980); Duncan 
(1975): FNA (1993, 1997); Godfrey (1988); Godfrey & Wooten (1979, 1981); Gould (1975); 
Radford et al. (1968) and Small (1933). For divisions Filicophyta (ferns), Lycopodiophyta 
(lycopods), Equisetophyta (horestails), and Coniferophyta (conifers), nomenclature fol- 
lows FNA (1993). For Magnoliophyta (angiosperms) subclasses Magnoliidae and 
Hamamelidae, nomenclature follows Ee (1997). For all remaining subclasses, nomen- 
clature follows Kartesz (1999).A pl rts were consulted to help identify prob- 
lematic taxa. These experts imetuele lanatles Allen, (Poaceae; NLU); Doug Goldman 
(Calopogon); John Hays (Agalinis; University of Louisiana, Monroe); Phil Hyatt (Carex; United 
States Forestry Service, Pineville, LA); David Moore (various rare taxa; United States For- 
estry Service, Pineville, LA); Guy Nesom (Asteraceae; North Carolina Botanical Garden); 
Latimore Smith (various rare taxa; National Heritage Program, Louisiana Department of 
Wildlife & Fisheries, Baton Rouge, LA). 

From an analysis of plant collection data and recent soil survey maps (Roy & Midkiff 
1988), vegetation regions within Calcasieu Parish were defined and mapped. 


RESULTS 


Collections made by the authors form the majority of the annotated list of vascular plants 
for Calcasieu Parish (Appendix |). The list includes 1,147 specific and subspecific entries 


oa 


364 SIDA 19(2) 


representing 147 families and 530 genera. Twenty-nine ferns, lycopods and horsetails 
account for about 3% of the Calcasieu Parish flora and six gymnosperms account for less 
than 1%.The remaining flora consists of angiosperms. About 15% of the Calcasieu Parish 
flora are introductions. 

One-hundred and thirty-four taxa reportedly collected in Calcasieu Parish 
(MacRoberts 1988; Thomas & Allen 1993, 1996, 1998) that were not discovered in field 
research nor located during our visits to other herbaria are listed in Appendix II. Likely 
reasons why we were unable to locate these specimens were that they were: 1) on loan; 
2) misfiled; 3) overlooked; 4) resident in herbaria not visited. When both confirmed and 
unconfirmed lists are added, the total number of native and naturalized taxa attribut- 
able to Calcasieu Parish is 1,281. 

Three-hundred and two species, not previously recorded in the Parish, were discov- 
ered during this study. Euphorbia texana Boiss. is a new record for Louisiana. Five veg- 
etation regions were identified and mapped for the Parish (Fig. 1 


—— 


DISCUSSION 


Calcasieu Parish is noteworthy for the extreme transition from pinelands in the north to 
prairie and brackish marsh in the south. Bottomland hardwoods and swamps are associ- 
ated with the Calcasieu, Sabine and Houston Rivers. Five vegetation regions in Calcasieu 
Parish are recognized (Fig. 1) and are discussed in terms of soil characteristics, current 
uses,and dominant species composition. 

Coastal Prairie —This region is characterized by soil that ranges from loamy to clayey, 
is medium in fertility and is used primarily for urban development, cropland and 
pastureland (Roy & Midkiff 1988). The main crops are soybeans and rice (Roy & Midkiff 
1988). Most of the rice farming is done on the prairie because of its low altitude, low 
relief and the impervious nature of its subsoil Jones et al. 1954). Extensive farming and 
urbanization has heavily impacted the prairie region. Dominant trees in the better-drained 
soils of higher elevations include Liquidambar styraciflua, Pinus elliottii, Pinus taeda, Quercus 
alba,and Nyssa sylvatica. Trees in poorly drained soils of lower elevations include Quercus 
nigra, Quercus phellos and Platanus occidentalis. Herbaceous plants typically are weedy 
and include many species of grasses and sedges. 

Coastal Marsh.—This region is very low in relief and rarely rises more than 1.5 meters 
(Jones et al. 1954). Soils in the coastal marsh region range from soft organic to firm min- 
eral clay. These soils are poorly drained, subject to flooding and used primarily for wildlife 
habitat, recreation and rangeland (Roy & Midkiff 1988). The coastal marsh region is in- 
habited mostly by grasses and sedges, and is typically devoid of trees Jones et al. 1954); 
however Sapium sebiferum and Zanthoxylum clava-herculis are common along roadsides. 
Marshes are either fresh or brackish. In general, marshes become more saline the closer 
they lie to the coast. 

Approximately 9,906 hectares of brackish marsh occur in the Parish. Salinity ranges 
from 0.42 to 28.08 parts per thousand (ppt) with a mean of 8 ppt (Roy & Midkiff 1988). 


NEYLAND ET AL., FLORA OF , LOUISIANA 365 


Marsh associated with the Calcasieu River may be saline up to a point just north of Lake 
mail: ee the river is dammed.Common brackish-marsh herbaceous plants include: 

ie niflora, Distichlis spicata, Juncus sp.and Scirpus sp.Approximately 7,020 hect- 
ares a fresh-water marsh occurs in the Parish. Common fresh-water herbaceous plants 
include: Alt thera philoxeroides, Eichhornia crassipes, Typha latifolia, Pontederia cordata, 
and Sagittaria sp. 

Swampland.—This region makes up about 4% of the parish. Soils are level, fluid,and 
loamy or clayey throughout (Roy & Midkiff 1988). Swamps are primarily used for wildlife 
habitat and for recreation. Major swamplands are associated with the Calcasieu River in 
the northeastern part of the Parish and with the Sabine River in western Calcasieu. Domi- 
nant trees include Nyssa aquatica, and Taxodium distichum. 

Longleaf Pineland.—This region was clear cut around 1900 and now serves prima- 
rily as forest range and cropland (Roy & Midkiff 1988). However, a few relatively undis- 
turbed sites within the Parish retain many of their original characteristics and are recog- 
nizable communities. We recognize four major communities within this region: hillside 
seepage bogs, semi-evergreen broadleaf acid seep forests, sodic flatwoods and acid 
flatwoods. 

A single hillside seepage bog occurs in Calcasieu Parish northeast of DeQuincy. This 
bog appears to be the headwaters of a small unnamed stream. A semi-evergreen broa- 
dleaf acid seep forest occurs along the sandy ravine adjacent to the bog. Distinctive 
herbaceous species in the bog include Rhynchospora oligantha, Sarracenia alata, and 
Utricularia cornuta. 

An area of sodic flat Is (sensu Smith 1996) occur in the western part of Calcasieu's 
longleaf pineland vegetation region. These flatwoods are typically saturated during the 
winter and spring but may become very dry Busing summer droughts. Herbaceous spe- 
cies are often prairie like and include Sparti Chaet a asteroides, Evolvulus 
sericeus, and Liatris punctata. The iacemion wooey yEnc oni is -sainied. 

A few recognizable acid flatwoods (sensu Smith 1996) occupy the central part of 
the longleaf pine region within Calcasieu Parish. This community is level to gently rolling 
with small elevated areas termed “pimple mounds” by Holland et al. (1952). Under natural 
conditions and frequent fires, these hs support a sparse canopy of longleaf pine 
with few other tree species (Bridges & Orzell 1989). Distinctive herbaceous species in these 
savannahs include Stylisma aquatica, and Platanthera nivea.The planting of pine planta- 
tions and protecting from fire have nearly eliminated these communities in the Parish. 

Bottomland Forests—This region occurs along the narrow flood plains of the 
Calcasieu River and its tributaries and along the Sabine River near the Texas border. Soils 
in this region are level, poorly drained,and loamy throughout (Roy & Midkiff 1988). Domi- 
nant trees that occur in bottomland forests include Liquidambar styraciflua, Quercus alba, 
Q, falcata, Carya sp., Platanus occidentalis, Taxodium distichum, Pinus taeda, and Fraxinus 


caroliniana. 


366 SIDA 19(2) 


APPENDIX 1 
Specific and subspecific entries are catalogued within their respective divisions. An- 
giosperms are further divided into the classes Liliopsida (monocots) and Magnoliopsida 
(dicots). Taxa are listed alphabetically by family, genus, and species within each category. 
The format used is species name, authority, reference, and whether the species is consid- 
ered native or introduced. Taxa that are designated by an asterisk (*) following the au- 
thor citation are introductions that appear to be naturalized. Authorities are abbreviated 
according to Brummitt and Powell (1992). A specimen reference includes the name of 
the collector, the collection number and the herbarium where the vouchered specimen 
is located. Frequently cited collectors are abbreviated as: A = Allen;B = Brown;C = Cocks; 
N = Neyland; M = Mayfield; T = Thieret; Th = Thomas. Unless otherwise indicated, speci- 
mens from the abbreviated collectors are housed in the following herbaria: A = NLU;B = 
LSU; C = NO; N = MCN; M = LSU; T = LAF; Th = NLU. One specimen each from Duke 
University (DUKE), New York Botanical Garden (NY), Missouri Botanical Garden (MO), United 
States National Herbarium (US) and Vanderbilt University (VDB) are referenced in the 
annotated list. Following the collection data is an abbreviation for the vegetation region 
where each species typically occurs: prairie (Pr); fresh marsh (FM); saline marsh (SM); 
swampland (S);longleaf pineland (Pi); bottomland forests (B). Although not a vegetation 
region, disturbed areas such as pastures, fallow fields, ditches, urban areas and roadsides 
are collectively abbreviated (D). Additionally, parasites are designated by (P), lithophytes 
by (L) and epiphytes by (E). Finally, taxa that have been designated as critically imperiled 
within Louisiana (S1),imperiled due to their rarity or vulnerability to extirpation (S2), rare 
and local throughout the state or found locally in a restricted region of the state (S3), 
reported from Louisiana but without conclusive evidence (SR), or of historical occur- 
rence but no records within the past 20 years (SH) are designated. a Natural Heritage 


Program of the Louisiana Department of Wildlife and Fisheries det I these rankings. 
APPENDIX 1 
ANNOTATED CATALOGUE OF SPECIES FOR CALCASIEU PARISH, LOUISIANA 
EQUISETOPHYTA Woodwardia virginica (L.) Sm., N 1293 Pi 
EQUISETACEAE DENNSTAED Te 
Pteridium — aquilin (L.) uhn ar. 


Equisetum hyemale L.ssp.affine (Engelm.) Calder 
& Taylor, N 564 B 


FILICOPHYTA 


ASPLENIACEAE 
Asplenium platyneuron (L.) B.S.P,, Maples 96 (MCN 
B 


So 


AZOLLACEAE 

Azolla caroliniana Willd., N 728,FM,S 
BLECHNACEAE 

Woodwardia areolata (L.) T. Moore, N 297 B,S 


pseudocaudatum (Clute) aE Wake 97 
(MCN) B, D, P 

DRYOPTERIDACEAE 

Athyrium Pie femina (L.) Roth ssp. asplenioides 
Michx.) Hulten, Maples 209 (MCN) B 

Onoclea sensibilis L. Maples 296 (MCN) B 

Polystichum acrostichoides (Michx.) Schott, N 
1133B 


LYGODIA 
glum pon cum (Thunb.) Sw.,* N 268 B, FM, 


NEYLAND ET AL., FLORA OF LOUISIANA 


OPHIOGLOSSACEAE 
Botrychium biternatum (Savigny) L.Underwood, 
Mani 106 wey) B 


tr hitnm dissecti 


3B 
ophogbsum soe, ae N 1215 


ee ioglossum nudicaule L.f,N1149D 

Ophioglossum petiolatum L., Parker 533 (NLU) B 

OSMUNDACEAE 

Osmunda cinnamomea L., N 

Osmunda regalis L.var. peubie ae ) A.Gray, 

POLYPODIACEA 

Fe naar ides (L.) E.G. Andrews & 

ham, Maples 95 (MCN) E 

PTERIDACEAE 

Pteris vittata LN 1415L 

SALVINACEAE 

vinia minima Baker, N 261 FM 

THELYPTERIDACEAE 

Macrothelypteris torresiana (Gaudich.) Ching,” 
Maples 213 (MCN) B 

Phegopteris hexagonoptera (Michx.) Fée, N 384 B 

Thelypteris dentata (Forssk.) E.P. St. John, N 1146 
B 


w 
Q 


ga a. (Dcne) Reed var. versicolor 
n) Lellinger, N 269 B, D, FM, $ 
ae pales: (Desv.) Morton, T 13387B 


LYCOPODIOPHYTA 


ISOETACEAE 

lsoetes melanopoda Gay & Durieu, Brooks & 
Maples 817 (MCN) D 

LYCOPODIACEAE 

Lycopodiella alopecuroides (L.) Cranfill, N 793 Pi 

Lycopodiella appressa (Chapm.) Cranfill, N 792 Pi 

Pseudolycopodiella caroliniana (L.) Holub,N 1087 

Pi 


SELAGINELLACEAE 
Selaginella apoda (L.) Spring, N 499 D, Pi 


CONIFEROPHYTA 


CUPRESSACEAE 
Juniperus virginiana L.var. virginiana, N 512 B, Pr 
Taxodium distichum (L.) Rich., N 607 FM, BM, S 


PINACEA 

Pinus ee P.Mill., Miller 5 (MCN) Pi, Pr 
Pinus elliottii Engelm.,* Miller 6 (MCN) D, Pi, Pr 
Pinus palustris P.Mill., N 797 Pi 

Pinus taeda L.,N 476 D, Pi, S 


MAGNOLIOPHYTA 
(Magnoliopsida) 
ACANTHACEAE 
Roe acustris (Schlecht. & Cham.) Nees, N 


ust ia ovata (Walt.) Lindau, N 64 

Walt Vn i. om D 
Ruellia humilis Nutt., i 1334D,P 

Ruellia ae Torr,A 15122 D, Pi, Pr, D 


ACERAC 

Acer pianee L,N1731D 

Acer rubrum var. drummondii (Hook. & Arn. ex 
Nutt.) Sarg., N 467 B,S 

Acer rubrum L. var. rubrum, N 1623 Pi 

Acer saccharinum L.,* N 1386 D 

AMARANTHACEAE 

Alternanthera philoxeroides (Mart.) Griseb.,* N 
1292 BM, D, FM, S 


Amaranthiis alhiys L.,Th 66 


904D 

Amaranthus blitum L.* 7 31617D 

Amaranthus cannabinus (L.) Sauer, N 1003 BM, 
FM 


Amaranthus spinosus L., N 812 D, Pr 

Amaranthus viridis L., N 1127 BM, FM, D 

Froelichia gracilis (Hook.) Mog,., Pias 4499 (LSU) 
, Pi 

ANACARDIACEAE 

Rhus copallina L., N 609 D, P 

Toxicodendron radicans (L.) Kuntze, N 1426 B, D, 
M, Pi, Pr 

BRIN HACERE 

imina (Michx.) Dunal, 729432 B 


APIACEAE 

Bowlesia incana Ruiz & Pav., John s.n. (NLU) D 

ie erecta (L.f.) Fern., N 666 D, Pi, Pr 

Chaerophyllum tainturieri Hook., N 448 D, Pr 

hee leptophyllum (Pers.) Sprague ex 
Britton & Wilson,* N 530 B, D, FM 

Cicuta maculata L. var. maculata, N 267 D, BM, 
FM 


Cynosciadium rere io A 16406 (LAF) D, Pr 
Daucus carota NT 

Eryngium f iat Walt., a 952 

Eryngium prostratum Nutt. ex DC., ee D, Pi 
Eryngium yuccifolium Michx., N 736 Pi, Pr 
Hydrocotyle bonariensis Comm. ex Lam., N 1352 


Hydrocotyl e umbellata L., N 1360 D,S 

Hydrocotyle Heuer eon N 706 D,S 

Limnosciadium pin m (DC.) Math. & 
Constance, N 538 DPr 


368 


Limnosciadium la (Engelm. & A. Gray) 
& Constance, A 16406 D, Pr 

Oxypolis fi lene (Wate Britton, N 784 

Ptilimnium capillaceum (Michx.) Raf., : 652 D, 


BM, FM, S 
ee corm (Ell.) i T 27955 B, Pi, S 
_N1 


Spermolepis ae inate ex DC.) Math. & 
nce, Th 14386 D,P 
faa. ses mee ex DC., Th 14422 B, 


Zizia aurea (L.) K.Koch, N 1750 B 


APOCYNACEAE 

Amsonia ludoviciana Vail, Gilmore & Smith 3543 
(LSU) Pr (S3) 

Amsonia tabernaemontana Walt., N 1218 D, Pi, 
Pr 


ees repens Shinners, Misgreth s.n. ee 

m cannabinum 
os di ifforme (Walt.) ae N658B 
AQUIFOLIACEAE 
x ambigua (Michx.) Torr. N& M 1587 (MCN) P 
llex coriacea (Pursh) Chapm., N 7507 Pi 
llex decidua Walt., N 477 B, Pr 
llex opaca Aiton, N 759 B, Pr, 
llex vomitoria Aiton, N 328 B, Pr, S 
ARALIACEAE 
Aralia spinosa L., N 892 B, Pr 
ARISTOLOCHIACEA 
Aristolochia sees Jacq., 129428 B 
Aristolochia serpentaria L., N 1418 Pi 
felpnenraiisis 


— 
ao 


Asclepias hirtell (Pennell) \ Woods., 1 67028 D 
As pies | Walt., N 713 Pi 
PECEDIOs engueNa hae N6/7 Pi 

EIl., N 280 Pi 


Asclepias perennis Walt., ee S 
Asclepias variegata L., N 1763 ae 
ie viridis Walt., N 543 
(Walt.) en N 688 B 


ASTER 
Acme ‘ppesitol var. repens (Walt.) R 


n, N 104 
Pes eee a L.,N 1027 BM, D, Pr 
Ambrosia psilostachya an . 1422 D,Pr 
Ambrosia trifida L., N 
at eel ovatum ine H.Robins., N 297 D, 
i, Pr 


wo 
es 


Ast dy umosus L 

Aster fragilis Willd. a : oe (MCN) B 

Aster lateriflorus (L) Britt., M & N 2628 (LSU) Pr 
Aster patens Aiton, McWilliams 244 (LAF) D, Pr 


SIDA 19(2) 


Aster praealtus Poir,, N 714 D,P 

Aster subulatus var.ligulatus Shinners, N 435 B,D, 
Pi 

poner ceOutiON ths i ae ee o 


BM, D 
Bidens aristosa ae a N 385 . FM, Pr, 


Bidens bipinnata L., N 1038 D, Pr 
Bidens frondosa L., N 1102 D, F 


onia asteroides (L.) UHér.,, N 1126 D, FM, Pi 

Boltonia diffusa Ell, N 937 BM, Pr 

(L.) DC, N693 BM 

Calyptocarpus vialis Less.,* N 635 D, Pr 

Chaetopappa asteroides (Nutt.) DC., N 1513 Pi 
(S2?) 


RQ gos Oe rs 
DOFTHICHHG HaLESCET IS 


Chaptalia tomentosa Vent., N 1194 Pi 
Centaurea biebersteinii DC.* N 1771 D 
Chrysopsis mariana L.,N&M 1593 (MCN) P 
Chrysopsis pilosa Nutt., N 942 Pi 
ISU HOM, on MIEN N 1255 D, Pi, Pr 
Conocliniu (L.) DC., N 263 D, Pi, Pr 
Conyza bonariensis (L L ) Crona,* Th 66922 D, Pr 
nee Sse (L.) Cronq., N 1458 D, Pr 


o ore opsstin incon la Nutt fe oe N 790 D,Pr 
Vahl) Cass., N 1390 D 
Echin cea sanguinea Nutt., N 657 D, Pi 

lpr prostrata (L.) L.,N 698 D,FM,S 
Lae 1105B 
Flephantopus nudatus A. Gray, ot) 

Elepha s tomentosus L.,N2 
ie eae lifolia (L.) Raf. oC, N1139D,B 
Frigeron philadel ph icus L.,N1 
Erigeron ee Muhl. ex ie, N 1773 D, Pr 
Erigeron tenu 


lenhantoniss caroliniqniyss 


ee capillifolium (Lam.) Small, N 397 D, 


Eu spatortun ee bee ia oo D, Pr 
II.,.N 
ol leucolepis (DC.) Torr. & i. Gray, N 973 


atorium nerfoliaty 


eupaoriv mohrii Greene, A 17133 (LAF) Pi 
N89 


pie rotundi folium ‘i. N 737 D, Pi 

Fupatorium semiserratum DC.,A 16120 (LAF) Pi 

Eupatorium serotinum Michx., N 957 BM, FM 

wee hemisphericus (Alexander) Nesom, N 
3 Pi 


hana gymnospermoides Greene, Th 98160 D, 
Pr 


NEYLAND ET AL., FLORA OF , LOUISIANA 


Euthamia leptocephala (Torr. & A. Gray) Greene, 


N 1045 BM, Pr 
Gaillardia aestivalis (Walt.) H.F.L. Rock, N 718 Pi 
Gaillardia pulchella Foug., N 1225 D 
ochaeta pensylvanica (Willd. 
M 1713 (MCN) D 
Gamochaeta purpurea L., N 1367 D, Pr 
Grindelia papposa Nesom & Suh Pigs & Thomas 
4515 (LSU) Pr 
ymnostyles anthemifolia Juss.,* T 30610 D 
Helenium amar af.) H.F.L.Rock, N 7389 D, Pr 
Helenium diamond H. ie ba N 5/77 Pi 
Helenium flexuosum Raf., N 
Helenium vernale Walt., ee 3589 (LSU) Pi 
Helianthus angustifolius L.,N 1119 Pi 
Helianthus annuus L., N 823 D, Pr 
Helianthus debilis ssp. cucumerifolius (Torr. & A. 
Gray) Heiser Correll & Correll 9625 (LSU) Pi 
Helianthus mollis Lam., N 
Heterotheca subaxillaris (Lam.) Britt. & Rusby, N 
D, Pr 
Hieracium longipilum Torr., C s.n. Pi 
Hymenopappus artemisiifolius DC., N 1276 D, Pi 
Hypochaeris microcephala (Sch.-Bip.) Cabrera,* N 
263D 


nae 


Cabrera, N & 


— 


ochaeris radicata L.* N& M 1722 (MCN) D 
onactis linariifolius (L.) Greene, N&M 1606 (MCN) 
Pi 


—~ 


lva angustifolia DC.,N 1120D 

lva annua L.,N 1090 D 

Iva frutescens L., N 1469 BM 

Krigia cespitosa (Raf) Chambers, N 1753 D 
Krigia dandelion (L.) Nutt., Hester s.n. (LAF) D 
Lactuca canadensis L., N 1446 D, Pr 

Lactuca floridana (L.) Gaertn., N 372 D,Pr 
Liatris acidota Engelm. & A. Gray, N 295 Pi 
Liatris aspera Michx., N 1515 Pi 

Liatris elegans (Walt.) Michx., N 919 Pi 


Marshallia caespitosa Nutt.ex DC., N 1280 Pi 
—— ee ee Bs as (Ell. 
eadle & F.E. Boynt., 
oro cro (L. f.) aa i ests D, ie S 
ns (L.) Willd., N 
Lea aiid (Torr. & A. ca el ie 


wel 


a 


Se ieee rigi tee var. glabratum (E.L. Braun 
Nesom, Cs.n 
Parthenium hysterophor L.*N1529D 
(Michx.) Nutt. N 855 Pi 
aan camphorata (L.) DC., N 375 BM, D,FM,S 
Pluchea foetida (L.) DC., N804 D, FM, Pi,S 


ityopsis 


369 


Pluchea odorata (L.) Cass., N 1002 BM, D, FM, Pr, 

Pluchea rosea Godfrey, N 292 BM, D, FM, Pi 

sei ctl obtusifolium (L.) ssp. 

usifolium Hilliard & Burtt, N 1035 Pi 

Perocoulon vigatin (L.) DC., 720612 Pr (S1) 

Pyrrhopappu Walt.) DC., Costanza 
67 (MCN D 

age columnifera (Nutt.) Woot. & Standl., T 

28897 Pr 


liniany 


eee lier (D. Don) J.F. Gmel. ex DC., 
Robichaux 4 (MCN) D, Pi 

Rudbeckia hirta L.,N 1330D 

Rudbeckia subtomentosa P ursh, N1500B 


Rudbeckia texana (Purdue) Cox & Urbatsch, N676 


Senesio gaeele US, Poir., ne we D, Pr, S 
hi ile A.Gra _ Pi, Pr 


wimoara 


marae radul a Nutt 

Silphium integrifolium ihe th 98155 D,Pr 
eee enn) lk, ey 

(L.) ven ex Small, N 


Solidago caesia L.,N 1556 Pi 

Solidago canadensis var. scabra Torr. & A. Gray, N 
D, Pi, Pr 

Solidago odora Aiton, N 1544 D,P 

Solidago rugosa P.Mill., N& M 1599 (MCN) D, Pr 

Solidago sempervirens var. mexicana (L.) Fern., N 


Solidago stricta Aiton Urbatsch 2090 (LSU) BM 


, Pi 

Solidago tortifolia Ell., N 1543 Pi 

Soliva mutisii Kunth,* N 7758 D 

Soliva sessilis Ruiz & Pav.,* N 457 D 

Sonchus asper (L.) Hill,*N 578 D 

Sonchus oleraceus L.* N518D 

Symphyotrichum pratensis (Raf.) Nesom, N& M 
N) Pi 


oo officinale G.H. Weber ex Wigg.,* N 


ee virginica L., N 1068 B,D 
silat cu tea (Walt.) Trel. ex Branner & 
Sp gigantea Pi 

Vernonia missurica Raf, N 1083 3 Pi, Pr 
eae texana (A. Gray) melee N 1430 Pi 
Youngia japonica (L.) DC.* 3D 
BERBERIDACEAE 

Podophyllum peltat. imL., N 1683 B 
BETULACEAE 

Betula nigra L.,N 1099 B 

Carpinus caroliniana Walt., N 464 B 

Ostrya virginiana (P. Mill.) K.Koch, N 758 B 


370 


BIGNONIACEAE 
Ri ; | 


Bignonia capreolata L., N 1203 B, Pi, S 
Campsis radicans (L.) Seem. ex Bureau, N 279 B, 
D, Pi, Pr 
Catalpa bignonioides Walt., N 1274 D, Pr 
BORAGINACEAE 
Heliotropium curassavicum L., N 1307 BM, FM 
Harel indicum L.,* N 993 D, ii 
opium procumbens P Mill, C 
he tis macros, perma Engelm., N 1 1333 B,D,S 
BRASSICACEAE 
Capsella bursa-pastoris (L.) Medik,* N 1152 D 
Cardamine bulbosa (Schreb. ex Muhl.) B.S.P, N 
179B 


Cardamine debilis D.Don, N 1212 D 
i Muhl.ex Willd., N 447 D, 
Coranopus didymus (L.) vn *N1171D,Pr 
Lepidium virginicum LN 173 D, 
Raphanus raphanistrum 2 T 28864 D 
Rorippa sessiliflora (Nutt.) Hitche., N 1205 D, Pr 
Rorippa teres (Michx.) Stuckey, N 1777 D, Pr, S 
BUDDLEJACEAE 

olypremum procumbens L., N 707 Pi 
CABOMBACEAE 
Brasenia schreberi J.F.Gmel., N870S 
Cabomba caroliniana A. Gray, N 1392 § 
CACTACEAE 
Opuntia humifusa (Raf) Raf. var. humifusa, N 1791 

D 


CALLITRICHACEAE 

Callitriche heterophylla Pursh, N 1168 D,S 
Callitriche peploides Nutt.,N 1759D,§ 
CAMPANULACEAE 

Lobelia appendiculata . re N 1298 D, Pi 
Lobelia cardinalis L., N 1S 

Lobelia flaccidifolia eit N 576P 

ay ia puberula Michx. var. eu a,N 1122 D, 


Lobel ia puberula var. pauciflora Bush, N 1570 Pi 

Triodanis ae iata var. biflora (Ruiz & Pavon) Bra- 
dley, N 

Triodanis ee (L.) Nieuwl. var. perfoliata, N 


, er 
CAPRIFOLIACEAE 
Lonicera japonica Thunb.,* N 347 B, D, Pi, Pr, FM, 
5 


pee Sere E EN ‘4 i 650 D, Pi, B 

613 D,FM, Pr,S 
Viburnum dentate L, finn B,D,S 
Viburnum nudum L., N 1 


SIDA 19(2) 


CARYOPHYLLACEAE 
Cerastium glomeratum Thuill,* N 438 D, Pr 
Sagina decumbens (Ell) Torr. & A. Gray, N 437 D, 


Silene antirrhina L., N 1209 B, 
Spergularia salina (L.) Bis N 1305 BM 
Stellaria media (L.) Cirillo,* N 474 D, Pr 


eee. 
aLl.N416B 


CERATOPHYLLACEA 
Ceratophyllum cea L., N 704 BM, FM 


CHENOPODIACEAE 

Chenopodium ambrosioides L.,* T 29422 D 
Salicornia bigelovii Torr, Montz 2296 (LAF) BM 
Suaeda linearis (Ell..) Moq., N 1433 BM 


CISTACEAE 
Lechea mucronata Raf., T 31636 D, Pi 
Lechea “oe Raf., N& Komo 1782 Pi 


CLUSIA 
ees Fenn (Spach) Steud., N 1404 
i 
Hypericum cistifolium Lam., Th 89297 FM, Pi 
Hypericum crux-andreae (L.) Crantz., N 946 D, FM, 
Pi,S 


Hypericum densiflorum Pursh, N 1403 B, D, Pi, Pr 

Hypericum v.& Hook.) Torr. & A. 
Gray, N 1473 B Pi 

Hypericum hypericoides (L.) Crantz, N 329 D,B, Pi, 
Pr 


Hypericum mutilum L., N 1075 B 
Hypericum setosum L., N 286 Pi 
Triadenum tubulosum Walt.,N 1111S 
Triadenum walteri J.G.Gmel.,N 1112S 


CONVOLVULACEAE 
cal ystedia sash (L.) R.Br,* N 1443 BM, FM 
ndra carolinensis Michx., N 452 D 
Evolvulus sericeus ae N 1761 Pi 
Ipomoea coccinea L.,* N 840 D,S 
{pomoea  cordatotriloba Dennst. — var. 
cordatotriloba, N 376 D, Pr, S 
oe ganesd L., N 1492 B, D,S 
ndurata (L.) G.Mey., N 781 B, D 
Ipomoea purpurea (L.) Roth,* N951D 
lpomoea quamoclit L.,* N 1523 D 
[pomoea sagittata Cav. N679 BM, FM 
[pomoea wrightii A. Gray,” N 847 D, Pr 
Jacquemontia tamnifolia (L.) Griesb.,, N 390 D 
Stylisma aquatica (Walt.) Raf, N 1382 Pi 


CORNACEAE 

Cornus drummondii C.A. Mey., N 1314B 
Cornus florida L., Costanza 69 (MCN) B,$ 
Cornus foemina P.Mill., N&M 1602 (MCN) B,S 


NEYLAND ET AL., FLORA OF , LOUISIANA 


CRASSULACEAE 

Penthorum sedoides L.,N 1100 B,D, S 

CURCURBIT. — 

Cucumis melo D 

Melothria facet L., fier B, D, FM, BM, S 

CUSCUTACEAE 

Cuscuta cuspidata Engelm., N 1482 P 

Cuscuta glomerata Choisy, N 1795 P 

Cuscuta indecora Choisy, N 689 P 

Cuscuta pentagona Engelm. var. pentagona, N 
1401 P 


CYRILLACEA 
Cyrilla heat L., N 856 B, Pi 


DROSERACEA 
Drosera paca Pursh, N 469 D, Pi 


EBENACEAE 

Diospyros he L.,N 71315 B, Pi 
ELATINACE 

Elatine ae Schkuhr, Givens 2788 (LSU) D 
ERICACEA 


Monotropa aoe L., Brooks 343 (MCN) B 
Rhododendron canescens (Michx.) Sweet, N 659 


=> 


Vaccir cinium eee Marshall, N 1237B 
Vacci stamineum L., N 494 B,S 
aie elliottii Chapm., N 1189 B,S 
Vaccinium virgatum Aiton, T 16966 Pi 


EUPHORBIACEAE 

Acalypha gracilens A. Gray, T 29398 D, Pr 

Acalypha monococca (Engelm. ex A. Gray) L., N 
1612 Pi 


peas ras Riddell, N 848 D 

Acalyp omboidea ay . | D 

eae vigiica - = ee 

Caperonia palus 

Chamaesyce eee ia i. vite : ee n. 


Chamaesyce hyssopifolia (L.) Small,* N 825 D 


Chamaesyce serpens (Kunth) Small, N 1432D 
Bu oa texanus (Muell. Arg.) Small, N 1272 


ee oa Michx., : 733D 
Croton alandiwlocic L., 

e118 willdenowi 7 Ls regen N1779D 

L., N 305 Pi, Pr 

Euphorbia dentata Michx,, N 1046D 

Euphorbia heterophylla L.,* N 1011 D 

Euphorbia meganoesos Featherman, N 686 D 

Euphorbia texana Boiss., N& M 1716 (MCN) D 


Phyllanthus polygonoides ie aa 66994 D 

Phyllanthus urinaria L.* 

Sapium sebiferum (L.) oun Ne B, BM,D, FM, 
Pr 


Sebastiania fruticosa (Bartr.) Fern., N697 $ 
Stillingia sylvatica Garden ex L., N 1846 Pi 
Tragia smallii Shinners, Th 66995 Pi 

Tragia urticifolia Michx., N& M 1591 (MCN) Pi 
Vernicia fordii (Hemsl.) Airy-Shaw,* N 1220 B 


FABACEAE 


(L.) Willd., N 620 i FM,S 
Acschvaomencinaiea L., N 849 D,P 

Albizia julibrissin Durz.,* 

Amorpha LN N 1377D, . 

ee amencang peas, N 1507 

Torr. & A. ne oul Pi 


hylia ( arisey caly N 608 


fruticosa 


Baptisia alba 


D, Pr 

Baptisia bracteata var. laevicaulis (A. Gray ex 
Canby) Isely, N 700 Pi 

Baptisia sphaerocarpa Nutt.Lasseigne 1417 
Pi 


—> 


LAF) 
Centrosema virginianum (L.) Benth., N 860 D, Pi, 
Pr 


Cercis canadensis L.,N 467 B,S 
Chamaecrista fasciculata (Michx.) Greene, N 340 


, Pr 
Chamaecrista nictitans (L.) Moench ssp.nictitians 
N9I8D 


Crotalaria sagittalis L. Lasseigne 1579 (LAF) Pi 

Dalea candida Willd., N 1775 Pr 

Dalea purpurea Vent., 1 31493 Pi 

Desmanthus illinoensis (Michx.) MacM. ex B. L. 
Robins. & Fern., N 784 D 

Desmodium ciliare (Willd.) DC., Cs.n.D, Pi 

Desmodium gene a DC., N 983 B 


NMecmod 


jm naniculatt m (| L)D 


C.,,N377B 
Dioclea multiflora (Torr. & A Gray) Mohr, N 1448 


Erythrina herbacea L., N 656 B, D, Pi 
Galactia volubilis (L.) Britt. N 937 D,$ 
Gleditsia aquatica Marshall, N & Komo 1737 


Gleditsia niecane thos L., N& Komo 1635 (MCN) § 

ee oe vesicarium (Jacg.) Harper, N 1027 D, 
FM 
Kum erowia striata (Thunb.) Schindl.,* 
Lasseigne 1434 (LAF) D, Pi 
Lesped neata (Dum-Cours.) G. Don,* 
Lasseigne 1597 (LAF) D 
Lupinus texensis Hook.,* N 1223 D 
Medicago arabica (L.) Huds.,* N & Komo 1670 
CN 


(MCN) D 
Medicago lupulina L.,* N 442 D 


372 


Meaicage minima (L.) L..* Conrad 1 


nolvm rpha L., 


1 (MCN) D 
4D 


Melilotus one inalis (L.) fine * Lasseigne 1564 
LAF) D 


Melilotus indicus (L.) All,* N 509 D 

Mimosa hystricina Small ex Britton & Rose) B.L. 
Turner, N 1344D 

Mimosa microphyl la an Zebryk 2252 (NLU) Pi 

Mimosa nuttallii B.L. Turner, Th 129572 D, Pi 

Mimosa strigillosa Torr.& A. Gray, N 752 D, Pr 

Neptunia lutea (Leavenworth) Benth., N 1348 D, 


r, Pi 
Orbexilum pedunculatum (P. Mill.) Rydb. var. 
pedunculatum, B 13531 (NLU) D, Pi, Pr 
Orbexi lum simplex (Torr. & A. Gray) Rydb., N 1343 


ss aculeata L.*N571D 

Pueraria montana (Lour.) eal var ae (Willd) 
rane & Almeida,* N 

Rhynchosia latifolia (Nutt. ie r A. Gray, Cheva- 
lier 16 (MCN) Pi 

Rhynchosia minima (L.) DC., N 1026 D, Pi 

Rhynchosia reniformis (Pursh) DC., Ware s.n.(NLU) 
D, Pi 


{hynchosia tomentosa (L.) Hook. & Arn,, Pias 4520 
(LSU) Pi 

Robinigq psel idoacaciaq L.* 

Senna obtusifolia (L.) Irwin & Barneby, N 753 D 

Sesbania drummondii (Rydb.) Cory, N6/8 D, BM, 
FM 


Sesbania herbacea (P. Mill.) McVaugn, N 830 D, 

Sesbania punicea (Cav.) Benth.,* Lasseigne 1564 
(LAF) D, BM, FM 

Strophostyles helvula (L.) Ell, N 928 D, Pr 

so es umbellata (Muhl.ex Willd.) Britton, 

N 1540 D, Pr 
losanthes bifl fa (lL. ) BSP, N 719 Pi, Pr 
robrychoides Nutt., N 1396 Pi 
Titoli ium mee Sturm, Walker s.n.(MCN) D 
Trifolium dubium Sibth.,* Milsom 3 (MCN) D 


Vigna luteola (Jacq.) Benth., N 997 D, BM 
Wisteria floribunda (Willd) DC.,* N 1232 D 
15 (L.) Poir., N 1740 B 


7 (MCN 


a 


AGACEAE 
Castanea pumila P. Mill, N& Komo 161 
i 
Fagus grandifolia Ehrh., N 458 B 


SIDA 19(2) 


Quercus acutissima Carruthers,* N 1785 D 
Quercus alba L.,N 1562 B 
Quercus falcata Michx., N 646 B, D, Pi 
Quercus hemisphaerica Bartr., 28035 D, Pi 
Quercus incana Bartr., 1 25652 Pi 
Quercus laurifolia Michx, i. 082 B,S 
Quercus lyrata Walt., N 667 
Quercus margarettiae rae ex Small, M 2920 Pi 
Quercus marilandica Muench, N 666 D, Pi 
Quercus michauxii Nutt., N 325 B, Pi 
Quercus nigra L., N 507 B, D, Pi, Pr, S 
Quercus pagoda Raf., N 1142 
Quercus phellos L., N 668 Pi 

uercus shumardii Buckl, N& M 1717B 
Quercus similis Ashe, T 33 i 
Quercus stellata Wangenh., N 1083 Pi 
Quercus texana Buckley, N & Kitt 1704 D, Pi 
Quercus virginiana P.Mill., N 475 B, D, Pi, Pr 


GENTIANACEAE 
Centaurium pulchellum (Sw.) Druce,* N 1307 D, 


r 
Eustoma exaltatum (L.) Salisb. ex G. Don., N 786 
D, BM, F 


Sabatia calycina (Lam.) Heller, N 1104 D, Pi, S 

Sabatia campanulata (L.) Torr., N 1429 D, Pi 

Sabatia campestris Nutt. N 73171 D, FM, Pi, Pr 

Sabatia dodecandra L. (B.S.P) var. foliosa (Fern.) 
ee Bae on =) Pi 


— 


EIL, N 739 Pi 
GERANIACEA 
Geranium ain L..N445 D,Pr 
GROSSULA dain 
Itea virginica L..N491§ 
HALORAGACEAE 


Myriophyllum aquaticum (Vell.) Verdc.,* N 336 D, 
Myriophyllum heterophyllum Michx., N 721D,FM, 
S 


aU a ial pinnatum (Walt.) B.S.P., Coe 8 
)D , , S 

Myriphylun tar N 1124 D, FM, BM 

P L.,N623 D,FM,$S 

Proserpinaca pectinata Lam, N 149 D, Pi, S 

HAMAMELIDACEAE 

Hamamelis virginiana L., N 323 B 

Liquidambar styraciflua L., N 1214 B, D, Pi, Pr 

HIPPOCASTANACEAE 

Aesculus pavia L.,N 1221 B 

HYDROPHYLLACEAE 

Hydrolea ovata Nutt. ex Choisy, N 300 D, Pi, Pr 

Nemophila aphylla (L.) Brummitt, N 579 D, B 

Phacelia hirsuta Nutt., N 480 D, Pr 


LOUISIANA 


NEYLAND ET AL., FLORA OF 


JUGLANDACEAE 

Carya alba (L.) Nutt. ex Ell, N 1084 Pi 

Carya aquatica (Michx.f) Nutt., N 1042 B 

Carya cordiformis (Wangenh.) K.Koch, N&M 1628 
MCN) B 


Carya glabra (P. Mill.) Sweet, N 7085 B, Pi 

Carya illinoinensis (Wangenh.) K.Koch, N 1728 D 
Carya ovata (P. Mill.) K. Koch, N 1061 Pi 

Juglans nigra L.* N 1512 D,P 


LAMIACEAE 

Hedeoma hispida Pursh, N 1326 D 

Hyptis alata (Raf.) Shinners, N 738 D, Pi, Pr 

Lamium amplexicaule L.,* Costanza 43 (MCN) D 

Lycopus rubellus Moench, N 1 oe D,FM 

Lycopus virginicus L.,N 1509P 

Monarda en A.Gray, Correll Correll 9572 
(LSU) P 

Monarda punctata L.,N 841 D,Pr 

Perilla frutescens (L (L.) Britt..* N 7268 D 

Physostegia digitalis Small, N 750 D, Pr 

Physostegia longisepala Cantino, N 643 D, Pi (S2) 

Physostegia virginiana (L.) Benth. ae praemorsa 
(Shinners) Cantino, alae 

Prunella vulgaris L., N 13 

Pycnanthemum ieee Torr. & A. Gray, N 903 

fr Pi, 

Pycnanthemum tenuifolium Schrad., N 678 D, Pi 

Salvia azurea Lam., N 1050 D, Pr 

Salvia coccinea Buchoz ex Etlinger,* 
(MCN) D 

Salvia lyrata L., Costanza 9 (MCN) D 

se ae cardiophylla Engelm.& A. set 123535 


Coto 45 


ee drummondii Benth., Th aa Pi 
Scutellaria integrifolia . : 578 B,P 

Stachys crenata Raf, N D 

Stachys floridana Shutlew ex es N1217D 


eucrium cubense Jacq., N 1 . 
Trichostema dichotomum L., N 1537 Pi 
LAURACEAE 


Cinnamomum camphora (L.) Presl,* N 606 B, D, 
Pr 


pasea palatite (Raf.) Sarg., N 904 B 

m (Nutt.) Nees., N 7 fe Pi, Pr 
LENTIBULARIACEAE 

Pinguicula pumila Michx., N 1253 Pi 

Utricularia cornuta Michx., N 1746 Pi 


eedane inflata Walt, N 473 FM,S 
wilariq jun loyal Vahl, N 1508 P 

ce macrorhiza LeConte, Curry 1479 (LSU) 
FM,S 


Utricularia radiata Small, N 495 § 
Utricularia subulata L.,N 1217 Pi 


LINACEAE 

Linum medium (Planch.) Britt. var. texanum 
(Planch.) Fern. N679 D, Pr 

Linum striatum Walt., N 1774 D, Pi 


LOGANIACEAE 

Gelsemium BUI (L.) J. St.-Hil., N & Komo 
1675 (MCN) B, D, P 

Mitreola petiolata (J. F. aa Torr. & A. Gray, N 


74 
Mitreola sessilifolia (J.F.Gmel.) G.Don, N 303 D,S 


Fenea eae 
i cinea Rottb., N 
Cuphea ae (Jacq, ) rae *N844D, 


Dips. diandra (DC.) Wood, N 1752 D 

Lyt alatum Pursch var./anceolatum (Ell.) Torr. 
s Gra ray ex Rothrock, N 339 D 

Rotala ramosior (L.) Koehne, N 924 D,FM 


MAGNOLIACEAE 

Liriodendron tulipifera L.,* N 1 bee D 

Magnolia grandiflora L., N 1381 

Magnolia virginiana L., N 1242 ‘ S 

MALVACEAE 

Cole panaver (Cav,) A. nee N1331D 

Walt., N 282 D, Pi 

Hibiscus eine L. een (Cav.) OJ. 
Blanchard, N 1427 D, BM, FM 

Hibiscus moscheutos L.ssp.moscheutos L., N 1444 


Kosteletzkya virginica (L.) Presl.ex A.Gray, N 1434 
BM 


Malvaviscus arboreus Dill.ex Cav.var.drummondii 
(Torr. & A. Gray) Schery,* N 1472 D, Pr 

Modiola caroliniana (L.) G.Don, N 1732 D, Pr 

Sida rhombifolia L., N 726 D 

Sida spinosa L., Th 66905 D 


MELASTOMATACEAE 

Rhexia alifanus Walt., N 284 Pi 

Rhexia lutea Walt., N 1385 Pi 

Rhexia mariana L.var.mariana, N 653 D, Pr, Pi 
Rhexia petiolata Walt., N 283 Pi 

Rhexia virginica L., N 290 Pi 

MELIACEA 

Melia a L.*N670D 
MENISPERMACEAE 

Cocculus carolinus (L.) DC.,N 1504D 


MOLLUGINACEAE 
Mollugo verticillata L.,N 1405 D, Pr 


MORACEAE 

Fatoua villosa (Thunb.) Nakai,* N 7743 B 
Maclura Aah een Schneider,* N 1036 B,D 
Morus alba L., 

Morus rubra L., N 1 me B, D, Pi, Pr 


MYRICACEAE 
Myrica cerifera L.,N351B 
Myrica eee Raf, oe i 1598 (MCN) Pi 


NELUMBONACEA 
Nelumbo lutea aie Pers., N 1005 BM, FM 


NYCTAGINACEAE 
Boerhavia erecta L.,N 1093 D 


NYMPHEACEAE 
Nuphar advena (Aiton) Aiton f, N 542 D, BM, FM, 
Ss 


Nymphaea elegans Hook., N 1006 BM (S1/S2) 
Nymphaea odorata Aiton, N 1064 D, BM, FM 


NYSSACEAE 

Nyssa aquatica L.,N 662 S$ 

Nyssa biflora (Walt) Sarg., N 333 Pi, Pr 
Nyssa sylvatica Marshall, N 1250 P 


OLEACEA 

eneee virginicus L., N 1210 

Fraxinus americana L., N Komo ce (MCN) B, 
Pi 


Fraxinus caroliniana P. Mill, N 664 $ 

Fraxinus pennsylvanica Marshall, N 1016 B 

Ligustrum japonicum Thunb.,* 

Ligustrum lucidum Aiton f.,* N & M 1595 (MCN) 
Pi 


Ligustrum sinense Lour.,* N 348 D, Pr 
ONAGRACEAE 
Gaura lindheimeri Engelm.& A. Gray, N 574 D, Pi, 


Gaura longiflora Spach, N 941 D 

Gaura parviflora Douglas ex Lehm., N 1406 D, BM 

Ludwigia alternifolia L., Brooks 394 (MCN) B, D, 
FM 


Ludwigia decurrens Walt., N 859 D, FM, Pr, S 

Ludwigia glandulosa Walt., N 922 D, FM, & 

Ludwigia hirtella Raf, N 1395 P 

Ludwigia leptocarpa (Nutt.) Hara, N 975 D, BM, 
FM 


Ludwigia | inearis Walt. iN 1053 D, FM, Pr, S 
_Orzell & bridges 5760 


(LSU) Pi (S1) 
—— octovalvis (Jacq.) PH. Raven, N 378 D, 
M,S 
a palustris (L.) Ell, N 923 D,S 
Ludwigia peploides (Kunth) PH. Raven, N 927 D,S 
Ludwigia pilosa Walt., Th 81866 D, FM, S 
Ludwigia sphaerocarpa Ell., N 1076 D, FM, Pi, S 


SIDA 19(2) 


Ludwigia uruguayensis (Cambess) H. Hara, N 740 
D, BM, FM, S 

Oenothera biennis L.,N 1022 D 

Oenothera grandis (Britt.) Smyth, Givens 1887 
LSU) D 


Oenothera heterophylla Spach, N 1453 D 
Oenothera laciniata Hill, N 560 D 

Oenothera linifolia Nutt., N 1241 P 

Oenothera spachiana Torr.& A. Gray, Givens 1888 


= 


, Pr 
Oenothera speciosa Nutt., N 13217 D 


OROBANCHACEAE 
Epifagus virginiana (L.) Barton, N419B 


OXALIDACEAE 

— debilis var. corymbosa (DC.) Lour,* N& M 
as Me D 

oa. a St.-Hil.* N 71237 D 

Oxalis ail ae ae Costanza 11 (MCN) D 

PAPAVERAC 

Argemone eae Hornem., N 1736 D 

PASSIFLORACEAE 

Passiflora incarnata L., N 346 D 

Passiflora lutea L.,N 273 B,D,§S 

PEATOEACEBEERE 


americang 


PLAN TAGINDS EAE 


ictota NAj 
WHC 


— 


Blantage heterophylla Nutt, eee MCN) D 
Plantago major L., N 642 


Plantago virginica L., N 444 D 


PLATANACE 
Platanus a L.N513B,$ 


POLEMONACEAE 
Phlox pilosa L., N 1277 D, Pi 


POLYGALACEAE 

Polygala crenata James, T 25650 Pi 

Polygala cruciata L., N 289 Pi 

Polygala incarnata L.,N 1760 P 

Polygala leptocaulis Torr.& A. on N712D,Pi,Pr 
Polygala mariana P.Mill., N 566 Pi 

Polygala nana (Michx.) DC, N675 Pi 


olygala verticillata L.,N 625 D 
POLYGONACEAE 

Brunnichia ovata (Walt.) Shinners, N 829 B,S 
Polygonum cespitosum Blume, N 1115 D, Pr 
Polygonum densiflorum Meisn., N 826 D,S 
Polygonum hydropiperoides Michx., N 344 D, FM, 


Polygonum pensylvanicum L., N 1009 D, FN, S 


NEYLAND ET AL., FLORA OF CALCASIEU PARISH, LOUISIANA 


at punctatum EI, N 382 D, ie Pr, S 
Moris, N8&27D 

He crispus Ls *N 1308 D 

Rumex hastatulus Ell, N592D 

Rumex obovatus Danser,* N 1362 D 

Rumex pulcher L.* Hebert 4 (MCN) D, Pr 

Rumex verticillatus L., N 1229 D, FM, § 


PORTULACACEA 

Claytonia virginica i. N1198B 
Portulaca oleracea L.,N811D 
Portulaca pilosa L., N 1886 D 


PRIMULACEAE 

Anagallis arvensis L.* N 1361 D, PI, PR 
Anagallis minima (L.) Krause, N 1182 D 
peaaenG radicans te N690S 

Kunth, N 71347 D, BM (S1) 
Samolus valerandi L.,N 1269 B,D,S 


RANUNCULACEAE 
Anemone caroliniana Walt, Brooks 379 (MCN) D, 
Di 


— 


i 

Clematis crispa L., N 762 B,D, S 

Clematis terniflora DC.,* N271D 

Delphinium carolinianum Walt., Lemmon 1204 

Pi 
Ranunculus fascicularis Muhl. ex Bigelow, 
Mistretta s.n.(NLU) D, Pr 

Ranunculus muricatus L.* N 1180 D 

Rgmeactlls palsies L.* Mistretta s.n.(NLU) D 
pusillus Poir., N 1169 D, FM, S 

ee aes sardous nae: N621D 

Ranunculus sceleratus L.,N 1164 D,FM,$ 

Ranunculus trilobus Desf.,* T 30626 D 


RHAMNACEA 
eee Seas (Hill) K. Koch, N 1397 B, D, 
i,S 


roa caroliniana (Walt.) A. Gray, N 1474 B 


OSAC 
ton crbtoo (L.) Ell, N 1574 Pi 
rshallii Eggl., N 1204 B 
a : ae aa Hook. & Arn., N 1683 B 
Crataegus viridis L.,N 1071 B, S 
Duchesnea indica (Andrz.) Focke,* Ashworth 19 
MCN) D 


Geum canadense Jacq., N 1354S 

Prunus caroliniana (P. ie : Aiton, N 1031 B,D 
Prunus serotina Ehrh., N ,D 

Prunus americana oo N& uA 
Pyrus calleryana Decne.,* N 1072 
Pyrus communis L.,* 

Rosa bracteata Wendl.,* N 1356 D 
Rubus argutus Link, N 1290 D, Pi, S 
Rubus trivialis Michx., N 1257 D, Pi, Pr 


1607 (MCN) B 


RUBIACEAE 

Cephalanthus occidentalis L.,N 715 FM,S 
Diodia teres Walt., N 955 D, Pi, Pr 

Diodia virginiana L., N 341 D, Pr, 
Galium aparine L., N 1165 B,D, Pr 
Galium parisiense L.,* T 29393 D 
Ga ne tinctorium (L.) Scop.,N 597 D, Pr, S 
Hedyo nigri ans (La mb.) Fosb., N 587 D, Pr 
Houstonia micrantha (Shinners) Terrell, N 449 D 
Houstonia pussila Schoepf,, N 440 D, Pr 
Mitchella repens L., N 276 (MCN) B 
Oldenlandia boscii (DC.) Chaom., N 1477 B, Pi 

Oldenlandia uniflora L., N & M 1610 (MCN) Pi 
Sherardia arvensis L.,* N439D 


RUTACEAE 

Poncirus trifoliata ‘ ’ be - 1213B 
Ban es LN 

Zan um clava- dc L., N 1306 D, Pr 


SALIC 

Populus ate Bartr. ex peat N 1370 B,D 
Salix babylonica L.,* N 
Salix humilis Marshall, ee Pr (S2) 
Salix nigra Marshall, N 468 D, S 


GW 


SAPINDACEA 

Caispermum na ledeneuae ie i 1000 D 
lreviteria pan iculata Laxm.,* 1554D 

SAPOTACEAE 


Sideroxylon lanuginosum  Michx. — ssp. 
ee (Nutt.) T.D. Pennington, N&M 


CN) P 
Sideroxylon ae L. N&M 1603 (MCN) P 
SARRACENIACEAE 
Sarracenia alata (Wood) Wood, N 1496 Pi 
SAURURACEAE 
Saururus cernuus L.,N 639 
SAXIFRAGACEA 
epee atte (Muhl,) Ell, N 7276 D 
SCROPHULARIACEAE 
Agalinis rena ts (Ell) Raf., N 7055 D, Pi, Pr 
Agalinis filicaulis (Benth.) Pennell, N 1538 Pi (S1) 
Agalinis heterophylla (Nutt.) Small ex Britt., N 1546 


Agalinis oligophylia Pennell, N 7 Wee Pi 

Agalinis purpurea L. Pennell, N 

Agalinis tenuifolia (Vahl) ee s.n.(NLU) D, Pi 

Agalinis viridis (Small) Pennell, Urbatsch 2087 
(LSU) D 


Aureolaria pectinata (Nutt.) Pennell, Givens 4265 
(LSU) Pi 


Aureolaria virginica (L.) Pennell, N 1486 Pi 
Bacopa caroliniana (Walt.) Robins., N 299 D, FM, 
S 


376 


eacona monnieri ite N 1400 D, FM, S$ 


N3 
Bacopa rotundl Holi id + (Michx) Wettst., N 1446 D, 


Buchnera americana L., N 1345 Pi 

Castilleja coccinea (L.) Spreng., Featherman s.n. 
(LSU) Pr 

Gratiola brevifolia Raf, ae : 

Gratiola pilosa Michx., N 1 

Gratiola ramosa Walt., oe oy (MO 

Gratiola virginiana L., N 460 D,S 

ia dubia (L.) Pennell var. anagallidea 

Michx.) Cooperrider, N 1454 D, FM, Pi, S 

Mazus pumilus (Burm.f) Steenis,* N 368 D, B 

Mecardonia acuminata (Walt.) Small, N 7703 D, 

M, Pi, 

Mecardonia procumbens (P.Mill.) Small, N 7490 D, 

S 


D,FM 


= 


— 


Micranthemum umbrosum (J.F. Gmel.) Blake, N 
6 


Mimulus alatus Aiton, N 902 § 

Nuttallanthus canadensis (L.) D.A. Sutton, N 563 
D 

Pediculari ad L. Pennell 1012 (NO) B 

Penstemon laxiflorus Pennell Small, N 1248 Pi 

Seymeria cassioides (Walt.) Blake, N 1525 Pi 

Scoparia dulcis L., N 1080 D, Pi, Pr, S 


Veronica ee Poir.* N 472 D 


SOLA 

Cali ee par iflora (Juss. i D’Arcy., N 1530 BM 
Lycium barbaru 

Physalis ene L, NI R 

Physalis cordata P.Mill. a & Correll 9566 (LSU) 
Phy. realic nifheccenc LN 26 D, PR 
Solanum carolinense su N 754 D,Pr 

Solanum pt) chanthium Dunal, N 766 D, Pr 
SPHENOCLEACEAE 

Sphenoclea zeylanica Gaertn.,* N 343 D, Pr 
STERCULIACEAE 

Melochia corchorifolia L.,* Ns.n.D 


Holes i fee ti hes e 


B,S 
Styrax grandifolius rey ce s.n. (LAF) B 
SYMPLOCACEAE 
Symplocos tinctoria (L.) L'Hér., N 465 B,S 


TAMARICACEAE 
Tamarix gallica L.,* N 1303 BM, D 


SIDA 19(2) 


TILIACEAE 
Tilia americana L.var.caroliniana (P.Mill.) Castigl., 
N 1558 Pi 


ULMACEAE 

Celtis laevigata Willd., N 1043 B, Pr,S 
Planera aquatica J.F.Gmel., 195B,S 
rile alata Michx.,N 611 B,P 

Ulmus americana L.,N&M 1 720 (MCN) B 


URTICACEAE 

Boehmeria cylindrica (L.) Sw., N318B,S 
Parietaria floridana Nutt., Montz 3030 (LSU) BM 
Pilea pumila (L.) A.Gray, N 1106 B,S 

Urtica sn te Pursh,N 1729D 


VALERIANACEAE 
Valerianella Piet (L.) Dufr., N 7340 D, Pr 


VERBENACEAE 
Callicarpa americana L., N 272 B, Pi, Pr, S 
Glandularia canadensis (L.) Small, Milsom 18 
D, Pi 
Glandularia pulchella (Sweet) Troncoso.,* N 1224 
D 


Lantana camara L..*N1116D 
Phyla Fi eae (Michx.) Greene, N 1369 BM, D, 


FM, P 
as dee flora (L.) E.Greene, N 1366 BM, D, FM, 


oe brasiliensis ae *N630D 
Verbena halei Small, N 

Verbena rigida Montes N544D 

Verbena scabra Vahl, N898 BM, FM 
Verbena xutha Lehm., Montz 2294 (LSU) D 


VIOLACEAE 
Viola affinis Leconte, N 1150 B,S 
Viola lanceol ata L, N 1252 Pi 
Viola n reene, tli 7 (LSU) B 
Viola pedata L Kina 7 (MCN) P 
iola X primulifolia (pro sp.), Nat 18 Pi 
iola pela Aiton, Cs.n. Pi 
| 
fol 


VISCA 

Phoradento ere aa (Raf.) Reveal & M.C. 
Johnston, N 1144 

VITACEAE 


Ampel yn arhoren 


L.) Koehne, N 612 B, FM, S 
Parthenocissus quinoueroli (L.) Planch., N 387 D, 
i, Pr 


Vitis cinerea Engelm., N 930 B 
Vitis mustangensis Buckl., N 702 D, BM 
Vitis rotundifolia Michx., N 533 B, Pi, S 
Vitis vulpina L., N 265 B, D, Pi 


NEYLAND ET AL., FLORA OF CALCASIEU PARISH, LOUISIANA 


MAGNOLIOPHYTA (Liliopsida) 
AGAVACEAE 
Manfreda virginica (L.) Salisb. ex Rose, N 782 D, 

i, Pr 
Yucca aloifolia L., N 998 D 
ALISMATACEAE 
Echinodorus cordifolius (L.) Griseb., N 1078 D, FM, 
Echinodorus berteroi (Spreng.) Fassett, N 1072 D, 
FM 


var.calycina, Th 30114 


Sagittaria calycina E 
D,FM 


Sagittaria pees L., N 1442 BM, D, FM 

Sagittaria graminea Michx., N 345 D, FM, Pr 

Sagittaria latifolia Willd., N 727 D, FM 

Sagittaria papillosa Buch., Tallman 19 (LSU) D 
FM 


Saaittaria platy phy Ilq Engelm., N 312 BM, D, FM 


ARACEAE 

Arisaema dracontium (L.) Schott, N 493 B 

Arisaema triphyllum (L.) Schott ssp. quinatum 
(Buckl.) Huttleston, N 500 B 

Colocasia esculenta (L.) Schott,* N 1037 D, FM 

Peltandra virginica (L.) Schott, N 492 D, FM, $ 

Pistia stratioites L.,N 869 D, FM 


ARECACEAE 
Sabal minor Jacq.) Pers., N638S 


Carex basiantha Steud., N&M 1728 (MCN) B 
Carex cherokeensis Schwein., N& M 1724 (MCN) 


Carex complanata Torr.& Hook., N 604 B 
Carex debilis Michx., N& M 1727 (MCN) B 
Carex di ioe Willd., M& N 2772 (MCN) B 
Carex wes osperma Dewey, N 553 Pr 


Car ankii Kunth, N 637 D, Pr 
carey glaucescens EIl., N 802 B, D, Pi 
x intumescens Rud dge, N 603 B 
Cre oo pes T 20606 B, FM 
ex leavenworthii Dewey, N& M 17175 (MCN) D 
Ese aulina Willd., Th 14405 D, Pr 
Carex microdonta Torr.& Hook., M& N 2765 (MCN) 


ps oxylepis Torr. & Hook, N&M 1723 (MCN) B 
I, M&N 2775 (MCN) 


B 
Carex styloflexa Buckl., N& M 1725 (MCN 
Carex verrucosa Muhl., Orzell & Bridges (N i‘ BM, 


Carex ales Michx., A 10027 (LAF) D, Pr 
ladium mariscu 


s (L.) Pohl ssp. jamaicense 
(Cran wk uke N 747BM,FM 
Dee N 984 BM, D, FM, S 


Cyp Th 66923 D, Pr 
Cyperus croceus ate N 745 “i Pr 
ig Neh us elegans L bea 
Cyperus oekler, Carter 8! 30 (VDB) D 
f s Muh, N 866 BM, FM 


Sabal palmetto (Walt.) Lodd.ex Schult. & Schult.* 
1506 D 


BROMELIACEAE 
Tillandsia recurvata (L.) L., N 1846 E 
Tillandsia usneoides (L.) L., N 1376 E 


BURMANNIACEAE 
Apteria aphylla (Nutt.) Barnh., N 1627 Pi 


COMMELINACEAE 

Commelina diffusa Burm.f., N 978 B, D, FM 
Commelina erecta L., N 809 D, Pr 

Commelina virginica L.,N 901 B,S 

Tradescantia hirsutiflora Bush, N 1207 D, Pr 
Tradescantia occidentalis (Britton) Smyth, Th 


= 


89238 Pr 
Tradescantia ohiensis Raf., N 1220 BM 
Tradescantia reverchonii Bush, Orzell & Bridges 
7112 (NLU) P 


CYPERACEAE 
Carex abscondita Mackenzie, N& M 1726 (MCN) 
B 


Carex albicans Willd. var. australis (Bailey) J.Rettig, 
M&N 2771 (MCN) B 

Carex annectens (Bickn.) Bickn.,M&N2766 (MCN) 
Pr 


Cyper us esculentus L., N 805 D, FM 
Cyperus flavescens L., N 1077 D, Pi 


Cyperus odoratus L., N 694 

Cyperus oxylepis Nees 2 seu, es D,FM 

Cyperus pilosus Vahl,* 

ee po fakin eae var. texensis (Torr.) 
n.,N 


= 


Steud., N 970 D, Pr 
Be retrorsus Chapm., N 834 D, Pr 
Cyperus rotundus L.,* N 1013 D, Pr 
Cyperus strigosus L, N91] 


Cyperus vi D,S 
neni Poe (L ) Roem.& Schult. 7h 87978 


Eleocharis agi (Torr.) Chapm., N a D 
rpa Torr, N615 D,P 
Eleocharis montana (Kunth) Roem. & Se huile, N 
05 D, Pr 


lo hoaric 


_N 528 D, Pr 
Eleocharis obtusa anita ane 7129415 D, Pr 


378 


Fleocharis parvula (Roem.& Schult.) Link, N 1034 


Eleocharis oe (Michx.) Roem. & 
Schult. N587 D 

Eleocharis ae Torr., Th 48235 Pi 

Eleocharis tuberculosa (Michx.) Roem. & Schult, 
N 949D,S 


“ve annua (All.) Roem.& Schult., N 964 D, 


aa wi autumnalis (L.) Roem. & Schult., N 
lis carolinia m.) Fern., ae a 

Fimbristyli s miliacea : ) ce N 863 P 

Fimbristylis puberula (Michx.) Vahl, Th 301 09D,Pr 


Fuirena breviseta (Coville) ae N oe D,FM 

Fuirena squarrosa Michx., N 950 FM 

Isoleppis carinata Hook. & Arn. ex Torr, N 1745 D, 
>) 


r 
Isolepis molesta (M.C. Johnson) S.G.Sm., 1 30614 
BM 


Kyllinga brevifolia Rottb., Th 66924 D, Pr 
Kyllinga odorata Vahl Lievens 1265 (LSU) D, FM 
Kyllinga pumila Michx., N 913 D, Pr 
Rhynchospora caduca Ell., N 908 Pr 
Rhynchospora cephalantha A. Gray, Orzell & 
Bridges 7404 (NLU) P 
Rhynchospora colorata (L.) H. Pfeiffer, Watson 90 
(MCN) D, Pi, Pr 
Rhynchospora corniculata (Lam.) A. Gray, Joyce 
407 (LAF) D, BM 
Rhynchospora divergens Chapm. ex M.A. Curtis, 
oe s.n. (LSU) Pi, S . : 
lliottii xD r., Th 14366 D 
myncospor fasci me (ane Vahl, N 


ie ee filifolia A. Gray, Orzell & Bridges 
LU) Pi 
oe globularis (Chapm.) Small, N 527 


ee aie glomerata (L.) Vahl., 7 31652 D, Pi 
Rhynchospora gracilenta A. Gray, N& M 1596 Pi 
es inexpansa (Michx.) Vahl, N 723 D, 


Mynchospr latifolia (Baldwin ex Ell.) Thomas, 
D, Pi, Pr 

Rhynchospora macrostachya A. Gray, N 969 D,& 

Rhynchospora microcarpa Baldwin ex A. Gray, 
Orzell & Bridges 7120 (LSU) D, FM 

Rhynchospora_ miliacea (Lam.) 
Featherman s.n. (LSU) B,S 

Rhynchospora mixta Britton ex Small, N 907 $ 

Rhynchospora nitens (Vahl) A.Gray, Cs.n. Pi 


A. Gray, 


SIDA 19(2) 
Rhynchospora oligantha A. Gray, N& M 1594 
(MCN) Pi 


Rhynchospora perplexa Britton, Th 89309 Pr 

Rhynchospora plumosa Ell. Joyce 288 (LAF) Pi 

Rhynchospora pusilla Chapm. ex M.A. Curtis, B 
13825 (NLU) D, BM 

ee rariflora (Michx.) Ell, Th et al.89449 


wa 


te ee tracyi Britton, C s.n. Pi, S (SH) 
Schoenoplectus maritimus (L.) Lye, T 30614 BM, 


Schoenoplectus robustus (Pursh) M.T. Strong, N 

, BM 

Schoenoplectus tabernaemontani (K.C. Gmel.) 
Palla, Montz 6497 (LSU) D 

Scirpus californicus (C.A.Meyer) Steud., Th 89218 
D, BM, FM 


cirpus cyperinus (L.) Kunth, 131682 FM, $ 
oe baldwinii (Torr.) Steud., Orzell & Bridges 
6870 (NLU) D, Pi 
ae ciliata Michx.,A 15119 Pi 
Scleria orgiana Core, B 13828 (NLU) Pi, S 
Scleria oligantha Michx. Joyce 405 (LAF) Pi 
Scleria pauciflora Muhl. ex Willd. Th 129557 Pi 
Scleria reticularis Michx., Pias ay 445 (LAF) Pi,S 
Scleria triglomerata Michx., N 1748 Pi 
Scleria verticillata Willd., bra | a Bridges 5/759 
(NLU) Pi, S 
DIOSCOREACEA 
Dioscorea Te : N 1264B 


EN LORAUENCEAE 


gulare \., N 796 
Lachnocaulon an ceps eee N 1219 Pi 
HYDROCHARITACEAE 


Egeria densa Planch.,* N 720 M,S 
Limnobium spongia (Bosc) Sone NI 1025 BM, 


Bae id alismoides Le ) ro 7 | 1007 BM 


N 1467 BM 
HYPOXIDACEA 
pee hirsuta (L core N 334 B, D, Pi 
IRID 


ees lahue (Molina) Goldblatt ssp. caerulea 

(Herbert) Goldblatt, N 7273 D, Pr 
Alophia drummondii (Graham) R. C. Foster, We- 

ber 26 (MCN) Pi 

Iris brevicaulis Raf. Whatley 2197 (LAF) FM 

Iris aaa = eee 29 (MCN) D 

Iris virginica L., N 

Sisyrinchium spear : Mill, N& M 1 

CN) B 

Sisyrinchium atlanticum Bicknell, N 526 D, Pi, Pr 


= 


NEYLAND ET AL., FLORA OF CALCASIEU PARISH, LOUISIANA 


Sisyrinchium rosulatum Bicknell, N 7791 D 
Sisyrinchium sagittiferum Bicknell, N 507 D, Pr 
JUNCACEAE 

Juncus acuminatus Michx.,, Giltner 393 (MCN) D, 


Juncus 5 biflorus Ell., Th 8924 
June a oe eG Ghee 608 (MCN) 


ee ae L., 723506 D, Pi, Pr 
Juncus dichotomus Ell, Brooks 610 (MCN) D, Pi, 
Pr 
diffusissimu ckl., N 722 BE i Pr 
Juncus dudleyi eee Th tea 
Juncus effusus L., N 543 D, F 
Juncus elliottii Chapm,, oe i (MCN) D, Pi, 


Juncus marginatus Rostk., Giltner 516 (MCN) D, 
Juncus Lee Coville, Giltner 391 
tee polycephalus Michx., Giltner 523 (MCN) D, 
hee repens Michx., Giltner 522 (MCN) D, Pi, Pr 
Juncus roemerianus Scheele, Giltner 487 (MCN 


= 


MCN) D, FM, 


SS 


Juncus scirpoides Lam., N 550 D, Pi, Pr 

Juncus tenuis Willd., N 756 D, S, Pr 

Juncus validus Coville, N 755 D, FM, Pi 

LEMNACEAE 

Lemna minor L., N 10878 

Lemna valdiviana Phil., N 744. 

Spirodela polyrrhiza (L.) Schleid., N 1086 § 

sepa punctata (G.F.W.Mey.) C.H.Thompson, 
8S 


Wee ek Weddell, N 1208S 
Wolffia columbiana Karst.,N 1751S 
Wolffiella lingulata (Hegelm.) Hegelm., N 1154S 


LILIACEAE 
Aletris aurea Walt., N 565 Pi 
Aletris farinosa L., N 566 Pi 
Allium canadense L. var. canadense, N 1228 B, D, 


Allium canadense var.mobilense (Regel) Ownbey, 
1297 B,D, Pr 

Cooperia drummondii Herbert, Th 66940 Pr (S1/ 

Crinum americanum L., N 785 BM, FM, S 

Habranthus tubispathus (L'Hér.) Traub, N 1784 D 

Bans lirlosome (Raf.) Shinners, N 1196 
BM, F i,S 

ena fee LN & Komo 1777 Pi 


Nothoscordum bivalve L., N 1089 D, Pr 
Schoenolirion croceum (Michx.) A. Gray, N 1218 
Pi 


379 


Trillium gracile Freeman, N 455 B 
Zephyranthes grandiflora Lindl.,N 1417 D 


NAJADACEAE 

Najas guadalupensis (Spreng.) Magnus, Haynes 
2504 (LAF) D, BM, FM 

ORCHIDACEAE 

Calopogon oki ee D.H. Goldman, Pennell 
10242 (NY) P 

Calopogon eed .) BSP, N 1341 Pi 

Corallorrhiza wisteriana Conrad, N 1706 B 

Epidendrum conopseum R. Br., Correll & Correll 
9660 (DUKE) E 

Habenaria repens Nutt., N 974 $ 

Listera australis Lindl., N 1689 B 

Platanthera ciliaris (L.) Lindl., N 1497 Pi 

Platanthera cristata (Michx.) Lindl., N 1790 Pi 

Platanthera flava (L.) Lindl., N& Komo 1781 Pi 

Platanthera nivea (Nutt.) Luer, N 1410 Pi 

Pogonia ophioglossoides (L.) Ker-Gawl., N 1336 Pi 

Spiranthes cernua var. odorata (Nutt.) Correll, N 
348 S$ 


Spiranthes longilabris Lindl., N& Komo 1804 Pi 
Spiranthes praecox (Walt.) S.Watson, N 582 Pi 
aa ae Engelm.& A. Gray, N 669 BM, 


rsd Ze (Pursh) Nutt., N 1624 B 


POACEA 
Agrostis ae is (Walt.) B.S.P., N 548 D 
eee? gerardii Vitman, Brooks 595 (MCN) 


Pi, 

Pen glomeratus (Walt.) B.S.P. var. 
glomeratus, N 1056D 

Andropogon gyrans Ashe var.gyrans, Bet al.8674 
Pi 


Andropogon ternarius Michx., 7 31890 Pr 
Andropogon virginicus L.,T 27991 D 
Anthaenantia rufa (Nutt.) Schult, Brooks 794 


N) Pi 
Anthaenantia villosa (Michx.) P. Beauv, Brooks 594 
(MCN) Pi 


Aristida longispica Poir., T 28008 D, Pi, Pr 
Aristida oligantha Michx., T 9644 D, Pi, Pr 

fore palustris (Chapm.) Vasey, N 1600 Pi 

stida ale Poir, Brooks 802 (MCN) D 
- 

Arun anne gigantea (Walt.) Muhl., Nee, Es Pi 

Axonopuls fissifolius (Raddi) Kuhlm 

Axonopus fae eee! Hitchc,, ei 8668 


Brtichoa exaristata (Nash) Henrard, Th 98153 
D, Pr 


Bothriochloa ischaemum (L.) Keng,* N 1018 
Bothriochloa laguroides (DC.) Herter ssp. 


torreyana (Steud.) Allred & Gould, Th 66885 
D 


Bothriochloa een (Gould) Allred & 
Gould, Th 8 D 

Bouteloua inate Lag., Cs. SR) 

Brachiaria platyphylla aa. ex Wright) Nash, 
N828D 


Brachiaria texana (Buckl.) S.T. Blake, Th 66891 
oled mi inor eae Costanza 3 Neg 

Vahl,* 

Bro omus japoni icus Th ae ex ae A 4608 D 
oe ] ee (Nutt.) Engelm.,* Chase 6114 


aa incertus M.A. Curtis, N 912 
Chasmanthium latifolium (Michx.) Yates, N 1057 


oem laxum (L.) Yates var. laxum, Th 
14372 B,S 

Chloris canterai Arechav.,* McKenzie 223 (LSU) D 

Chloris virgata Sw., N 

Cinna arundin 

Coelorachis rugosa (Nutt) Nash, A 16163 D, Pi 

Coelorachis cylindrica (Michx.) Nash, B 13871 
(NLU) D, Pi, Pr 

Cynodon dactylon (L.) Pers.,* N 537 D 

Dactyloctenium aegyptium (L.) P Beauv.,* N 894 


Nichanthels q ula 


ar rs ex Poir.) Gould & 
CA. Clark, Betal. 5646 Pi, P 
Dichanthelium acuminatum Sw) se & CA. 
Clark var.acuminatum 
Dichanthelium acuminatum (Sw.) Gould var. 
eet (Nash) Gould & C. A. Clark, A 4601 


an commutatum Schult. N317D 

Dichanthelium dichotomum (L.) Gould var. 
dichotomum, A 16403 Pi 

Dichanthelium dichotomum (L.) Gould var. 
ensifolium (Baldw. ex Ell.) C. A. Clark, A 3596 
Pi 


Dichanthelium dichotomum (L.) Gould var. tenue 
(Muhl.) Gould & C.A. Clark, 725624 Pi 

Dichanthelium laxiflorum (Lam.) Gould, B et al. 
8598 D 


Dichanthelium leucothrix (Nash) Freckman, A 
4419 (LAF) Pi 

Dichanthelium scabriusculum (Ell.) Gould & CA. 
Clark, N 586 Pi 

Dichanthelium scoparium (Lam.) Gould, N 598 D 

Dichanthelium sphaerocarpon (Ell.) Gould var. 
sphaerocarpon, N 498 D 

seer teed strigosum (Muhl.ex Ell.) Freckman 

ucoblepharis (Trin.) Freckman, B 8669 Pi 
e Highs (Retz.) Koeler, A 4593 (LAF) D 


SIDA 19(2) 


Digitaria cognata (J. A. “ti Pilger var. 
cognata, Bet al. 8661 

Digitaria filiformis (L.) Koclee Brooks 792 (MCN) 
Pi 


Digitaria ischaemum (Schreb. ex Schweigger) 
Muhl.,* N 7094 D 
Digitaria violascens Link* N.& Komo 1619D 


Echinochloa crus-galli (L.) P. Beauv, *N852D 
Echinochloa muricata (P. Beauv.) Fern.,* Brooks 
300 (LSU) D 
Echinochloa walteri (Pursh) Heller,* N 865 BM, 
FM,S 
Eleusine indica (L.) Gaertn.* Gaskin 30 (MCN) D 
Elymus virginicus L., N696 D 
Eragrostis bahiensis Schrad.,* A 4432 (LAF) D 
Sago barrelieri pee * Th 66893 D 
Th 66892 Pi, Pr 
Eragrostis glomerata ie L.H. Dewey, N 7 
B,D 


Eranr, amnectr 


Eragrostis hirsuta (Michx.) Nees, | 20620 D, Pr 

Eragrostis hypnoides (Lam.) B.S.P, N 970 B,D 

Eragrostis lugens Nees, Bet al. 8622 D, Pr 

Eragrostis pectinacea (Michx.) Nees, N 970 B,D 

Eragrostis refracta (Muhl.) Scribn., B et al. 8652 D, 
Pi 


Eragrostis secundiflora Presl|, N 939D 
Fragrostis spectabilis (Pursh) Steud., N 400 D, Pr 
Eremochloa ophiuroides (Munro) Hack.* N & M 
1627 (MCN) D 
Eustachys caribaea (Spreng.) Herter* McKenzie 
14(LSU) D 


Eustachys petraea (Spreng.) Herter, N 985 D 
Festuca arundinacea Schreb.* N1117D 


Leersia lenticularis Michx,, N Le D 

Leersia oryzoides (L.) Sw., N 

ee fascicularis ere : Gray, Th 66898 
BM 


Petit mucronata (Michx.) P.Kunth, Th 66899 
B,D 


Leptochloa nealleyi Vasey, N 842 BM 
Leptochloa uninervia (Presl) Hitchc. ‘i Oe Th 
1568D 


Limnodea arkansana (Nutt.) L.H. Dewey, s. nom. 

s.n. (LAF) D 
lium arundinaceum S.J. Derbyshire,* Edgerton 

s.n. (LSU) D 

Lolium perenne L..* N1772D 

Melica mutica L., N 444 Pi 

Muhlenbergia capillaris (Lam.) Trin., A 16179 Pi, 
Pr 


NEYLAND ET AL., FLORA OF CALCASIEU PARISH, LOUISIANA 

Oplismenus setarius (Lam.) Roem. & Schult.,* 7 
20534 Pi 

Oryza sativa L.* N 891 D, Pr 


Panicum anceps Michx. var.anceps, N 797 D 
Panicum brachyanthum Steud, Brooks 795 (MCN 
D 


= 


Panicum dichotomiflorum Michx., N 893 D 

Panicum repens L., N 989 BM, D, FM 

Panicum rigidulum var. pubescens (Vasey 
LeLong, Bet al. 8677 Pi 

Panicum rigidulum Bosc ex Nees var. rigidulum 
Gilmore & Smith 3461 (NLU) D, FM, Pi 

Panicum verrucosum Muhl., N 7 oe 7 Pi 

Panicum virgatum L., N 836 BM 

Paspalum bifidum (Bertol.) ee Brooks 807 

CN) Pi 


SS 


Paspalum boscianum Fluegge, B 5370 D 
Paspalum eonsoailin Bergius, 7 20530 D 

im Poir.,* Augustus 50 (MCN) D 
Pes balun distichum L., N 992 BM, D, FM 
es floridanum Michx.,A 16138 (LAF) D, Pi, 


ee hydrophilum Henrard,* N 854 D 

Paspalum laeve Michx., Augustus 80 (MCN) D 

eee iad Trin., TA te BM, D, FM 

pau mm che eat 1316 

An pa ar. saurae eee 
79 (MCN) E D 

Paspalum plicatulum Michx., Augustus 117 
D 


(MCN) 


Paspalum praecox Walt, Brooks 805 (MCN) Pi, Pr 
Paspalum setaceum Michx., B et al. 8611 D 
Paspalum urvillei Steud.,* N 940 D 

Pennisetum glaucum (L.) R.Br.* N 779 D 
Phalaris angusta Nees ex Trin., Dutton & Taylor 


pnalails eaouniang Walt., N 551 D 
op) Vasey, N 1108 B, S 
f ee australis is (Cav) Tin, N'787 BM 
Poa annua A 
Poa autumn able ex Ell., Griggs 17 (LSU) B 
Polypogon es (L.) Desf.,* N 1222 D 
Rottboellia tae (Lour.) W. Clayton,” 
a n.( C ) D 
rum lwinii Spreng., N 1052 D, Pi 
Saccharu rum gi iganteum (Walt.) Pers., N 1047D 
Sacciolepis striata (L.) Nash, B et al. 8679 D, Pi 
Schizachyrium scoparium (Michx.) Nash, Brooks 
806 (MCN) D, P 
Schizachyrium tenerum Nees, Th 98157 Pi 
Setaria faberi Herrm.,* T 28865 D 
Setaria parviflora (Poir.) Kerguelen, N 945 D 
Setaria pumila (Poir.) Roemer & Schult.,* 
McKenzie & Urbatsch 135 (LSU) Pr 


381 


Sorghastrum elliottii (Mohr) Nash, N 1533 D, Pi 
Sorghastrum nutans (L.) Nash, 727992 D, Pi, Pr 


Spartina alterniflora Loisel., N 1153 BM 
Spartina patens (Aiton) Muhl., N 1408 BM 
Spartina spartinae (Trin.) Hitche., N 1557 BM, Pi 
Sphenopholis obtusata (Michx.) Scribn., A & Vin 
cent 10022 (LAF) D, Pi 
seen compositus (Poir.) Merr. var. 
positus ,B8 8658 BM, FM, Pr 
Sporabal compositus var. eee (Trin.) 
Kartesz & Gandhi, B 8620 is 
Hele Gt indicus (L.) R.Br, N iD 
bolus junceus (Michx.) ae N 1608 D, Pi, 


ee silveanus Swall., 7 31899 Pi (S2/S3) 
Steinchisma hians (Ell.) Nash, N601 D 
Stenotaphrum secundatum (Walt.) Kountz, N 705 


Tridens ambiguus (Ell.) Schult. 7h 98149 Pi, Pr 
Tridens flavus (L.) Hitchc., B et al. 8583 D, Pi 
Tridens strictus (Nutt.) Nash, N 7552 D 


7 

Vulpia octoflora (Walt.) Rydb., N 552 D 

Zizania aquatica L., N 1459 BM, FM 

Zizaniopsis miliacea (Michx.) Doell. & Asch., N 
1008 BM, FM, S 


EAE 


P 
Fichhornia iL .) Solms,* N 1409 D,BM, 
Heteranthera limosa (Sw.) Willd., N 845 D, BM, FM 
Pontederia cordata L., N 266 D, F 


POTAMOG 

Promeaeto ie a. Raf., N 799 D,FM,S 
Potamogeton nodosus Poir., N 1047 FM, BM, S 
pales busi us L.,N 1393 BM, FM 
SMILACACEAE 

Smilax bona-nox L., N 486 B, D, Pi, Pr, S 

Smilax glauca Walt., N& M 1630 (MCN) B, D 
Smil ax | laurifolia LN 1413 Pi, 


Smilax tamr a N&M 
srnilax vaked Pursh, N 731 = S 
TYPHACEA 
Typha angustfo Lae 1364 BM, FM 
s, N 1318 BM,D 
Typha latifolia LN Bieb: FM,S 
XYRID 
Xyris Hei Bey.ex Kunth, N 795 D, Pi 


minnoenc 


382 


Xyris baldwiniana Schult., N 1768 Pi 

Xyris caroliniana Walt., N i 

Xyris difformis var. curtissii (Malme) Kral, N & M 
1614 (MCN) Pi 

Xyris difformis Chapm. var. difformis, Th 14369 D 
Pi 


~ 


Xyris drummondii Malme (MacRoberts & 
MacRoberts 3905 MCN) Pi 


SIDA 19(2) 


Xyris fimbriata Ell. hie 7 se 7407 (LSU) Pi 
Xyris jupicai Rich., NI 
Xyris laxifolia var. ir irae Ghee N 335 Pi, 


ie 5 Chapm.,N8 
torta Sm.in Rees, ae i 1592 (MCN) Pi 


APPENDIX II 


Taxa reportedly collected in Calcasieu Parish by either MacRoberts (1988) or by Thomas 
and Allen (1993 1996 1998) that were not discovered during our field research nor lo- 
cated during our visits to other herbaria are listed.Taxa are listed alphabetically by family, 


genus, and species within each division. 


APPENDIX II 
UNVERIFIED TAXA REPORTED FOR CALCASIEU PARISH 


DRYOPTERIDACEAE 

Woodsia obtusa ees Torr. 
ACANTHACEA 

Ruellia Pais ate (Nees) A. Gray 
Ruellia strepens 


NNONACEAE 
Asimina triloba (L.) Dunat 
APIACEAE 
Polytaenia nutta 


nels 


li DC. 


Chloracantha spinosa (Benth. YN 

Cirsium carolianum oe Fern. & aie 

Coreopsis gladiata W. 

Coreopsis pubescens Ell var. debilis (Sherff) E.B. 
Smith 

ne angustifolia DC. 

Echinacea pallida (Nutt.) Nutt. 


x 


Euthamia graminifolia (L.) Nutt. 
fires simulans E EE: ee 


Symphyotrichum subulatum (Michx.) Nesom 


BRASSICACEAE 

Lepidium sa Schrad. 
Sinapis arv 
CALLITRICHACEAE 
Callitriche fats Nutt. 


Lobelia nuttallii i. ales 

CISTACEAE 

Helianthemum carolinianum (Walt.) Michx. 
Lechea minor L. 

Lechea r losa Michx. 

Lechea tenuifolia Michx. 


CONVOLVULACEAE 
Convolvulus arvensis L. 


ERICACEAE 
Vaccinium corymbosum L. 


E 

Baptisia nuttalliana Small 
Desmodium glutinosum (Willd.) Wood 
Desmodium obtusum (Muhl. ex Willd.) D 
Desmodium perplexum Schub. 
Kummerowia striata (Thunb.) Schind. 

Neptunia pubescens Bent 
Trifolium cate nianum Michx. 
GENTIANACEAE 
Sabatia stellaris Pursh 
Sabatia angularis (L.) Pursh 
Sabatia brachiata Ell. 
CLUSIACEAE 
Hypericum galioides Lam. 


NEYLAND ET AL., FLORA OF CALCASIEU PARISH, LOUISIANA 


Hypericum gentianoides (L.) B.S.P. 
Hypericum gymnanthum Engelm. & A. Gray 
Hypericum nudiflorum Michx. ex Willd. 


EUPHORBIACEAE 
lates hia (L ) Millsp. 
amaesy Aiton) Small 
a indheimenianus Scheele 
Croton monanthogynous Michx. 
Euphorbia spathulata am. 
ma eue x. 
Phyl lanthus Coe Walt. 
LAMIA 
Monarda oe 
Physostegia een (Nutt.) Engelm.& A. Gray 
Pycnanthemum muticum (Michx.) Pers 


Scutellaria parvula Michx. 

LENTIBULARIACEAE 

Utricularia ak L. 

LINACEA 

Linum aoe ook 

Linum flor ae ane ) Trel. 

Linum sulcatum Ridde 

LOGANIACEAE 

Spigelia marilandica L. 

LYTHRACEAE 

Lythrum lineare L. 

MALVACEAE 

Callirhoe involucrata (Torr.& A. Gray) 
ACEAE 

Boerhaavia diffusa L. 


ONAGRACEAE 

Ludwigia repens J.Forst. 
OXALIDACEAE 

Oxalis violacea L. 
PLANTAGINACEAE 
Plantago rugelii Decne. 
Plantago wrightiana Dene. 
POLYGALACEAE 

Polygala curtisii A. Gray 
Polygala cymosa Walt. 
Polygala grandiflora Walt. 
Polygala lutea L 
POLYGONACEAE 
Polygonum convolvulus L. 
RANUNCULACEAE 
Delphinium virescens Nutt. 
Ranunculus laxicaulis Darby 
Ranunculus platensis Spreng. 


RHAMNACEAE 
Ci aAnothiic americantis< 


ROSACEA 

Cra ataegus aestivalis L. 
Crataegus crus-galli L. 
Crataegus sabineana Ashe 
saa spathulata Michx. 


aie pet ia Marshall 


RUBIACEAE 
Galium pilosum Aiton 
Galium trifidum L. 

a ium uniflorum Michx. 
Houstonia caerulea 
Houstonia purpurea : 
SCROPHULARIACEAE 
Agalinis pinetorum Pennell 
Castilleja indivisa Engelm 
Gratiola neglecta Torr. 
Verbascum virgatum Stokes 
SOLANACEA 
Physalis heterophylla Nees 
Solanum americanum P. Mill. 
Solanum eleagnifolium Cav. 
TAMARICACEAE 
Tamarix ramosissima Ledeb. 
URTICACEAE 
Parietaria pensylvanica Muhl. ex Willd. 
VERBENACEAE 
Verbena bonariensis L. 
Verbena litoralis Kunth 
VIOLACEAE 
Viola langloisii Greene 
AGAVACEAE 
Yucca filamentosa L. 
CYPERACEAE 
Carex leptalea Wahl 
Cyperus polystachyos ae var. polystachyos 
Eleocharis elongata Cha 

Eleocharis macrostachya 
Eleocharis palustris (L.) Poe & Schult. 
IRIDACEAE 
Sisyrinchium langloisii Greene 
LEMNACEAE 
Lemna obscura a Daubs 
ORCHIDACE 
Calopogon eae: (Walt.) Ames 


383 


384 SIDA 19(2) 


Platanthera integra (Nutt.) A. Gray ex Beck Digitaria sanguinalis (L.) Scop. 
Spiranthes cernua (L.) L.C. Rich. Digitaria villosa (Walt.) Pers. 
Spiranthes ovalis Lindl. Eragrostis elliottii S. Wats. 


Hackelochloa granularis (L.) Kuntze 


PONCE AE It rn virgini nica le 


Schu 
Aristida ramosissima Engelm. ex A. Gray 
Biamnuehacemosus is um langei (Fourn, ) Nash 


Dichanthelium ovale var. addisonii (Nash) Gould Paspalum repens se 
acleee Paspalum vagina 
Saccharum ee (Mich) Pers. 


Dichanthelium ovale var. ovale (Ell.) Gould & C.A. 
Clare Triticum aestivum 


lar 
Dichanthelium villosissimum 


— 


Nash) Freckman 


APPENDIX III 
REJECTED TAXA PREVIOUSLY REPORTED FOR CALCASIEU PARISH 
APIACEAE LAMIACEAE 
Ptilmnium nuttallii (DC) Britt. Physostegia pulchella Lundell 
ASTERACEA Salvia reflexa Hornem 
rah spicata i .) Willd. LAURACEAE 
arshallia obovata ig Beadle Persea borbonia (L.) Spreng. 
rude fulgida A . RANUNCULACEAE 
Al a cnet at Anemone berlandieri Pritz. 

Solidago radula Nutt. 
EUPHORBIACEAE Ss Michx. 
Cnidoscolus stimulosus (Michx.) Engelm.& A.Gray 

ACENIACEAE 


FABACEAE 

Baptisia bracteata os ex Ell. var. leucophaea 
Lespedeza capitata Mich 

JUGLANDACEAE 

Carya texana Buckl. 


Sarracenia flava L 


CEAE 
Aristida longispica Poir.var.geniculata (Raf.) Fern. 


ACKNOWLEDGMENTS 


We thank R. Dale Thomas (NLU), Charles Allen (NLU), Garrie Landry (LAF), and Anne S. 
Bradburn (NO) for assisting us in locating specimens at their respective herbaria; Latimore 
Smith (Louisiana Natural Heritage Program) for his technical assistance; Paul Kitt and 
Robert Rumsey (McNeese State University) for helping us locate several uncommon spe- 
cies, and Heather Harrington for her curatorial work. Funds were provided, in part, by 
McNeese State University through the Shearman Research Initiative Fund. 


REFERENCES 


AtLen, C.M. 1992. Grasses of Louisiana 2nd Edition. Cajun Prairie Habitat Preservation Soci- 
ety, Eunice LA. 

Brioces, E.L.and S.L.Orzell.1989. Longleaf pine communities of the West Gulf Coastal Plain. 
Nat. Areas J.9:246-263 


NEYLAND ET AL., FLORA OF CALCASIEU PARISH, LOUISIANA 385 


Brummit, R.K. and C.E. Powett. 1992. Authors of plant names. Royal Botanic Gardens, Kew. 

Corrett, D.S.and H.B. Corrett. 1941.A collection of plants from Louisiana. Amer. Midl. Natu- 
ralist 26:30-64. 

Corrett, D.D. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner, TX. 

Cronauist, A. 1980.Vascular flora of the southeastern United States, Vol. 1. Asteraceae. Uni- 
versity of North Carolina Press, Chapel Hill. 

Duncan, W.H. 1975.Woody vines of the southeastern United States. The University of Geor- 
gia Press, Athens. 

Fenneman, M.N. 1938. Physiography of eastern United States. McGraw Hill Book. Co. Inc., 
New York & London. 

Flora of North America Editorial Committee. 1993. Flora of North America north of Mexico. 
Vol. 2. Pteriodophytes and Gymnosperms. Oxford University Press, New York & Oxford. 

Fiora oF NortH America EpitoriaL Committee. 1997. Flora of North America north of Mexico. 
Vol 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford University Press, New 
York & Oxford. 

Goorrey, R.K. 1988. Trees, shrubs and woody vines of northern Florida and adjacent Geor- 
gia and Alabama. The University of Georgia Press, Athens and London. 

Goorrey,R.K.and J.W.Wooten. 1979. Aquatic and wetland plants of the southeastern United 
States. Monocotyledons. University of Georgia Press, Athens. 

Goorrey,R.K.and J.W.Wooren. 1981.Aquatic and wetland plants of the southeastern United 
States. Dicotyledons. University of Georgia Press, Athens. 

Goutp, FW. 1975. The grasses of Texas. Texas A & M University Press, College Station. 

Haroner, A.H. 1960 The geology and ground-water resources of Calcasieu Parish, Louisi- 
ana. United States Government Printing Office, Washington. 

HoLtano, W.C., L.W. Houco and G.C. Murry. 1952. Geology of Beauregard and Allen Parishes. 
Louisiana Geol. Surv. Bull. 27. 

Jones, PH., A.N. Turcan, and H.E. Skisitzke. 1954. Geology and ground-water resources of 
southwestern Louisiana. Department of Conservation, Louisiana Geological Survey, 
Baton Rouge. 

Kartesz, J.T. 1999.A synonomized checklist of the vascular flora of the United States, Canada, 
and Greenland. 1st. ed. In: Kartesz, J.T.and C.A. Meacham, eds. Synthesis of the North 
American flora, Version 1.0.North Carolina Botanical Garden, Chapel Hill, NC. 

Mac Roserts, D.T. 1988.A documented checklist and atlas of the vascular flora of Louisiana. 
Published by the author, Shreveport, LA. 

Raveoro, A.E., H.E. ALES, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill. 

Roy, AJ.and C.T. Mipkirr. 1988. Soil survey of Calcasieu Parish, Louisiana. USDA, Soil Conser- 
vation Service. 

Smatt,J.K. 1933. Manual of the southeastern flora. University of North Carolina Press, Chapel 
Hill. 


386 SIDA 19(2) 

Smith, LM. 1996. The rare and sensitive natural wetland plant communities of interior 
Louisiana. Louisiana Department of Wildlife and Fisheries, Natural Heritage Program, 
Baton Rouge. 

THomas, R.D.and C.M. Atten. 1993. Atlas of the vascular flora of Louisiana. Vol.|: Ferns & fern 
allies, conifers, & monocotyledons. Louisiana Department of Wildlife & Fisheries, Natu- 
ral Heritage Program, Baton Rouge, LA. 

THomas, R.D. and C.M. Aten. 1996. Atlas of the vascular flora of Louisiana Volume II: Dicoty- 
ledons Acanthaceae - Euphorbiaceae. Louisiana Department of Wildlife & Fisheries, 
Natural Heritage Program, Baton Rouge, LA. 

THomas, R.D.and C.M. ALLEN. 1998. Atlas of the vascular flora of Louisiana Volume Ill: Dicoty- 
ledons Fabaceae - Zygophyllaceae. Louisiana Department of Wildlife & Fisheries, Natural 
Heritage Program, Baton Rouge, LA. 


ESTABLISHING ETHNOBOTANICAL CONSERVATION 
PRIORITIES: A CASE STUDY OF THE 
KALLAWAYA PHARMACOPOEIA 
Kevin D.Janni 


Botanical Research Institute of Texas 


509 Pecan Street 
Fort Worth, TX 76102-4060, U.S.A. 
kjanni@brit.org 
Joseph W. Bastien 


Department of Anthropology and Sociology 
niversity of Texas, Arlington 
Arlington, 1X 76019, U.S.A. 


ABSTRACT 


The Kallawaya herbalists of Bolivia follow a healing tradition over a millennium old. Twenty-eight 
plants of the indigenous pharmacopoeia have been documented in both 16" and 17" century 
historical literature and contemporary ethnobotanical research. This continuity has bio-cultural 
implications representing not only useful medicinal plants but also traditional cultural knowledge. 
These plants and their antiquity of use represent the cultural heritage of the Kallawaya and should 
be considered along with the biodiversity priorities of local conservation programs. The rapid loss 
of biological diversity and local knowledge is putting modern ethnobotany at the risk of becom- 
ing decadent, before becoming a discipline. Analysis of indigenous pharmacopoeias can be an 
effective means of determining bio-cultural importance and establishing ee conser- 
vation PHlOnies: TS ue case study discusses a selected grOup of plants that have 

use utili f these plants over time 
represents their bio- cultural importance and priority for ere eicn and sustainability efforts. 


KEY WORDS: Bolivia, Conservation, Ethnobotany, Kallawaya, Pharmacopoeia 


RESUMEN 


Lost i Kall de Bolivia si d ion de mas de un milenio. Se han 
documentado veintiocho pants ae a farmacopea iglosnes en ve Ure fetvle histdrica de los siglos 
XVI y XVIl y de la investigacié idad tien e limplicaciones 
bio-culturales que eerie no solo plantas medicinales litiles sino también ul- 
tural tradicional. Estas plantas y su antiguedad de uso representan la hevencis eultuial da los 
Kallawaya y debe ser considerada en los programas de conservacién junto con las prioridades de 
biodiversidad. La pérdida rapida de diversidad biolégica y conocimiento local esta poniendo a la 
etonotanica ede lia en el ESeO de pee ooo antes de llegar a ser una disciplina. El 

| | wingers Se of ecialenia importancia 
io-cultural y establec idadesd caso de estudio 
discute | un gripe selecto ‘de plantas que eel romania ethnomédica ume con usos 


-o 


an ews s.La ial t 


bic: cull y ‘ls mies para la conservacién y usO eer ido. 


SIDA 19(2); 387 — 398. 2000 


388 SIDA 19(2) 


INTRODUCTION 


The Kallawaya herbalists of Bolivia are renowned throughout Argentina, Bolivia, Chile 
and Peru (Bastien 1987) (Fig. 1). Living at altitudes of 2700-4300m and frequently travel- 
ing to communities in varied ecological zones, the Kallawaya have not only established 
a continuity in Andean folk medicine, they have also had the opportunity to greatly 
augment their pharmacopoeia along the way (Bastien 1983; Abdel-Malek et al. 1996; 
Janni & Bastien n.d). They follow a medical tradition from the Tihuanaco (400-1145), 
Mollo (1145-1453), Inca (1438-1532), Spanish (1532-1825), and Bolivian Republic (1825- 
present) periods (Oblitas-Poblete 1969; Bastien 1982, 1983, 1987; Abdel-Malek et al. 1996). 
The Kallawaya utilize nearly 900 plants (Girault 1987) of the 2000 medicinal plants re- 
ported in all of Bolivia (De Lucca & Zalles-Asin 1992).In contemporary times a syndicate 
of herbalists known as the Society for Bolivian Traditional Medicine (SOBOMETRA) has 
been responsible for preserving and disseminating Kallawaya herbal knowledge, while 
the Servicio Integrado en Salud (SIENS) clinics in La Paz utilize both physicians and 
Kallawaya herbalists to provide integrated ethnomedical and biomedical treatment 
(Abdel-Malek et al. 1996), 

Compiled from a survey of historical and ethnobotanical literature are 28 plants 
that have been present in the health and healing practices of the Kallawaya since pre- 
Columbian times. This indigenous cultural knowledge extends deep into Kallawaya 
ethnohistory and offers compelling reasons for the conservation of bio-cultural diver- 
sity. By targeting plants in indigenous pharmacopoeias for conservation priority we not 
only help sustain traditional cultural knowledge and biological diversity, but also the 
ethnomedical practices of the community. Focusing ethnobotanical research on com- 
munity level priorities helps target plants of cultural importance that frequently go un- 
noticed by global conservation programs that are,"...the action of outsiders who are 
culturally and politically detached from the threatened environments and who identify 
species for conservation through western economic models” (Etkin 1998). Discussion of 
the conservation priorities of ethnobotanical research has been unjustifiably rare (Alcorn 
1995; Benz et al. 1996; Etkin 1998; Eisner & Beiring 1994; Laird et al. 1997; Posey & Balée 
1989) and deserves further attention and investigation. 


METHODS 


Data on pre-Columbian (before European invasion) uses of medicinal plants were com- 
piled by a survey of the historical literature (Anonimo 1703; Calancha 1638; Cobo 1891, 
1892, 1893; Contreras & Valverde 1650; Jimeneza 1965; Monardes 1569; Oviedo 1535; 
Polo de Ondegardo 1585;Vega 1609; Yacovleff & Herrera 1935).Information on medicinal 
plants was often fragmentary and incomplete, but fourteen plants had two or more 
references. Information from modern ethnobotanical literature was considerably more 
substantial and was compiled to see what plants had persisted since pre-Columbian- 
era. Data on the Kallawaya pharmacopoeia were compiled by Bastien (1982, 1983, 1987), 
including information on therapeutic uses and non-medicinal or economic uses. The 


JANNI AND BASTIEN, CTUNOROATANICAIL 389 
1. 
< 
a 4 a R 
Le a | : e 
a : : 
z lL. aa ) / 
ie 
®@ Cuzco i Boo L re 
; / 
e | 
CharazaniB einer b ‘ ) 
Lake ‘ 
Titicaca Ley, a eee, 
. e@elLaPaz “% ~ 7 
Arequipa 6 lan, e@cochabamba te 
“, Chr; *Santa Cruz 
D) ‘es ruro @ "lille On, ) 


c:,@ Sucre Mal 
Potosi e ey, pare 
ape 
Pieces. as | 
oa wo NS : 
SO @ Tucuman ) 
ad 


A\ Kallawaya Ayllus 
@ Cities & Towns 


ARGENTINA 


? e Mendoza 


ya Indians. (From Bastien 1987 with permission of University of Utah Press). 


Kallawaya reportedly have an unwritten pharmacopoeia of over 900 medicinal plants. 
For the purpose of this study, the pharmacopoeia is limited to published information 
and follows the nomenclature therein (Bastien 1982, 1983, 1987). 

Relative importance values were calculated for each plant using a normalized num- 
ber of pharmacological properties (PH) and a normalized number of body systems (BS) 
treated. This approach follows the one used by Bennett and Prance (2000) for measuring 
the relative importance of plants in indigenous pharmacopoeias and is primarily a mea- 


390 SIDA 19(2) 


sure of the versatility of each plant in relation to the pharmacopoeia as a whole. For 
example, Minthostachys andina is used to treat 10 body systems, the most of any plant in 
the pre-Columbian pharmacopoeia. Therefore, it has a normalized BS value of 1 (10/10). 
Erythroxylum coca is used to treat six body systems, four less than Minthostachys andina. 
The BS value for E. coca is 0.6 (6/10). Minthostachys andina has seven pharmacological 
properties, again the most of any pre-Columbian plant in the Kallawaya pharmacopoeia. 
Thus its PH value is 1(7/7). The combined PH and BS values of M. andina equal 2.0, which 
is then divided by two and multiplied by 100 to calculate the relative importance of pre- 
Columbian M.andina, 100.This approach is useful for calculating the relative importance 
of a plant by taking into account the differences in number of pharmacological proper- 
ties and body systems treated. For example, F. coca has five pharmacological properties 
and treats six body systems giving it a pre-Columbian relative importance of 65. Nicoti- 
ana rustica also has five pharmacological properties, but treats only five body systems, 
giving it a relative importance of 60.The relative importance scores for each plant reflect 
differences in versatility. 

The relative importance of each pre-Columbian plant is then analyzed compara- 
tively with its relative importance in contemporary times. By taking an average of pre- 
Columbian relative importance and contemporary relative importance we find the overall 
relative importance of each plant. For example, £. coca has a pre-Columbian relative im- 
portance of 65 and a contemporary relative importance of 70, thus having an overall 
relative importance of 68. By comparative analysis, those plants used in pre-Columbian 
times are recognized for their continuity. Seven plants, Psoralea pubescens, Mutisa 
acuminata, Salvia haenkii, Verbena hispida, Peperomia anaequifolia, Gnaphalium 
quadichaudium, Ambrosia peruviana, are included in this table because they were cited 
in the literature without specific pharmacological or therapeutic details other than be- 
ing medicinal. There is no known pre-Columbian importance for these plants, but their 
contemporary relative importance is included. This comparative analysis is designed to 
show the changes in number of pharmacological properties and body systems treated 
between pre-Columbian and contemporary times. Medical terminology follows that of 
Dorland’s Medical Dictionary (1980) and discussed in Bastien (1982, 1983, 1987). 


RESULTS 


There are 28 plants in the Kallawaya pharmacopoeia cited in historical literature (Table 
1). This is a small portion of the Kallawaya pharmacopoeia and information on these 
plants and others in the modern literature is considerably more comprehensive. These 
plants have a wide variety of therapeutic uses (19 in all) mainly as analgesics, diuretics, 
antiseptics, and expectorants (Table 2). Aside from medicinal uses there are 12 plants 
that overlap economically as aromatics, ornamentals, dyes, foods, intoxicants, etc (Table 
3).Minthostachys andina recorded an overall relative importance of 80, the highest of all 
28 pre-Columbian plants (Table 4). Three plants, Erythroxylum coca, Urtica flabellate and 
Nicotiana rustica, scored in the sixties and only one plant, Polypodium angustifolium, scored 


JANNI AND BASTIEN, ETHNOBOTANICAL 


Taste 1.Pre-Columbian medicinal plants of the Kallawaya pharmacopoeia. 


Genus, species (Family) Vernacular Name References 
Ambrosia peruviana Willd. (Asteraceae) Malco 5,9, 10 
Azorella biloba Schlecht. (Apiaceae) Yareta 3 
Baccharis pentandii DC. (Asteraceae hilca 3,5 
Calceolaria cuneiformis R&P. (Schrophulariaceae) Ava Zapatilla 3 
Calceolaria aff. eae (Schrophulariaceae) Puru Puru 3 
Chenopodi br L.(Chenopodiaceae) — Paico Lombrio 5,6, 8, 10 
hona calvisa Wedd. (Rubiaceae) Quina Cascarilla 2,3,10 
Datura pangniaed L. (Solanaceae Floripondio 2,3 
Edt g HBK. (Equisetaceae) Cola de Caballo 3 
ie xylum coca Lam. (Erythroxylaceae) Coca 3,6 
Gentiana lutea L. (Gentiana Pencacuc 3 
aD apna ue quadichavdum BC. (Asteraceae) Wira Wira 4 
hys andina nth. (Lamiaceae) una 3,10 
Mutisa acuminata R&P. een Chinchircuma 10 
ae ylon balsamum L., (Fabaceae) Quina Quina 3,10 
N ti fficinale R.Br. (Brassicaceae) Berro 9 
Nicotiana rustica L. (Solanaceae) Sayre 3,6,7,8 
Peperomia inaequalifolia R&P. (Piperaceae) Congona 6 
Plantago tomentosa Lam. (Plantaginaceae) Llanten 35,10 
beste angustifolium SW. (Polypodiaceae) Calaquala 3,10 
ileatum SW. aaa — Macho 3,10 
see ee Pers. (Fa bacea Bilye 1 
Psittacanthus cuneifolius R&P. (Loranthaceae) Suelde con Suelda 3 
Salvia haenkii Benth. (Lamiaceae) Savia Grande 3,10 
Senecio tephosiodes Turcz. (Asteraceae) Mamanlipa 3 
Solanum radicans Lf. (Solanaceae) K’umasillo 3,9 
Urtica flabellate H.B.K. (Urticaceae) rtega 3,10 
Verbena hispida R&P. (Verbenaceae) Verbena 3,9 


].Anonimo 1703 

2. Stee ne 

3.Cobo 

4. eae me ens 1650 


5. ees la Espada 1965 
6.Vega 

7. Polo ns Oncegsdo 1585 
8. oe 

9. Oviedo 


10. ae ar Herrera 1935 


in the fifties (Table 4). Twenty-three of the 28 pre-Columbian plants have a relative im- 
portance under 50.Seven plants, Polypodium angustifolium, Plantago tomentosa, Cinchona 
calvisa, Gentiana lutea, Polystichum aculeatum, Psittachanthus cuneifolius,and Solanum 
radicans, show an increase in pharmacological properties and body systems treated 


392 SIDA 19(2) 


TABLE 2. Weer ie Mee of Pre-Columbian 
A=analgesic.A1 = antiseptic ce = astringent.C = cardiotonic. a bi eee D1 = Diuretic. 
D2 = tae D3 - ea Icent.E = emetic. E1 = expectorant. F = febrifuge. R = cae ny R1 

= resolvent. S = stomachic. $1 = eae $2 = Stimulant. T = pera ee = vermifuge. V1 
vulnerary. 


Species A Al A2 C D D1 D2 D3 E E1 FR R1S $1 S2 TV V1 


Ambrosia peruviana Xx x 
Azorella biloba x 

Baccharis pentandii x x 

Calceolaria cuneiformis x 

Calceolaria engleriana x 


Ampbrocinide x x 


Che vavalin 
saad 
Cinchona calvisa Xx Xx 
Datura sanguinea Xx x 
Earsicetiim hoagntence 
a | 


Y 


Ervthr, vit q 
ythroxy x x x x x 


Gentiana lutea x 
Gnaphalium ae x 

Minthostachys andina x x x x Xx x 
Mutisa Peete x x 

A Ny i} bal 


qq wrth ; | 
INI tort fh x x 


Nicotiana rustica x x x 
Sa Le lifoliu xX x 


Pols war Im angi ictifolirim x x x x x 


Poly ictichiym acrleatiim xX x x x 


Psoralea eee cens x Xx xX x 
Psittacanthus cuneifolius x 
saivia den nil x x 


a 
tephosiodes x x x 


Solanum radicans x x x 
Urtica flabellate x x 
Verbena hispida x x 


Totals 96 5 15.8 3 1 27 6117 $2 33.21 


throughout time. The other 21 plants show a decrease in PH and BS values between pre- 
Columbian usage and present. These differences could be attributed to the scant and 
fragmentary historical documentation of pre-Columbian medicinal plants. 


OBSERVATIONS AND DISCUSSION 


Analysis of the pre-Columbian pharmacopoeia elucidates the importance of medicinal 
plants to the health and healing practices of the Kallawaya. Outside the pharmacopoeia 
these plants are culturally useful as food, for hygienic purposes, ornamental, and other 
purposes. These twenty-eight plants are arguably important parts of the traditional cul- 


JANNI AND BASTIEN, ETHNOBOTANICAL 393 


Tasle 3. Other uses of Pre-Columbian medicinal plants in the Kallawaya pharmacopoeia. A = aro- 
matic. B = used to make broom. D = dye. E = embalming. F = food.H = horse injuries.H1 = Hygiene. 
I = intoxicant. O= ornamental. R = resin to catch birds. 


Species A B D E F H HI I fe) R 


Ambrosia peruviana Xx 

Baccharis pentandii x 

Calceolaria ee x 
Pe sangup x 


a gual x x 


Gi +7 inn! x x 

Nicotiana rustica x 

Peperomia inaequalifolia x x 

peCamn enon x 
olanum n x 


tural, environmental, and biological knowledge of the Kallawaya. Along with plants hav- 
ing a pre-Columbian continuity, the Kallawaya have integrated exotic plants into their 
pharmacopoeia (Janni & Bastien n.d.). Despite the integration of exotic species into 
the pharmacopoeia, the 28 plants discussed herein have retained much of their cultural 
and medicinal importance for over a thousand years. 

The diversity of therapeutic uses of these plants is remarkable. Clearly, by the time 
of Spanish invasion, the Kallawaya had actively investigated the phytomedicinal poten- 
tial of the local and regional flora. The diversity of therapeutic uses indicates the 
ethnomedical sophistication of the Kallawaya, and offers a picture of the health and 
healing concerns of pre-Columbian Kallawaya culture. This information is useful in un- 
derstanding the epidemiological fluctuations of the Kallawaya throughout time by re- 
vealing the patterns of health and sickness that enable us to ask questions as to why 
they changed. Also, by compiling such data we find the plants that are not only impor- 
tant medicinally, but also have been an integral part of Kallawaya cultural heritage. 

Many ethnobotanical investigations compile information in an effort to identify 
potential new drugs; we have compiled this information to identify plants of cultural 
importance as well as plants of priority for biological conservation and sustainability 
programs. Local efforts in conservation offer greater potential results than those of west- 
ern economists because they represent the intimate local knowledge of the native ecol- 
ogy and long experience with the species in question (Etkin 1998). Assessing the 
significance of specific taxa with cultural and ecological importance gives us a frame- 
work by which conservation of local biota is based on local values (Benz et al. 1996). 

The list of 28 plants discussed herein describes a portion of the pharmacopoeia 
that has been analyzed based on local values. The long-standing persistence of these 
plants in the Kallawaya pharmacopoeia indicate continued cultural reliance on these 
taxa in ethnomedical and economic practices. The versatility of these species within the 


Table 4. Relative importance values of selected medicinal plants in the Kall n 16-17' PH = number of pharmacological wean es ae 
ner ee 17" cana: Benine 16-17" Rel PH = relative number of pharmacological properties sited in 16 &17" ae. iterature. (norm 

maximum value of 1). 16-17" BS = number of body systems treated, sited in 16 & 17% Century ese 16-17° Rel BS = bes number 0 
systems treated, sited in 16 & 17 Century Literature. (normalized to the maximum ve of 1). 16-17" RI = relative importance of 16 & 17 Century 
Medicinal Plants. 20** PH = number of pete ob ie: properties, 20 Century. 20 Rel PH = relative number eel properties. (normalized 
to the maximum value of 1). 20" BS = number of body systems treated, 20" Century. 20° Rel BS = relative number of pharmacological properties, | 
Century. 20" RI = relative importance of medicinal plants, 20" Century. ORI = overall relative importance. RI = ((Rel PH + Rel BS)/2) x 100. ORI = (16-17% 
+ 20° RI)/2. 


16-17% = 16-17 = 16-17t = 16-17% = 16-17th =. 20th = 2th 20% = =—.20¢ 20° 
Species PH Rel PH BS Rel BS RI PH Rel PH BS Rel BS RI ORI 
Minthostachys andina 7 1.0 10 1.0 100 3 0.6 4 0.6 60 80 
Erythoxyl 5 0.7 6 0.6 65 3 0.6 > 0.8 70 68 
Urtica flabellata 5 0.7 7 0.7 70 3 0.6 3 0.5 55 63 
Nicotiana rustica > 0.7 5 0.5 60 2 0.4 5 0.8 60 60 
Polypodium angustifoli 2 0.2 ] 0.1 15 5 1.0 6 1.0 100 58 
Baccharis pentandii 2 0.2 7 0.7 45 2 04 4 0.6 50 48 
Equisetum bogotense 5 0.7 5 0.5 60 2 04 2 0.3 35 48 
Plantago tomentosa 2 0.2 3 0.3 25 4 0.8 4 0.6 70 48 
Senecio tephosiodes 5 0.7 4 0.4 55 ] 0.2 4 0.6 40 48 
Myroxylon balsamum 7 1.0 > 0.5 75 1 0.2 1 0.2 20 48 
Cinchona calvisa 2 0.2 2 0.2 20 3 0.6 5 0.8 70 45 
Chenopodi b 2 0.2 4 0.4 30 2 0.4 4 0.6 50 40 
Azorella biloba 3 0.4 2 0.2 30 2 0.4 3 0.5 45 38 
Psoralea pubescens 0 0.0 0 0.0 00 4 0.8 4 0.6 70 35 
Mutisa acuminata 0 0.0 0 0.0 00 2 0.4 6 1.0 70 35 


v6E 


(Z)6L Vals 


Table 4. continued 


16-17 = 16-17 = 16-17 = 16-17% = 16-17% }=— 20% = 20th 20% =. 20th 20th 
Species PH Rel PH Rel BS H Rel PH BS Rel BS Ri ORI 
Datura sanguinea 3 04 3 0.3 35 1 0.2 3 0.5 35 35 
Gentiana lutea 1 0.1 ] 0.1 10 3 0.6 4 0.6 60 35 
Polystichum aculeatum 2 0.2 ] 0.1 15 3 0.6 3 0.5 55 35 
Psittacanthus cuneifolius 2 0.2 2 0.2 20 2 0.4 4 0.6 50 35 
Solanum radican ] 0.1 1 0.1 10 2 0.4 3 0.5 45 28 
Salvia haenkii 0 0.0 0 0.0 00 3 0.6 3 0.5 55 28 
Verbena hispida 0 0.0 0) 0.0 00 4 0.8 2 0.3 55 28 
Calceolaria cuneiformis 3 0.4 1 0.1 25 ] 0.2 2 0.3 25 25 
Peperomia inaequifolia 0 0.0 0 0.0 00 2 0.4 3 0.5 45 23 
Gnaphalium quadichaudium 0 0.0 0 0.0 00 2 0.4 3 0.5 45 23 
Calceolaria engleriana 2 0.2 2 0.2 20 ] 0.2 i 0.2 20 20 
Nastri fficinal ] 0.1 ] 0.1 10 ] 0.2 2 0.3 25 18 
Ambrosia peruviana 0 0.0 0 0.0 00 2 0.4 2 0.3 35 18 


WOINVLOGONALS NalLSVa UNV INNVI 


S6E 


396 SIDA 19(2) 


pharmacopoeia and outside it also makes conservation priorities more compelling. Sev- 
eral taxa are used for more than one therapeutic (Table 2) or pharmacological (Table 4) 
property and several are used for a variety of economic uses (Table 3). Conservation 
initiatives focused on biological diversity alone neglect the socio-cultural importance of 
some taxa. By using ethnobotanical research to target taxa of such importance we have 
a chance to protect biological diversity and in the process also protect and sustain tradi- 
tional cultural knowledge, indigenous health care systems, and plants of particular cul- 
tural significance. Conservation and sustainability initiatives that focus on community 
level priorities allow us to work on several problems at once. Direction based from an 
ethnobotanical perspective (e.g. analyzing indigenous pharmacopoeias) gives us the 
opportunity to address problems of social, cultural and biological importance. The data 
discussed here provides only a part of a broader investigation that should include field- 
work and interviews to more accurately determine current social and biological needs. 

The loss of local knowledge and biological diversity should be the primary concern 
of every ethnobotanist. Organizing at local levels with an ethnobotanical framework 
simultaneously protects biological and cultural diversity. Losing tribal elders and the 
knowledge they encapsulate is losing information on the biological environment of the 
surrounding area. Conservation and sustainability programs that are sensitive to bio- 
cultural issues like these will help protect traditional cultural knowledge and biodiversity 
for future generations. 


CONCLUSION 


Analysis of the pre-Columbian continuity of the Kallawaya pharmacopoeia reveals plants 
of cultural and medicinal importance that should be targeted for conservation and 
sustainability programs. The persistence of use of these plants throughout centuries of 
healing, as well as their role outside the pharmacopoeia for a variety of economic uses 
elucidates the importance of these plants to the cultural heritage of the Kallawaya. By 
focusing conservation efforts on plants of cultural importance we not only recognize 
indigenous environmental knowledge, we also have th tunity to protect biologi- 
cal diversity. With overwhelming predictions of 60,000 sie niger sient species to be- 
come extinct by the middle of the next century, ethnobotany must be a leader in iden- 
tifying conservation and sustainability priorities by analysis of local needs. Just as local 
knowledge has been tapped for new drug leads, we must go to it in the future to deter- 
mine conservation priorities. 


ACKNOWLEDGMENTS 
Thank you to the following individuals who read and commented on an earlier version 
of this manuscript: Ted Barkley, Carol Janni, and Debra Trock. Also, thanks to the staff at 
BRIT and UTA libraries for assistance. 


JANNI AND BASTIEN, ETHNOBOTANICAL 397 


REFERENCES 


Appet-Ma ek, S., J.W.Bastien, W.F. MAHLER, Q. JIA, M.G. REINecKE, W.E. Rosinson, Y. SHU, and J. ZALLES- 
Asin. 1996. Drug leads from the Kallawaya herbalists of Bolivia. 1.Background, rationale, 
protocol and Anti-HIV activity. J. Ethno-pharmacol. 50:157-166. 

Acorn, J.B. 1995. Economic botany, conservation, and development: What's the connec- 
tion. Ann. Missouri Bot. Gard. 82:34-46. 

Anonimo, Jesuita. 1703. Technologia indigena. Revista Inca, 1923. Volume 1: Nos. 2 and 3. 
Lima. 

Bastien, J. W. 1982. Herbal curing by Qollahuaya Andeans. J. Ethno-pharmacol. 6:13-28. 

1983. Pharmacopoeia of Qollahuaya Andeans. J. Ethno-pharmacol. 8:97-111. 
1987. Healers of the Andes: Kallawaya herbalists and their medicinal plants. 
University of Utah Press. Salt Lake City. 

Bennett, B.C. and GT. Prance. 2000. Introduced plants in the indigenous pharmacopoeia 
of Northern South America. Econ. Bot. 54:90-102. 

Benz, B.F,, Cevattos, E., E. Munoz M., and F. Santana M. 1996. Ethnobotany serving society: A 
case study from the Sierra de Manantlan Biosphere Reserve. Sida 17:1-16. 

CALANCHA, Fr. ANTONIO DE LA. 1638.Cronica Moralizadora del orden de San Angustin. Barcelona 


and Lima. 

Coso, Fr. Bernabe. 1891-1893. Historia del Nuevo Mundo. In Yacovleff and Herrera. 1935. El 
Mundo Vegetal de Los Antiguous Peruanos. Revista del Museo Nacional, Lima. 

Contreras, VASCO DE and Vatverbde. 1650. Relacion de la ciudad del Cuzco. In Yacovleff and 
Herrera. 1935.El Mundo Vegetal de Los Antiguous Peruanos.Revista del Museo Nacional, 
Lima. 

De Lucca, M.and J. ZALLES-ASIN. 1992. Flora medicinal Boliviana.Los Amigos del Libro, La Paz, 
Cochabamba. 

Dortanp’s Mepicat Dictionary. Shorter Edition. 1980.The Saunders Press, Philadelpia, PA. 

Eisner, T.and E. BeIrRING. 1994, Biotic exploration fund: Protecting biodiversity through chemi- 
cal prospecting. BioScience 44:95-98. 

Etkin, N.L. 1998. Indigenous patterns of conserving biodiversity: Pharmacological impli- 
cations. J. Ethno-pharmacol. 63:233-245. 

Girautt, L. 1997. Kallawaya, curanderos itinerantes de los Andes. UNICEF, ORSTOM, La Paz. 

Janni, K.D. and J.W. Bastien. n.d. Exotic botanicals in the Kallawaya pharmacopoeia. Econ. 
Bot. Submitted. {available from author} 

JIMENEZ DE LA Espapa, Marcos. 1965. Relaciones geograficas de Indias- Peru. Ediciones Atlas, 
Madrid. 

Lairb, J.W., MJ. Batick, and S.A. Lairp. 1997. Medicinal plants: Can utilization and conserva- 
tion coexist? New York Botanical Garden, Bronx. 

Monaroes, N. 1569. Historia medicinal de las cosas que traen de nuestras Indias occidentals, 
que sirven de medicina.|In Yacovleff and Herrera 1935.E] mundo vegetal de Antiguos 
Peruanos. Revista del Museo Nacional, Lima. 


398 SIDA 19(2) 


Ostitas-Postete, E.1969.Plantas medicinales de Bolivia. Farmacopea Callawaya.Los Amigos 
del Libra, Cochabamba. 

OvieDo, GONZALO FERNANDEZ DE. 1535. Carta dirigida al emperador Carlos V. in Collecion de 
documentos ineditos relativos al descumbrimiento, conquesta y colonizacion de las 
possesiones espanoles en America y Oceania. 1865. Madrid, Spain. 

POLO DE ONDEGARDO, JUAN. 1585. Lor errors y supersticiones de los Indios. In Yacovleff and 
Herrera 1935. El Mundo Vegetal de Antiguos Peruanos. Revista del Museo Nacional, 
Lima. 

Posey, D.A.and W. Bate (eds.) 1989. Resource management in Amazonia: Indigenous and 
folk strategies. New York Botanical Garden, Bronx. 

VEGA, GARCILASO DE LA. 1609. Comentarios reales de los incas. In Libreria Internacional del 
Peru. 1959, 

Yacovterr, E.and F. Herrera. 1934-35. El mundo vegetal de los Antiguos Peruanos. Revista 
del Museo Nacional, Lima. 


CRYPTOCORYNE BECKETTII (ARACEAE), 
A NEW AQUATIC PLANT IN TEXAS 


David J.Rosen 
Bes 5. ye aa oe ineers 
1229 
Ga Weston, TX i 


The genus Cryptocoryne Fisch. ex Wydl. comprises approximately 50 species distributed 
on islands and coastal areas of South East Asia (Muhlberg 1982). Cryptocoryne beckettii 
Thw. ex R.Trim., a native of Sri Lanka (MUhlberg 1982), was collected in 1996 in the San 
Marcos River in the City of San Marcos. This taxon has not been previously reported in 
Texas according to Jones et al.(1997), Hatch et al.(1990), and Correll and Johnston (1970). 
Large, naturalized colonies of C. beckettii were observed growing in open shallow riffles 
as well as in shaded deep pools. Cryptocoryne beckettii is a valued aquarium plant col- 
lected in the wild and widely exported (Nicolson 1987).The occurrence of this species in 
the San Marcos River is likely due to escape from cultivation or dumping of aquariums as 
has been proposed for the introduced aquatic fern Ceratopteris thalichtroides by Hannan 
(1969). A description of C. beckettii modified from Nicolson (1987) and illustration (Fig. 1) 
follow. 

Cryptocoryne beckettii [hw. ex R. Trim. J. Bot. 23:269. 1885. 


Perennial, rhi tous emergent-submerged herbs. Leaves basal with elongate, sheath- 
ing petioles to 15 cm; blades glabrous, ovate to narrowly ovate, 3—-9(-13) cm long, (1-)1.5- 
3.5(-4) cm wide, upper surface green to dark green to brown and marbled to red-brown, 
lower surface red-tinged to more or less brownish or green; veins usually conspicuously 
red; apex acute to acuminate; base obtuse to cordate, margin entire, sometimes undu- 
late; submerged specimens mostly with larger, thinner leaves, often brownish marbled. 
Inflorescence (not seen) short peduncled;spathe 4—12(-20) cm long, limb greenish brown, 
narrowly ovate, 0.5-1.2 cm wide, 1.5-3 cm long, twisted, upright to somewhat recurved 
and twisted; spadix 1.0 cm long. 

There are three closely related species, C. walkeri Schott, C. wendtii de Wit, and C. 
undulata Wendt. A key to od the four taxa can be found in Nicolson (1987). 


Ri f + Tt 


Voucher specimen: TEXAS. Hays Co.: treat 
ment plant,08 Aug 1996, Rosen 202 aes SWT). 


ACKNOWLEDGMENTS 
| am indebted to Don Bryne (Suwannee Labs, Florida) for his determination of the 
Cryptocoryne material to species and Collete Jacono (USGS) for her enthusiasm and as- 
sistance. Thanks are expressed to Stanley D. Jones and two reviewers for their helpful 
comments. 


SIDA 19(2): 399 - 401. 2000 


SIDA 19(2) 


400 


Wl 


t 


Fic. 1.C 


ROSEN, CRYPTOCORYNE BECKETTII IN TEXAS 401 


REFERENCES 


Corrett, D.S. and M.C. Jonnston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation. Renner 

HANNAN, H.H. 1969.The introduction and establishment of Ceratopteris in Texas. Amer.Fern 
J.59:122. 

Hatch, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas 
Agric. Exp. Sta. Bill. MP-1655. 

Jones, S.D., J.K. Wipre, and RM. Montcomery. 1997. Vascular plants of Texas: A comprehensive 
checklist including synonymy, bibliography, and index. University of Texas Press, 
Austin. 

Munveerc, H.1982.The complete guide to water plants.EP Publishing Limited, GDR.391 p., 
with 109 color and 112 B&W photographs and 59 illustrations. (Translated from the 
German by Ilse Lindsay.) 

Nicotson, D.H. 1987. Araceae. In:M.D.Sassanayke and F.R. Fosberg, eds. Flora of Ceylon (Vol. 
VI). Smithsonian Institute and the National Science Foundation, Washington, D.C. 
Amerind Publishing Co. Pvt. Ltd., New Delhi. Pp. 117-101 


402 SIDA 19(2) 


BOOK REVIEW 


D.M. Persatt. 2000. Paleoethnobotany. A Handbook of Procedures, second edition. 
cademic Press, 525 B Street, Suite 1900, San Diego, CA 92101-4495, U.S.A. 


Eleven years after its debut, Paleoethnobotany: A Handbook of Procedures returns in second edition. 
This classic by University of Missouri-Columbia anthropologist Deborah Pearsall is back -700 pages 
in all- in updated form. In the time between editions major advances were made in the field of 
phytolith analysis, as well as a general growth in paleoethnobotanical literature. Pearsall makes 
appropriate updates and responds to the critics of the first edition by making this edition more 

Obal in scope, correcting an imbalance in the first book in favor of New World examples. Also new 
is the final chapter on integrating biological data into paleoethnobotanical analysis. This chapter 
makes a case that reconstructing past human diets may be the biggest and most immediate con- 
tribution by the field of paleoethnobotany. 

k is laid out in six chapters followed by references and an index. Chapter one,"The 
Paleoethnobotanical Approach” includes an introduction to the field, a historical overview and a 
discussion of the nature and status of ethnobotany. Chapters two and three are devoted to 
macroremains and the techniques for recovering, identifying and interpreting them. Chapter four, 
“Pollen Analysis” covers the nature and production of pollen, field sampling, laboratory analysis, 
and a discussion of issues and directions in archaeological pollen analysis. Chapter five, “Phytolit 
Analysis," covers the nature and occurrence of phytoliths and methods of field sampling, lab test- 
ing, and interpreting the results. Chapter 6, "Integrating Biological Data,” is the most welcome 
addition to this volume. Broken into two parts,"Indicators of Diet and Health" and “The Interplay of 
Dietary Indicators," respectively, Pearsall gives a welcomed big-picture framework to the field. Part 
one of chapter six discusses indirect dietary factors, such as botanical and faunal data, and then 
discusses direct indicators like gut contents and coprolites, stable isotopes, trace elements, and 
skeletal indicators of nutrition and health. Part two of chapter six discusses predictions from di- 
etary indicators, combined indicators for eight neotropical diets, and two archeological case stud- 
ies from Ecuador and Peru. This chapter is a valuable addition to the annals of paleoethnobotanical 
literature by developing an approach for integrating and evaluating multiple lines of evidence 
concerning diets. Pearsall discusses eight diets chosen to represent diets spanning the transition 
from hunting and eee MS Ach oaatsy in the New World tropics, which is then illustrated by 
two Neotropical onstructing diets is an He ela se and bio-anthro- 
pological issue by which investigati | understand how vived and 
prospered. This chapter is followed bya list of rercieness and closed - a general Hae Black and 
white se and tables appear frequently throughout to illustrate concepts discussed therein. 

Paleoethnobotany: A Handbook of Procedures is valuable for its comprehensive outline of ap- 
proaches and techniques of ee ce more mpertanty. i exe as ais only peek of its a 
Pearsall has made an effort to make t 
succeeds admirably. My only complaint is the $150 price that is simply an outrageous amount to 
ask from a student or professional.| would be more tempted to require a smaller, less comprehen- 
sive, text for a class in lieu of the heavy price tag.—Kevin D. Janni. 


SIDA 19(2): 402. 2000 


THYMELAEA PASSERINA (THYMELAEACEAE) 
NEW TO TEXAS 


Walter C. Holmes 


Department of Biology 
Baylor University 
Waco, TX 76798-7388, U.S.A. 
walter_holmes @baylor.edu 


John F. Pruski 


United States National dial 
we of Botany 
Smithsonian Institution, Washington ee 0166, U.S.A. 
pruskijohn@nmnh.si.edu 


Jason R. Singhurst 
Wildlife Diversity Program 
Texas Parks and Wildlife al 
Austin, TX 78704, 
jason. pee Sie state.tx.us 


Thymelaea passerina (L.) Cosson & J.Germain was first reported in the United States by 
Pohl (1955). In reporting T. passerina as new to Ohio, Vincent and Thieret (1987) pre- 
sented an expanded account of the occurrence of the species in the United States, men- 
tioning its presence in Illinois, lowa, Kansas, and Nebraska. Additionally, Thymelaea 
passerina has also been reported from Alabama (Webb et al. 1997), Mississippi (Wofford 
& DeSelm 1988), Washington (USDA, NRCS 1999), and Wisconsin (Harriman 1979). The 
species is not cited in any of the recent references or checklists treating the flora of Texas 
(Correll & Johnston 1970; Hatch et al. 1990; Johnston 1990; Jones et al. 1997), nor is it 
included in Diggs et al. (1999) in their flora of the north central part of the state. A de- 
scription of the species and pertinent synonymy follow. 

Thymelaea passerina (L.) Cosson & J.Germain, Syn. Fl. Env. Paris, ed. 2, 360. 1859. (Fig. 1). 

Stellera passerina L., Sp. Pl. 559. 1753. Complete synonymy given by Tan (1980). 

Taprooted annual to 55-60 cm tall; stems erect, slender, simple or more commonly with 
few to several ascending branches in distal half, glabrous or weakly pilose distally, yel- 
lowish green; leaves alternate, simple, exstipulate, sessile or nearly so, linear-lanceolate, 
7-15 mm long, 1-2 mm wide, stiffly chartaceous or subcoriaceous, glabrous or less com- 
monly abaxially weakly puberulent, apex acute, margins entire. Inflorescence axillary, 
flowers often 3 but ranging from 1-7 in distal leaf axils, cluster commonly subtended by 
2 green bracts, bracts lanceolate to broadly so, to ca. 1.5 mm long, basally ciliate; flowers 
bisexual, actinomorphic, perigynous, sessile, 2-3 mm long, corollas absent, sepals 4, weakly 


SIDA 19(2): 403 — 406. 2000 


404 SIDA 19(2) 


£ Dink 


. 
J r or 


losed fruit. [F 1 Thieret in Sida 12:77. 1987. Used with permission. ] 


petaloid, manifested as minute (ca. 0.5 mm long) ovate to lance-ovate greenish yellow 
lobes with obtuse apices on top of the urceolate, persistent hypanthium, hypanthium 
and calyx lobes substrigose, stamens episepalous, 8, bicyclic, included, filament much 
shorter than the anther thecae, ovary superior, apically hirsute-villous, subtended by a 
hypogynous disk, bicarpellate but unilocular by abortion, style one, terminal, short, ovule 
one, pendulous. Fruit a dry, indehiscent, 1-seeded, 1-locular capsule, apically beaked, en- 
closed within the persistent hium;seed ovoid, 2-3 mm long, brown to black. Chro- 
mosome number 2n = 18. 

This Eurasian weed of Thymelaea section Lygia (Tan 1980) has been reported within 
the United States from Alabama (Webb et al. 1997), Illinois (Mohlenbrock & Ladd 1978), 
lowa (Pohl 1955), Kansas (McGregor et al. 1986), Mississippi (Wofford & DeSelm 1988), 
Nebraska (Pohl 1955), Ohio (Vincent & Thieret 1987), Washington (USDA, NRCS 1999), 
Wisconsin (Harriman 1979), and is here newly documented for the flora of Texas. It has 
also been introduced into Australia. Flowering time for the North American populations 
ranges from (April) June to September. 


a, 


HOLMES ET AL., THYMELAEA PASSERINA NEW TO TEXAS 405 


Specimens cited: TEXAS. Denton Co.: approximately 5 mi E of Sanger on FM 455, 24 Jun 1999, 
Singhurst 8156 (BAYLU, TEX, US). Fannin Co.: 0.5 mi E of Haile Community on FR 1550,N 33.51173,W 
96.05437,5 Jun 1999, Holmes 10173 & Singhurst foil TEX, US). 


We believe that the species was likely introduced to Texas through the use of agricul- 
tural machinery imported from further north and used to harvest wheat, the dominant 
crop in both areas. The Denton County specimen is from the Cross Timbers and Prairies 
vegetational region of the state, an area characterized by slightly acidic to acidic sandy 
loam soils (Correll & Johnston 1970). Associated species included Dalea purpurea, 
Indigofera miniata (Leguminosae), Froelichia floridana (Amaranthaceae), Helianthemum 
georgianum, Lechea mucronata, L. tenuifolia (Cistaceae), Hypericum drummondii 
(Hypericaceae), and Krameria lanceolata (Krameriaceae). The Fannin County specimen 
occurred in the margins of roads and wheat fields in clay over limestone “chalk” on the 
Gober Limestone Formation of the Blackland Prairie vegetation region of the state.Com- 
mon associates included Asclepias asperula (Asclepiadaceae), Forestiera pubescens 
(Oleaceae), Hypericum perforatum (Hypericaceae), Rhus aromatica (Anacardiaceae), Se- 
dum pulchellum (Crassulaceae), and Sophora affinis (Leguminosae). 

The doc tati f Thymelaea passerina as new to Texas is not only a report of a 
new genus and species to the known non-cultivated flora of the state, but another fam- 
ily, the Thymelaeaceae. Jones et al. (1997), in their checklist of the vascular flora for the 
state, included two species of Thymelaeaceae, Daphne cannabina Wall. (= D. papyracea 
Wall. ex Steud. according to Huxley 1992) and D. cneorum L., but both of these are culti- 
vated and not known to escape in Texas. 


ACKNOWLEDGMENTS 


We thank James Solomon (MO) and Michael Nee (NY) for searching for specimens of 
North American Thymelaea passerina in MO and NY, respectively; and Aaron Goldberg 
(US) and Monique Reed for helpful comments on the manuscript. We are grateful to 
Tom Wendt (TEX) for providing other assistance and to Michael Vincent (MU) and John 
Thieret (KNK) for their helpful reviews and permission to use the illustration. 


REFERENCES 


Corrett, D.S. and M.C. Jounston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner. 

Dices, G.M, Jr., B.L. Liescome, and RJ. O’kennon. 1999. Shinners & Mahler's illustrated flora of 
north central Texas. Botanical Research Institute of Texas, Fort Worth. 

Harriman, N.A. 1979. Four additions to the Wisconsin flora. Michigan Bot. 18:143-145. 

Hatch, S.L., K.N. GANDHI and L.E. Brown. 1990. Checklist of the vascular plants of Texas. The 
Texas Agricultural Experiment Station, Texas A&M University, College Station. 

Huxtey, A. (Ed.-in-Chief). 1992. The new Royal Horticultural Society Dictionary of garden- 
ing, vol. 2,D to K, Macmillan Press, London & Stockton Press, New York. 

JoHnston, M.C.1990.The vascular plants of Texas. A list, up-dating the manual of the vascu- 
lar plants of Texas. Published by the author, Austin. 


406 SIDA 19(2) 


Jones, S.D., J.K. Wiper, and PM. Montcomery. 1997. Vascular plants of Texas:a comprehensive 
checklist including synonymy, bibliography, and index. University of Texas Press, 
Austin. 

McGrecor, W.L., TM. Barktey, R.E. Brooks, and E.K. ScHofieto (eds.). 1986.Flora of the Great Plains. 
University Press of Kansas, Lawrence. 

MoHtensrock, R.H. and D.M. Lavo. 1978. Distribution of Illinois vascular plants. Southern 
Illinois University Press, Carbondale. 

POHL, R.W. 1955. Thymelaea passerina,a new weed in the United States. Proc. lowa Acad. 
Sci. 62:152-154. 

Tan, K. 1980. Studies in the Thymelaeaceae Il: a revision of the genus Thymelaea. Notes 
Royal Bot. Gard. Edinburgh 38:189-246. 

USDA, NRCS. 1999. The PLANTS database (http://plants.usda.gov/plants). National Plant 
Data Center, Baton Rouge, LA 70874-4490 USA. 

Vincent, M.A. and J.W. THieret. 1987. Thymelaea passerina (Thymelaeaceae) in Ohio. Sida 
(25-20, 

Wess, D.H., H.R.DeSetm, and W.M. Dennis. 1997. Studies of prairie barrens of northwestern 
Alabama. Castanea 62:173-184. 

Worroro, B.E.and H.R.DeSeto. 1988. Distribution of and first report of Thymelaea passerina 
from the southeastern United States. Castanea 53:305-306, 


REDISCOVERY OF RIBES NIVEUM (GROSSULARIACEAE) 
IN COLORADO 


Timothy W. Chumley! and Ronald L. Hartman 
Department of Botany 
University of Wyoming 
Laramie, WY 82071-3165, U.S.A. 


While conducting a floristic survey of central Colorado (Chumley 1998), two collections 
of a gooseberry from the Cafion City area, Fremont County, proved to be Ribes niveum 
Lindl. These collections represent the rediscovery of a taxon known in Colorado from a 
single, neglected gathering byT.S.Brandegee in 1873 (Brandegee 697, Sinnott 1985). These 
new collections come from the drainage of Cottonwood Creek, 13 to 15 miles north- 
west of Cafion City. Wilson Creek, the Brandegee locality, is 3-4 miles to the east or ca.8 
miles northwest of Cafion City.Field work in 1997 verified that R.niveum still occurs along 
Wilson Creek. 

The Brandegee specimen was cited by Porter and Coulter (1874) as R. irriguum Dou- 
glas, a species of the Pacific Northwest. Rydberg (1906) excluded this taxon from his 
treatment of the Colorado flora. Presumably, he did not examine the material and ig- 
nored the taxon due to its great disjunction from its normal range (being one of several 
species “accredited to Colorado but not the intervening states," page xii). The occurrence 
of’R. irriguum" has been similarly ignored or overlooked in subsequent treatments of the 
flora (Harrington 1954;Weber 1953, 1967, 1990; Weber & Wittmann 1992, 1996). Sinnott 
(1985) examined the Brandegee collection in preparing his treatment of Ribes section 
Grossularia and determined it to be R.niveum, another species of the Pacific Northwest, 
rather than R. irriguum (R. oxyacanthoides L. ssp. irriguum (Douglas) Q.P. Sinnott). This is 
consistent with Porter and Coulter's description of the specimen, which matches R.niveum 
and not R. irriguum. Sinnott, however, failed to relocate the Colorado population. Exami- 
nation of the specimen (Brandegee 697) at the Missouri Botanical Garden confirmed his 
determination, and the new collections document its persistence in Colorado for over 
120 years. 

The normal geographical range of R.niveum is in three separate areas: the northern 
group of populations are in southeastern Washington, eastern and northeastern Oregon, 
and adjacent western and central Idaho;the middle group, southeastern Oregon, south- 
ern Idaho, and adjacent northeastern Nevada; and the southern group, west-central 
Nevada (Churchill, Lander, Pershing, and possibly Nye counties; Holmgren 1997; Sinnott 


Current address: Section of Integrative Biology, School of Biological Sciences, University of Texas at Austin, 
Austin, TX 78712, U.S.A. 


SIDA 19(2): 407 — 410. 2000 


408 SIDA 19(2) 


1985).The marked disjunction to the east or southeast is most curious. A possible clue to 
its occurrence in Colorado prior to settlement was found in an account of the Long 
expedition to the Rocky Mountains in 1820 (Goodman & Lawson 1995). Edwin James, 
the botanist of the expedition, collected a specimen of Ribes in the area that was deter- 
mined by Torrey and Gray as R. irriguum.A description of the fragment provided by Torrey 
and Gray (1838-1843) indicates that it was not R.niveum but most likely R. inerme Rydb. 
Goodman and Lawson were unable to locate the specimen and thus could not confirm 
its identity. The presence of R.niveum in the region therefore is not documented before 
Brandegee's collection in 1873. Prior to that time, Cafion City was a major gateway and 
supplier to the gold and silver mines of South Park and Leadville, and it is possible that R. 
niveum was introduced into Colorado from the Pacific Northwest during the 1860s by 
gold or silver miners. It is also possible that Native Americans may have been respon- 
sible for dispersal since the localities are close to old Ute trails into South Park. Brandegee’s 
collection may thus represent the parental population at or near its initial point of es- 
tablishment. 

The plants were in flower by mid-June, in fruit by late July. Fruits persisted through 
at least early October. They were found growing along creeks or dry washes in pinyon- 
juniper on sandy soils derived from gneiss. Interestingly, R. niveum in Colorado always 
grows in clumps with other woody taxa including Ptelea trifoliata L., Ribes cereum Dou- 
glas, Ericameria (Chrysothmanus) nauseousus (Pall.ex Pursh) G.L.Nesom & G.I, Baird, Rhus 
trilobata Nutt.,and Quercus gambellii Nutt. This could indicate dispersal of seeds by birds. 

In floral features, the Colorado material of R. niveum is a close match with material 
from the main geographical range.The most striking characters of R.niveum are those of 
the stamens.With the petals extended or erect, the stamens surpass them by 2 to 3 mm; 
filaments range from 7 to 9 mm in length. Furthermore, both the anthers and filaments 
are at least moderately pubescent with prominent silky hairs. All other Colorado species 
have glabrous filaments and anthers. Vegetatively, R. niveum has a distinctive rich reddish 
brown bark on new growth and is unarmed except for stout spines at the nodes. Ribes 
lacustre (Pers.) Poir. may have a similar hue, but the internodes are usually quite spiny. 
Another species, R. inerme, shares similarities in leaf shape, but the young twigs are pale 
yellow. With age the bark of most Colorado gooseberries becomes gray and exfoliates, 
exposing a dark reddish brown interior. 

Leaf blades of specimens of R. niveum collected at the three known sites in Colo- 
rado are moderately to densely covered with minute, stalked glands mixed with erect to 
curved eglandular trichomes. Those in the main populations to the west and northwest 
vary from glabrous and ciliolate to densely clothed in minute, erect to curved eglandular 
trichomes only,although the petioles often have stalked glands. Due to the fickle nature 
of glandular trichomes in many groups of plants and the variability in pubescence in 
populations over the main geographical range of this taxon, this difference may not be 
significant. 

In Weber and Wittmann (1996), R. niveum keys to lead 10b, where it matches fruits 


CHUMLEY AND HARTMAN, REDISCOVERY OF RIBES NIVEUM 409 


and flowers glabrous externally, but differs in anthers purple, greenish with age vs.white. 
roi anther color, it would key to R.inerme (1 1a), although not easily. For an excellent 
treatment of Ribes that is largely relevant to Colorado, the reader is referred to the /nter- 
mountain Flora (Holmgren 1997). All but two Colorado taxa are treated: R. americanum 
Mill, a species of the plains and northeastern North America, and the questionable in- 
troduction (W. Jennings, Louisville, CO, pers. comm.), R. divaricatum Douglas. Ribes 
coloradoense Coville is placed in synonymy with the major disjunct (to the northwest), R. 
laxiflorum Pursh. 

It is always possible that additional populations of Ribes niveum remain to be dis- 
covered east of the Continental Divide in Colorado. Two recent floristic projects have 
contributed substantially to our understanding of the ebtaloution of species of Ribes, 
and to vascular plants in general, in this area (see http://www.rmh.uwyo.edu). The first, 
the Central Colorado project, which led to the rediscovery of R niveum (Chumley 1998), 
included the Mosquito and Rampart ranges and Pikes Peak. It extended from Morrison 
(near Denver) west to Webster and Hoosier passes, south on the west side along the 
crest of the Mosquito Range to near Salida and south on the east side to Pueblo. The 
second, the Sangre project, included the Sangre de Cristo and Wet mountains, Mesa de 
Maya, and the Spanish Peaks. It was to the south from the first area, with the Arkansas 
River, in part, forming the boundary in common. Canon City is on the Arkansas River, 
north side, midway along this line of contiguity. The Sangre project extended from Pueblo 
west to Monarch Pass, south on the west side to North Pass, Saguache, San Luis, and the 
New Mexico line and south on the east side to Trinidad and Branson (collections mostly 
by B. Elliott and R. Hartman). Together, these two study areas cover nearly 10,000 mi? for 
which ca. 27,500 numbered collections were obtained. In the process, 322 populations 
of nine of Ge 13 ea of Ribes reported for Colorado (Weber & Wittmann 1992 and 
this report) were sampled during four field seasons (1995-96, 1998-99). Despite the high 
level of ene, in collecting, no new sites for R. niveum were discovered south of the 
Arkansas River. 


a eas late COLORADO. Fremont Co.: Sand Gulch south along a drainage intersecting 
15 air mi NW of Canon City, 16 Jun 1995, Chumley 895 (RM); Cottonwood Creek, 
ca. 3.135 air mi an of Cafion City, 25 Jul 1995, Chumley 2454 (RM); along Wilson Creek and sur- 
rounding hills to the east, ca. 7.8 air mi NW of Cafon City, 28 Jun 1997, Chumley 6887 (RM). 


ACKNOWLEDGMENTS 


This work was funded by a cost-share agreement between the Rocky Mountain Her- 
barium and the U.S.D.A., Pike and San Isabel National Forests, and grants from the Colo- 
rado Natural Areas Program and the Colorado Native Plant Society. It was part of a M.S. 
thesis by the senior author at the University of Wyoming. The authors would like to thank 
the following herbaria for the use of their collections and facilities: University of Colo- 
rado Museum (COLO), University of New Mexico (UNM), University of Texas Plant Re- 
source Center (TEX), and Missouri Botanical Garden (MO). 


410 SIDA 19(2) 


REFERENCES 


CHumtey, TW. 1998. A floristic inventory of the East Slope, central Colorado. M.S. thesis, 
Botany. University of Wyoming, Laramie 

Goopman, G.J.and C.A. Lawson. 1995. Retracing Major Stephen H.Long’s 1820 expedition: 
The itinerary and botany. University of Oklahoma Press, Norman. 

Harrincton, H.D. 1954. Manual of the plants of Colorado. Sage Press, Denver. 

Houmaren, PK. 1997.1n:A.Cronquist, N.H. Holmgren, and P.K.Holmgren.|ntermountain flora: 
Vascular plants of the intermountain West, U.S.A. Volume 3A. New York Botanical Gar- 
den, Bronx. Pp. 12-26. 

Porter, T.C. and J.M. Coutter. 1874. Synopsis of the flora of Colorado. Department of the 
Interior, U.S.Geological and Geographical Survey of the Territories, Miscellaneous Publ. 
no. 4.Washington, D.C 

RypBerG, P.A. 1906. Flora of Colorado. Agric. Exp. Sta. Agric. Coll. Colorado Bull. 100. 

Sinnott, Q.P. 1985. A revision of Ribes L.subg. Grossularia (Mill.) Pers. sect. Grossularia (Mill.) 
Nutt. (Grossulariaceae) in North America. Rhodora 87:189-286. 

Torrey, J.and A. Gray. 1838-1843. A flora of North America.Wiley & Putnam, New York. 

Weer, W.A. 1953. Handbook of plants of the Colorado front range. University of Colorado 
Press, Boulder. 

Weer, W.A. 1967.Rocky Mountain flora. University of Colorado Press, Boulder. 

Wester, W.A. 1990. Colorado flora: Eastern slope. University Press of Colorado, Niwot. 

Weeer, W.A.and R.C. Wittmann. 1992. Catalog of the Colorado flora: A biodiversity baseline. 
University Press of Colorado, Niwot. 

Weser, W.A.and R.C. WITTMANN. 1996. Colorado flora: Eastern slope. Revised edition. Univer- 
sity Press of Colorado, Niwot. 


ALOPECURUS MYOSUROIDES AND 
SCLEROCHLOA DURA (POACEAE) NEW TO LOUISIANA 


John K. Saichuk 
LSUAGCenter 
Cooperative Extension Service 
Crowley, LA 70527-0497, U.S.A 
Charles M. Allen 
Biology Department 
Oe of Louisiana at Monroe 

Monroe, LA 71209, U.S.A 
Wiliam D. pecs 
iology Departmen 
ube of Louisiana at eae 
Lafayette, LA 70504-2451, U.S.A. 


Two recent collections from Morehouse Parish in northeast Louisiana are apparently the 
first records for the state. Specimens of Alopecurus myosuroides Huds. and Sclerochloa 
dura (L.) Beauv. were collected from the same field on the same day. Neither species is in 
Allen (1993) or Thomas and Allen (1993). Sclerochloa dura (L.) Beauv was reported for 
Louisiana by MacRoberts (1977) and then excluded by Allen (1980). Brandenburg et al. 
(1991) did not find any records for this species from Louisiana but did report it for all 
three adjacent states: Arkansas, Mississippi, and Texas. Alopecurus myosuroides Huds. is 
listed by the USDA-NRCS (1999) database from adjoining Mississippi and Texas as well as 
Alabama, Kansas, New Mexico, and most east and west coast states. 


kL 


Voucher specimen (Alopecurus myosuroides Huds.): LOUISIANA.M Parish: Rice field area 
on the Harold Tucker farm, about 7.7 mi due S of the Arkansas/Louisiana state line near the inter- 
section of Sunshine and Tucker roads, E side of Tucker road about 0.5 mi S of Sunshine Road, field 
fallow through the winter and not flooded, scattered throughout the field and along the edges, 27 
Apr 2000, Saichuk s.n. (LAF, NLU). 


Voucher specimen (Sclerochloa dura (L.) Beauv.): LOUISIANA. Morehouse Parish: Rice field area 
on the Harold Tucker farm, about 7.7 mi due S of the Arkansas/Louisiana state line near the inter- 
section of Sunshine and Tucker roads, E side of Tucker Road about 0.5 mi S of Sunshine Road, field 
fallow through the winter and not flooded, growing along the field road, 27 Apr 27 2000, Saichuk 
s.n. (LAF, NLU). 


REFERENCES 


ALLEN, C.M. 1980. Grasses of Louisiana. University of Southwestern Louisiana, Lafayette. 
Auten, C.M. 1992. Grasses of Louisiana,second edition.Cajun Prairie Habitat Pres. Soc., Eunice, 


SIDA 19(2): 411 — 412. 2000 


412 SIDA 19(2) 


Brandensurc, D.M.,J.R. Estes, and J.W. THieret. 1991. Hard grass (Sclerochloa dura, Poaceae) in 
the United States. Sida 14:369-376. 

MacRoserts, D.T. 1977. Additions to the Louisiana flora. Sida 7:220-222. 

THomaAS, R.D. and C.M. Atten. 1993. Atlas of the vascular flora of Louisiana, Vol. 1: Ferns and 
ferns allies, conifers, and monocotyledons. Louisiana Department of Wildlife and Fish- 
eries, Baton Rouge, LA. 

USDA, NRCS. 1999. The PLANTS database (http://plants.usda.gov/plants). National Plant 
Data Center, Baton Rouge, LA 70874-4490 USA. 


GAMOCHAETA SIMPLICICAULIS 
(ASTERACEAE: GNAPHALIEAE) IN GEORGIA 
Guy L.Nesom 


North Carolina Botanical Garden 


University of North Carolina 
Chapel Hill, NC 27599-3280, U.S.A. 


Nae 


Two collections extend the known range of Gamochaeta simplicicaulis (Willd.ex Spreng. 
Cabrera into Georgia. This native South American species was previously first reported 
from North America in North Carolina, South Carolina, Florida,and Alabama (Nesom 1999). 
An additional collection from Florida confirms its presence in that state; the previously 
cited Florida specimen also was collected in Walton County by H.A. Davis. These new 
records are unequivocal in identity. The hospitality of the staff at BRIT/VDB is greatly 
appreciated. 

Voucher specimens: FLORIDA. Walton Co.: 4.5 mi S of Freeport, by overflow pond, 12 Jul 1972, 
Davis 16204 (VDB). GEORGIA. Bulloch Co.: wet pine woods 1 mi S of Statesboro, 8 Aug 1975, Kral 
56228 (VDB). Wayne Co.: longleaf pine-turkey oak sandridge by US 301 at S side of Jesup, 8 Aug 
1975, Kral 56320 (VDB). 


REFERENCE 


Nesom, G.L. 1999. Gamochaeta simplicicaulis (Asteraceae: Gnaphalieae) in four southeast- 
ern states and new for North America. Sida 18:1259-1264. 


SIDA 19(2): 413. 2000 


414 SIDA 19(2) 


BOOK NOTICES 


Bossy J. Warp. 1999. A contemplation Upon Flowers: Garden Plants in Myth and Lit- 
erature. (ISBN 0-88192-469-5, hbk.). Timber Press. Inc., The Haseltine Building, 133 
S.W.Second Avenue. Suite 450. Portland, OR 97204-3527, U.S.A. (503-227-2878, 503- 
227-3070 fax; www.timberpress.com). $24.95 hbk. 447pp. Line drawings. 


This work represents extensive reading and intensive delving into the subject. Not only does the 
author describe the genus and species of some eighty-three plants and their natural origins, he 
also relates them when possible to native wild species. Beginning with etymological accounts of 
scientific names and common names, he narrates myths and legends and cites poetry and prose 
referring to the species from the works of authors throughout the range of Western literature. Well- 
indexed by authors’ names and, separately, by plants (binomial as well as common), this work be- 
sides being pleasurable reading may have particular value as a source of tales and quotations to 
enliven talks to general audiences.—Joann Karges. 


Haroid W. Lawrence. 1999. Douglas Chandor: An English Artist and His Texas Garden. 
(No ISBN, hbk.) Antler Press, 317 Cleveland Ave., Weatherford, TX 76086, U.S.A. (No 
price given), 173 pp.,b/w photographs. 

Douglas Chandor painted portraits of royalty, of world leaders, of business tycoons and society 

matrons, and in the rocky ground of Weatherford, Texas, he created an extensive garden that even- 

tually reached national recognition."White Shadows," later Chandor Gardens, opened to the public 
in the 1940s. Eclectic in style, the gardens employed principles of English landscape gardening 
with many Oriental features. Chandor is quoted as having said,"“God gave me the talent to paint 
the pictures so that | could sell them to get the money to build the garden.” 
Contents.—Introduction. 1) Early Years in the U.S.;2) Chandor and Kuteman Family Matters; 3) 

Douglas and Ina Chandor: A life shared; 4) The Studio and the Garden; 5) Public Response to the 

Garden; 6) Journals; 7) Chandor's Garden EDI). Pppenalx ~ BOR Estiel mppenals B: Malcolm 

Vaughn's Brochure; Appendix C:A list of Doug g Bi Joann Karges. 


Andreé Millar. 1999. Orchids of Papua New Guinea. (ISBN 0-88192-438-5, hbk). Timber 
Press. Inc., The Haseltine Building, 133 S.W. Second Avenue, Suite 450, Portland, OR 
97204-3527,U.S.A.(503-227-2878, 503-227-3070 fax; www.timberpress.com). $34.95 
hbk. 128 pp, 271 color photos, 14 b/w illustrations, 1 map 


From the close-up and intimate photos or towers, to habitat shots to beautiful landscapes, the 271 
color photographs by Ron and b Sinead kay are enough to make you want to grow orchids or 
at po visit Papua New Guinea. What a passion or should | say an obsession! 

om the dustjacket.—" " thoroughly revised and updated pag al Millar's Sales |. on 
the same subject, Millar is the first to attempt this vast subject since 
Guinea ended more than sixty years ago. Covering both ere ne andi ial orchid = thie ne 
is a truly personal account of the author's eles “— the oneness Decades of siuidy<cobse 
sion? love?—of the orchids native to th nd have produced an incredibly 
detailed and authoritati = fecona orwial grows where and ea it Boks ike ‘The 203 species are 
accounted for in 26 subtri t tions asin > ee ren S leuidacede 
of German New Guinea; species in each section are in alphabetical order.—Barney Lipscon 


SIDA 19(2): 414. 2000 


GEORGE JONES GOODMAN 
(1904-1999) 
A MEMOIR 
Cheryl A. Lawson 


1 W. MacArthur 
Shawnee, OK 74804, U.S.A. 
The University of Oklahoma and the State of Oklahoma lost their most distinguished 
and renowned plant taxonomist on May 23, 1999, with the death of Dr. George J. 
Goodman, Regents Professor Emeritus and Curator Emeritus of the Bebb Herbarium. 

Dr. Goodman was born November 5, 1904, in Evanston, Wyoming, to Arthur and 
Elizabeth Jones Goodman. As a young girl his mother had come in 1886 to this country 
from Wales, and Dr.Goodman would proudly point out that half of him was a first-gen- 
eration American.A ranch, located some twenty-five miles south of Evanston and home- 
steaded by his Grandfather Goodman in 1883, is where Dr.Goodman spent the first five 
years of his life. His love of the outdoors and the West thus began. 

After graduating from Evanston High School in 1922, Dr. Goodman, who had no 
plans to attend college, worked as a ranch hand, as a clerk ina drug store,as a sacker ina 
grain elevator, as a salesman of pianos and phonographs and of contract printing, and as 
a compass man for the U.S. Forest Service in the Kaibab of Arizona. About 1925 and 
during the time he was delivering groceries in Ogden, Utah, Dr 's father bought 
another ranch which was located due south of Evanston a mile over the Utah line. Dr. 
Goodman went to that ranch in the spring of 1926, and very soon thereafter an event 
occurred that would begin his botanical career which lasted nearly three-quarters of a 
century! 

Dr.Edwin Payson, a botany professor from the University of Wyoming who had done 
his graduate work on the Cruciferae at the Missouri Botanical Garden under Dr. Jesse M. 
Greenman, and his wife Lois came to Evanston to find someone to take them up into the 
Uintah Mountains to collect plants. It was there that they learned of George Goodman, 
who by that time had already climbed several peaks in the Uintah Mountains. They made 
their way to the Goodman Ranch and camped there for a week or two. Dr. Goodman 
took them up to Stillwater Fork and Hayden’s Peak where Sereno Watson had collected 
in 1869 on the King Expedition. 

Impressed by the young man who became their guide, the Paysons suggested to 
Dr.Goodman that he should go to college and offered to give him a room for the year 
and to try to find him a job on the Laramie campus. After a discussion with his parents, 
who decided a year of college couldn't hurt, Dr.Goodman agreed to go.While still camped 
at the Goodman Ranch, the Paysons wrote to Dr. Aven Nelson, botany professor and 
herbarium curator, and told him they had someone they thought would make a good 


SIDA 19(2): 415 — 420. 2000 


416 SIDA 19(2) 


person to work in the herbarium mounting plants. Dr. Nelson wrote back, and when the 
Paysons told Dr. Goodman he had the job in the herbarium, he had no idea what an 
herbarium was. He didn't ask, but learn he did! 

The Medicine Bow Mountains, located just to the west of Laramie, became a favor- 
ite collecting site during the three years and a summer Dr. Goodman spent obtaining 
his bachelor of arts degree with honors in botany. It was to these mountains that he 
would return many, many summers throughout his life. Whether he was there to teach 
at the University of Wyoming Science Camp or just to enjoy his cabin at 10,000 feet, he 
was in the mountains he loved. 

In 1929 Dr. Goodman received a Rufus J. Lackland Fellowship from Washington 
University in St. Louis to do graduate work at the Missouri Botanical Garden under Dr. 
Jesse M.Greenman.His new and soon-to-be closest friend was C.L. Hitchcock 
(Hitchy) who would become professor of botany and curator of the herbarium at the 
University of Washington, Seattle,and a major contributor to the Flora of the Pacific North- 
west. Lured by Eriogonum and Lycium, Goodman and Hitchcock headed out to collect in 
the West in the summer of 1930. 

This trip which began in June followed Route 66 through southern Missouri, Okla- 
homa, and the Texas panhandle. This was the first time Dr.Goodman had been in Okla- 
homa, and while no plants were collected until they reached Portales, New Mexico, on 
June 14,he remembered seeing in Oklahoma a curious, dicotyledonous plant with flow- 
ers that looked like an orchid. Three years later, after coming to the University of Okla- 
homa, he learned that he was looking at Krameria. 

The Goodman/Hitchcock collecting trip was bountiful. Over five-thousand sheets 
were collected and later divided into sets which were then sold mostly to large herbaria 
throughout the country in order to finance the trip. Sites such as the Chiricahua Moun- 
tains of Arizona and the Abajo Mountains of Utah had been infrequently collected. From 
their collections a few new taxa resulted. 


lengthily to two gentlemen on where to buy the cheapest gas in Monument Valley,One 
of the men proved to be the vice-president of Bethlehem Steel! Hitchy never did agree 
with Dr.Goodman on whether they had seen a mountain lion or a wolf in the Kaibab forests 
in northern Arizona. Nor could Hitchy believe that the herds of horses there were wild. 

The two repaired in one day as many as fourteen blowouts on the tires of their 
panel truck, lived on strawberry jam, and had a lifetime’s worth of fun. Even as late as 
1992 Dr.Goodman clearly recounted for me the events of that trip. 

Dr. Goodman completed his doctoral degree at Washington University and the 
Missouri Botanical Garden in 1933 and came once again to Oklahoma, but this time as 
an assistant professor of botany and curator of the herbarium at the University of Okla- 
homa, Norman. However, a job offer in 1936 from lowa State College took Dr.Goodman 
to Ames. 


LAWSON, GEORGE JONES GOODMAN 417 


During three of the summers in lowa Dr.Goodman had the opportunity to return 
to the southwest to collect in the Lukachukai Mountains in the northeast corner of Ari- 
zona. He and Hitchy had seen these mountains in the distance as they drove north to- 
ward Mexican Hat, Utah. Very few botanists had by that time collected in the Lukachukai 
Mountains. Among those who accompanied him was Lois Payson, who had been instru- 
mental in beginning Dr.Goodman’s botanical career and who had been widowed in the 
spring of 1927.The group camped near Canyon de Chelly and collected on the Navajo 
Reservation. Lasting friendships developed with the wonderful Navajo.On one occasion 
at least they were invited to an Indian ceremony, which lasted the entire night. This asso- 
ciation with the Navajo people made a lasting impression upon Dr. Goodman. 

Fortunately for those of us who came later as students to the University of Okla- 
homa, Dr.Goodman accepted in 1945 the offer from Dr. George L. Cross, a former col- 
league and close friend in the OU Botany Department and by then president of the 
University of Oklahoma, to return to OU and the Bebb Herbarium. During his tenure as 
curator, Dr.Goodman built the collection from a few thousand plant specimens into one 
that before his retirement in 1975 would amass nearly a quarter-million specimens. To- 
day the Bebb Herbarium houses the world’s finest collection of the flora of Oklahoma, 
along with strong holdings of the flora from surrounding states, the southwestern United 
States, the Great Plains, and Mexico. 

The University of Oklahoma recognized Dr.Goodman’s exceptional contribution to 
the University and to the Bebb Herbarium by appointing him Regents Professor in 1967. 
He was also awarded the Distinguish Service Citation, the University’s highest honor,in 1978. 

Dr. Goodman possessed not only a lively intellectual curiosity and an exhaustive 
knowledge of plants, but also a warmth and cordiality that would make the herbarium a 
place where colleagues and students gathered over the years for meetings, parties, and 
informal coffee and conversation. 

During his career as a botanist Dr.Goodman came to be known as a leading expert 
in the field of plant taxonomy for Oklahoma and the western United States. He authored 
seventy-three publications, described thirty-six new plant taxa, made eleven new com- 
binations, and had four plants named for him. 

Dr.Goodman was a charter member of the American Society of Plant Taxonomists, 
the International Association of Plant Taxonomists, the Society for the Study of Evolution, 
the Southwestern Association of Naturalists, and the Colorado-Wyoming Academy of 
Science.|n addition, he was a member of Phi Beta Kappa, Phi Kappa Phi, Phi Sigma, Sigma 
Xi,and the Oklahoma Academy of Science.He received the Phi Sigma Ortenburger Award 
and the Oklahoma Academy of Science Award of Merit in addition to the Distinguished 
Service Citation from the University of Oklahoma. 

| was Dr.Goodman's last graduate student. We continued to work side by side both 
in the field and in the herbarium for nearly a quarter century after his so-called “retire- 
ment.” During the field work for our book, Retracing Major Stephen H. Long’s 1820 Expedi- 
tion: The Itinerary and Botany (OU Press, 1995), we traveled over 10,000 miles through 


f4hnc th Diatto Di 


George J.G 


hos 
(He Lidsm O 


rtne sOutn Fiatte iver, 


Douglas Co., Colorado, June 30, 1981. 


8Lb 


(2)6L Vals 


LAWSON, GEORGE JONES GOODMAN 419 


George J. Goodman, spring, 1975. 


420 SIDA 19(2) 


Nebraska, Colorado, New Mexico, Texas, and Oklahoma. Our field trips in the West follow- 
ing the route of this expedition were full of episodes which we recounted in our book. 
Working as | did with Dr.Goodman was an incredible experience filled with indelible memo- 
ries. 

Dr. Goodman was as enthusiastic about life as he was about plants. Once when 
asked how he would describe his life, Dr. Goodman replied,"It’s been a blast!” It, too, was 
a“blast” for those of us lucky enough to have had him in a part of our lives! 

In August, 1999, Dr.Goodman’s ashes were taken by Marcia, his wife of fifty years, his 
daughter Sula Grace Henrichsen, and other relatives and friends to the vicinity of West 
Glacier Lake near Centennial, Wyoming, in the Medicine Bow Mountains. The circle closed, 
and he was back for perpetuity in the mountains he loved. 


NESTA DUNN EWAN 
NOVEMBER 8, 1908-SEPTEMBER 13, 2000 
A MEMOIR 
Anne S. Bradburn 
Tulane University Herbarium 
Dept. of Ecology and Evolutionary Biology 


University 
New Orleans, LA 70118-5698, U.S.A. 


The History of Natural History has lost another champion. Although she was most often 
in the background, those of us who were fortunate enough to know the Ewans were 
always aware that Nesta’s contributions to Joe’s writing were indispensable. She was a 
meticulous researcher spending days tracking down the most minute details. Hers was 
the final authority on matters of grammar, style, and punctuation. 

Born near Saskatoon, in Saskatchewan, Canada, Nesta with her family moved to 
California where she attended the University of California at Los Angeles, earned a de- 
gree in botany in 1933, and met another young botanist, Joseph Ewan, whom she mar- 
ried in Reno, Nevada in 1935. They produced three daughters, Kathleen Harris of 
Mandeville, Louisiana who with her husband Dick did so much to make their latter years 
enjoyable, Dorothy Nemecek of Chattanooga, Tennessee, and Marjorie Ewan who now 
lives in Albuquerque, New Mexico. All share their parents love of nature, if not book col- 
lecting, and have happy memories of family camping trips. There are five grandchildren. 

During their next 64 years together the Ewans also produced nearly 500 titles. Be- 
ginning in 1963 with John Lyon, Nurseryman and Plant Hunter, and his Journal, 1799— 
1814” published in the Transactions of the American Philosophical Society (n.s. 53: pt.2.), 
Nesta was frequently listed as co-author. Her participation was more widely recognized 
after “Ewania: the writings of Joe and Nesta Ewan’ was published in 1989 (The American 
Botanist, Booksellers, Chillicothe, Illinois). 

During their 40 years at Tulane, Nesta was famous for providing fabulous Sunday 
brunches to generations of graduate students. David White, now Professor of Biology at 
Loyola University, fondly remembers devouring stacks of buckwheat cakes loaded with 
jams and jellies, a tradition which continued after the Ewans moved to the Missouri Bo- 
tanical Garden in Saint Louis. 

Once established in the old Museum building where they had palatial quarters and 
for the first time ample space for their burgeoning library, Nesta told me “When | die | 
don't have to go to heaven, | am already there.” Joe and Nesta were both great lovers of 
plants, animals, and the great outdoors, and they reveled in the morning walks through 
the splendor of the Gardens. So while we at Tulane missed them sorely, we were glad 
indeed that they had found a wonderful place to continue their studies. 


SIDA 19(2): 421 — 423. 2000 


SIDA 19(2) 


422 


==s-osS Soest 


—— is ares iron 


i 


Nesta Dunn Ewan 
Mandeville, Louisiana 


BRADBURN, NESTA DUNN EWAN 423 

A lasting tribute was made to the couple in 1994 when they were jointly presented 
the Henry Shaw Medal by the Garden for their many contributions to botany and the 
history of science. 


Small in statue, Nesta was a giant of a woman.A true personification of the adage 
“Behind every great man, there is an equally great woman.” We shall not see her like 
again soon. 


Nesta and be Ewan 
September 1994 


424 SIDA 19(2) 


BOOK REVIEWS 


Ricard V. FRANCAvIGUA. 2000. The Cast Iron Forest: A Natural and Cultural History of the 
North American Cross Timbers. (ISBN 0-292-725 15-9, hbk.;0-292-725 16-7, pbk,). 
Univ.of Texas Press, PO. Box 7819, Austin, TX 78713-7819, U.S.A (512-47 1-4032).$45.00 
hbk., 24.95 pbk. 269 pp., 15 color and 52 b&w photos, 8 maps, 12 line drawings, 


"Ss gn 


More than anything else, The Cast Iron Forest portraits the Cross Timbers, conceptualized as a re- 
gion, as having a life of its own, growing and changing with time and captivating those that enter, 
including the author.While the book is an objective analysis of historical maps, travel diaries, eco- 
nomic statistics, and census records, many passages reveal an intensely personal fascination with 
the Cross Timbers. Richard Francaviglia is a historical geographer at the University of Texas at Ar- 
lington. Like many professionals and others transplanted to Texas, he has been nourished by Texas 
pride and now finds that his roots have grown deep into the Cross Timbers sandy cla 

Francaviglia begins the book by developing the concept of “Cross Timbers" from several per- 
eee ae Cross Timbers...are a forested archipelago largely surrounded by a sea of prairie. 
Centered roughly between the 97" and 98" meridians, the Cross Timbers vegetation comprises 
generally north-south trending belts of scrubby oak trees.” Next,”...the Cross Timbers typically 
appears as dense stands of post oak and blackjack oak trees that fey exceed pene thirty feet in 
height, but that are visible for a considerable distance across the prairie.” And”...the term...Cross 
Timbers referred to a large area that consisted of a swath of trees stretching north er Wace along 
the Brazos River of Texas and extending far north into. ..present-day Oklahoma."He th 
the natural history—the geology, soil, and climate correlated with the vegetation, and the impor- 
tant constituent species. The conclusion is:"...they are considered a separate region largely be- 
cause they are mostly surrounded by prairie, and would otherwise would not be so noticeable. 
Throughout the Cross Timbers, then, the distinctive oak forests are framed by prairie and devel- 
oped on sandstone [emphasis his]." 

The bulk of the book, however, is an examination of historical maps and writings to trace the 
history of the conceptualization of th 1 the minds of the Native Americans, Spanish/French 
explorers, and Anglo settlers and ae tnd finally the contemporary urbanites. This is the real 
strength of the book and a significant contribution to the non-technical literature. In so doi 
Francaviglia is able to trace the perceptions through 5- and 10-year intervals. Use of these rene 
data necessarily involves interpretations of why certain features were or were not added to maps 
and diaries, and readers will want to keep that in mind. However, his interpretations appear to be 
reasonable and without particular ideological bias. Especially fascinating is the argument that pre- 
vailing perceptions (e.g."dangerous and evil,""barrier to travel,”"plentiful source of wood and game") 
often influenced political, economic, and cultural responses of the public and governments. 

As a reader not interested so much in the pitone gaa | ae on in some of the 
detail.|t seemed to me that surely every map hed before 1900 is 
described. In fact, the author appears also to have aren somewhat lost in the details and intro- 
duced some inconsistencies. For example, he discusses the Cross Timbers on important maps of 

1849 and 1853, concluding (p. 104),"By the mid-nineteenth century, the Cross Timbers appear on a 
wide variety of popular maps” and “Historical geographer Ralph Brown nee that the Cross 
Timbers ‘achieved much notoriety of a favorable sort’ by the mid-nineteenth century.” However, 
just two paragraphs later (p. 106) states, “By the mid-nineteenth century, the Cross Timbers thus 
began to eck their status as a landmark.” 

uthor's transition to the twentieth aaa is aed by a move from documennng 
travels, nace skirmishes, and log-cabin architectur 


vt rh t 


SIDA 19(2): 424. 2000 


BOOK REVIEWS 425 


popular regional identity, and increasing conservation awareness. Even regional literary works, 
graphic art, and music are reviewed. One of une nicest features of the book is,in the last chapter, a 
four-page bulleted eee of the top nd a six-page summary of the author's interpre- 
tation of the region's hist 

Before The Cast Iron oe was written, the literature on the Cross Timbers region was largely 
scattered and not readily available. However, this book provides, without being academically tech- 
nical, references to most or all of the historically significant writings. Although enthusiasts of re- 
gional heritage will find it especially interesting, it is essential reading for anyone wanting a good 
overview and knowledge of as geulc and ental ey of this “cast iron” region.—Roger W. 
Sanders, Associate Collecti g titute of Tex 


Dennis W. Woop anb. 2000. Cont PI S tics, Third edition. (1-883925- 
25-8, hbk.). Andrews University Press, 2 3 Information Services Bldg., Berrien Springs, 
MI 49104-1700, U.S.A (616-471-6134, http://www.andrews.edu/press) $64.99 hbk. 
560 pp., numerous b&w photos and line drawings, CD of over 4,700 color images 


pean Plant Systematics flows from Woodland’s en uals to fill the needs of his 
wn students. His dea in wating a text is to provide“a well-illt d, bro ao; view, beginning text 
at would th wherever they may live in the world, sufficie 1d 
ing of aceclat sclents that would utilize the changing world of global Pa ene "In some ways 
he has succeeded and in others he has not. 
book is divided roughly into three portions. Printed page tabs, which mark the chapters 
and other divisions, are a helpful innovation. The first four chapters cover an introduction of sys- 
tematics as a subject, nomenclature, identification tools, and field and herbarium methods. The 
second section (5 chapters) is by far the largest. It encompasses an extensive survey of pterido- 
d angiosperm families with each group preceded by an explanation of 
special characteristics and terms. The last six chapters cover more advanced topics: history of the 
field, including contemporary systems and methods of analysis; pertinent literature; origins of the 
vascular plant groups;a ae of morphological, chemical, and other criteria; conservation issues; 
and the role of botanical g ns 
The field and ee potion (early chapters) is thorough, but the other chapters on 
principles sometimes lack depth. For example, speciation is covered in one page and one diagram. 
oe in the discussion of phenetic and cladistic methods, relevant terms are defined and sample 
nd ms and cladograms are illustrated. Hoe there is neither an attempt to explain how 
the disances are calculated nor examples worked out to help the student assimilate the methods. 
An appendix with simple four-taxon problems, mathematical algorithms, character and distance 
oo and step-by-step solutions would be very helpful. 
oodland is to be congratulated for the innovative chapter on conservation. In it he covers 
the Ba conservation issues that pertain to preserving biodiversity, including genetic aaa 
ing Since oes Somewelony is soe phase by texts in plant Syeteiietiees but is an endeavo 
that systematis to during their careers. His survey of b | gardens is a 
nice addition, but many instructors would rather exchange this information for more detail on 
principles. The chapter on taxonomic criteria is also strong because each major topic is contrib- 
uted by an authoritative researcher: Nels Lersten on plant anatomy, Rolf Sattler on morphology, 
Loren Rieseberg on molecular systematics, Cliff Crompton on palynology, Peter Holland on ecol- 
and biogeography, and Woodland on cytolo 
The weakest portion is - large, page- pomtntive survey of families. Each family is treated 


SIDA 19(2): 425. 2000 


426 SIDA 19(2) 


in a box with a set of drawings and short »stem,and/or chemical charac- 
ters; akg reproductive structures; distribution; economic value: and fossil record. Woodland is to 
ommended for sneer ae meat Mop lcat ang ante “adapted families of angiosperms and, 
thus, giving a global perspectiv rally of poor quality. Much 
page space is wasted and ee for printing has thickened and ‘ised lines and obscured de- 
ne that might have ce in the originals. Most of the close-ups do not show a distinctive fea- 
at Anther flaps are not shown on of Lauraceae 
ee panies of fe 27 coun) eee aee homosporous ferns are treated; neither 
the drawings nor the descriptions are especially helpful to the novice in differentiating or recog- 
nizing any particular segregate family. To a certain extent, the included CD of the University of 
Wisconsin's Photo Atlas of the Vascular Plants makes up for these deficiencies, but | found the inter- 
face to be less than self-explanatory and the images to include too few floral dissections. 

There is a number Sean minor ploplens (the examples listed here either remain un- 
corrected from the second edition or were introduced with the third). The hardback binding is low 
quality and probably will not hold un up wall with extended use. The typesetting was poorly proofed. 
For example, on page 410 one finds “from a wide spectra [sic] of sources” and on page 416 “phyl- 
lomes (e.g., leaf and leaf homologs), and trichomes (homes [sic])."In Table 1.1 illustrating the taxo- 
nomic hierarchy, the Magnoliineae is listed as the suborder under the Asterales and above the 
Asteraceae! A aot of the i were redrawn with permission from various sources, espe- 
cially Cronquist’s An Integrated System of Classification of Flowering Plants. In an apparent oversight, 
Correll and a s Flora of the eee he in which Pricilla Fawcett's plate of Zamila pumila 
was published (Fig. 25), is not cited as the source of the original from which the illustration of 
Zamia (p.64) was undoubtedly redrawn. 


| 


es they posed to illust 


wg 


hether other instructors will find this book useful for adoption will depend on how closely 
their cae backgrounds and needs and their own teaching philosophy matches Woodland's. 
Contemporary Plant Systematics would be more cost-effective if the family surveys (and illustrated 
glossary of morphological terms) were replaced with a short, several-page synopsis of groups of 
families and if the book were designed to use a professionally illustrated companion atlas, such a 
Wendy Zomlefer's Guide to Flowering Plant Families.—Roger W. Sanders, Associate Collections Men. 
ager, Botanical Research Institute of Texas. 


M.S. Mani and J.M. Saravanan. 1999, Pollination Ecology and Evolution in Compositae 
(Asteraceae). (ISBN 1-57808-058-4, pbk.). Sciences Publishers, Inc., PO. Box 699, May 
Street, Enfield, NH 03748, U.S.A.$49.50 + shipping. 166 pp.Line drawings, tables, and 
figures 


Mani and Saravanan have produced a very thorough account of our current understanding of the 
details of pollination in the largest and most complex family of flowering plants. The book begins 
with a brief summary of past and current research on pollination in general and on Compositae in 
particular. This section is particularly well referenced, and is a wonderful resource for anyone inter- 
ested in the topic. An entire section of the book is devoted to an analysis of the groups of insects 
that are commonly found on the flowers in the Composite family, and on their effectiveness as 
pollinators. Special note is made of those insects that, while serving as pollinators, may also do 
considerable damage to the plants as well. There are six chapters devoted to detailed discussion of 
specific parts one compose capitulum, and their development in relation to pollination. One 
chapter presents f the overall structure and multitude of variations found in 
the capitula of this family. This enaptel is ese well illustrated with line drawings, showing 


SIDA 19(2): 426. 2000 


BOOK REVIEWS 427 


detailed pari f the arrangement of florets, and the involucres.A second, particularly well- 
illustrated chapter, discusses the variety of shapes, sizes and colors of the florets, and how this 
diversification can be directly linked with effectiveness of pollination. An entire chapter is also de- 
voted to the wide variety of shapes and sizes of stamens, styles and stigmas. This chapter points 
out the oo of the structural differences of these organs, and how pollinators have influ- 
ences the evolution of those differences. Chapters on nectaries and pappus complete the discus- 
sion of the ie found in composite capitula. Mixed in with the chapters on each of the 
floral organs is a section on sexual polymorphism within the Compositae. In this author's opinion, 
this chapter is a bit misplaced, and would better fit in with the discussion on the overall structure 
of the capitulum. However, other than being misplaced within the overall context of the book, the 
chapter is well written and well referenced. The final three chapters bring all of this information 
together with a well-presented discussion of floral biology, pollination and evolutionary trends 
within the family. This book is very well written and thoroughly researched. It includes a glossary of 
terms which is quite useful, especially for those not familiar with terminology specific to the 
Compositae. The entire volume is well references, with nearly 450 citations contributing signifi- 
cantly to the importance of this work. | would recommend this volume to all students of the 
Compositae, and to anyone who is interested in pollination biology in general.—Debra Trock. 


Avan Hopkins (Editor). 1999. Grass: Its Production and Utilization. Third Edition. (ISBN 
0-632-05017-9, pbk.). Blackwell Science Ltd., 350 Main Street, Malden, MA 02148- 
5018, U.S.A. Distributed for Blackwell Science by lowa State University Press, 2121 
South State Street, Ames, |A 50014-8300, U.S.A. (Orders:800-862-665 7, 515-292-0155; 
www.isupress.edu). $49.95 pbk. xiii + 440 pp., numerous figures. 


It has been a treat to review this book, and here is why: for many years | was a biology professor at 
Kansas State University, which is located in the tallgrass prairie of a notably grassy state. There | was 
familiar with the native grasslands, plus the management and utilization of grasslands for many 
purposes. The book under review centers upon British grasslands, and it offers a different dimen- 
sion to an American's thinking about grasslands 
he text consists of fifteen chapters that are essentially free-standing essays of about 15-30 
pages, each with a several page list of references. Most of the cited papers are from the past de- 
cade. There are chapters on pastures, herbage production, weeds & pests in grasslands, feeding 
values of grass (50 pages), conservation, forage and grazing behavior, landscape and wildlife, etc. 
There is no coverage of grassland systematics. Two chapters drew my attention; one on sward 
establishment and renovation, and the other on amenity grassland. The climate of Britain comfort- 
ably supports lawns of great expanse (swards), and tough turf for athletic or decorative purposes 
(amenity grass), and the approach to lawn and turf management is rather different from what we 
are accustomed to. The introductory chapter is an absolute gem for summarizing the thrusts of 
book for supplying a grass-oriented land-use map of Great Britain. 
user woule was book be ° a Boe palates elon Anyone dealing with the 
lied evant, especially anyone whose studies 
Focus on intensely managed raiesslande The iceratne citations are partieneny valuable, for they 
are largely from the European literature that is less commonly surveyed by Americans.—Theodore 
M. Barkley, Botanical Research Institute of Texas. 


= 


SIDA 19(2): 427, 2000 


428 SIDA 19(2) 


CHERYL HAZELTINE and Barry Lovelace. 1999. The New Central Texas Gardener. (/SBN 0-89096- 
871-3, pbk.). Texas A&M University Press, College Station, TX 77843-4354, U.S.A.(409- 
845-1436, 409-847-8752 fax). $24.95 hbk., $14.95 pbk. 204 pp., 4 pp. color plates, 
b/w illustrations. 

This “basic gardening text for Central Texans,"is a slightly updated version of the 1980 publication 

by Hazeltine and Joan Rival On entitled The pee Texas Gardener.The authors’ stated purpose in 

this new edition is “to pro entral Texas information and 
advice about isieecapng ‘design for function and esthetic oe. wise plant selection; and gar- 
den peak that work.” 

the 1980 book, Hazeltine and Lovelace provide information on plants suited to the 
seni aan. of this area. A new feature of The New Central Texas Gardener is a chapter on 

“Tools of the Trade,” which includes ee aura rie of garden Seal and Aelia 

sources—books, magazines, and internet sites. In addition ardens 

and ponds are briefly described and Te Illustrative material ine lnidies maps, ink vee eee of 
plants by Kate Bergquist, and an 8-page section of color photographs. A month-by-month calen- 
dar for oe concludes the boo 

hough advertised for’ amate eur and seasoned gardeners,” the latter will find it somewhat 
aes |, while those familiar with the earlier edition ta with recent central Texas gardening 
literature will discover little new information.—Joan H. Swai 


Rogert W. Kicer, CHARLOTTE A. TANCIN, and Gavin D.R. Brinson. 1999. Index to Scientific Names 
of Organisms Cited in the Linnaean Dissertations Together With a Synoptic 
Bibliography of the Dissertations and a Concordance for Selected Editions. 
(ISBN 0-913196-67-3, hbk.). Hunt Institute for Botanical Documentation, oy 
Mellon ea Pittsburgh, PA 15213-3890, U.S.A. Price unknown. 300 pp. 6 b/w 


figures, 8" 
Carl Linnaeus, the 18th Century grandsire of systematics, saw 186 dissertati prepared under his 
direction at the University of Uppsala. They represent a trove of Hua some of which was 
undoubtedly prepared by Linnaeus himself. The original dissertations were printed and circulated 


at the time of their defense, but were later Collected: ne pu alene in en editions, two of 
which were edited by Linnaeus under the title Am 

s book indexes all of the formal (i.e., eaenthc ‘) names used int the 
and oe where each name is found in — original dissertation and in . collected editions. 
There is a bibliographic catalog of the origin plus a succinct overview of the several 
collected editions. A concordance is anc to tie original dissertations to their locations in the 
collected editions. 

sity, the introductory essay is brief, but clear and to the point. The authors use con- 
ventional bibliographical styles and the work is easy to navigate. Were | to quibble, | would wish for 
an eae of the Lidén sae which are used to catalog the dissertations. 

e dissertations retain some currency in contemporary biology because many are signifi- 
cant in aie poe aic) priority, and because the texts of all represent the thinking and 
attitudes of a group of capable people who were among the earliest to consider the flora and 
fauna from a world perspective. 

Biologists of every stripe always have been addicted to scribbling, and their products crowd 
the libraries, often to be slowly forgotten. But, a fact is a fact regardless of when it was observed, 
and the writings of the past are not merely space-fillers. Those among us who have the skill and 
facilities to make the great wealth of old information comfortably accessible to the rest of us de- 
serve our anens we Lig wishes. May they continue their work.—Theodore M. Barkley, Botanical 

bh] 


Recenn 


tati 


SIDA 19(2): 428. 2000 


BOOK NOTICES 


Marie CAILLeT, J. FARRON CAMPBELL, KEVIN C. VAUGHN, and Dennis VERCHER (editors). 2000. The Loui- 
siana Iris; the taming of a native American wildflower. Second Edition. (ISBN 0- 
88192-477-6, hbk.). Timber Press. Inc., The Haseltine Building, 133 S.W. Second Av- 
enue, Suite 450, seals OR 97204-3527, U.S.A. (503-227-2878, 503-227-3070 fax; 

m). $34.95 hbk. 254 pp., 111 color photos, 5 watercolors, 11 
ee photos, 14 ine ae 1 tables 

The beardless Louisiana irises (subsection Apogon) belong to series Hexagonae of the genus /ris in 
the family lridaceae. The greatest concentration of species is in the State of Louisiana, hence the 
name Louisiana irises. They comprise a unique group in the iris family and perhaps are the most 
distinctive in color and in form. I've seen some in cultivation and | can attest to that fact. The main 
objective of tt people all over the Id in growing Louisiana irises. The ealtals 
have given growers an incredible amount of information about Louisiana irises and h mph 

the how-to-aspects of adaptability, propagation, culture, and hybridizing. These wonderful irises 

will grow with moderate to excellent results in most of the United States and in many foreign 

counties.’In a very short amount of time, the Louisiana iris had made its way into the hearts of both 

Galeeniel ang almiatelt Hy eIaZeKs Members of the Society for Louisiana irises and the current edi- 

tors are 1 for having done a terrific job in promoting the beautiful Louisiana Irises. 

Conten ts—1) History of Louisiana Irises; 2) Classification and Species; 3) Description of the 

Louisiana Iris; 4) Collecting of the Species and Natural Hybrids; 5) Propagation of Louisiana Irises; 6) 

History of Hybridizing to 1988; 7) History of Hybridizing since 1988; 8) Tetraploid Hybridizing; 9) 

Objectives in Hybridizing; 10) Culture; 11) Landscaping with Louisiana Irises; 12) Flower Arranging 

with Louisiana Irises; Appendix A, Society for Louisiana Irises; Appendix B, Popular Cultivars of Loui- 

siana Irises; Appendix C, Mary Swords DeBaillon Medal; Glossary; References; and Index of Plant 

Names.—Barney Lipscomb. 


f th Cc th hi Mountains (ISBN 


STANLEY L. BENTLEY. 2000. Native Orchids o 

78-2563-8, Abie 0- 8078-4872-7,pbk.). Univ. of North Carolina Press, PO. Box 2288, 

Grave Hill, NC 27515-2288, U.S.A. (800-848-6225, 919-966-3829 fax; www.uncpress. 
unc.edu). $39.95 hbk; $24.95 pbk. xviii + 256 pp., 119 color photographs. 


Another orchid book. Well, maybe not! This is quite a guide filled with great photography of 52 
species of orchids found in the mountains of southern Appalachians. That is to say, a region en- 
compassing western Virginia and North Carolina and eastern West Virginia, Kentucky, and Tennes- 
see. This book is quite a treat for you amateur naturalists or professional botanists who happen to 
live, vacation or botanize in the southern Appalachians. Each species is provided with a scientific 
name and common name(s), a description of the flower (color, shape, and size), information on 
flowering time, distribution and typical habitat. The author offers much advice when hunting for 
orchids, but one piece of advice | thought was most interesting was to carry binoculars. Have you 
ever used binoculars to look for orchids? Well, you just might want to consider it according to the 
author.’Time after time, when | have spotted a plant high on a bank, well past the ‘telling for sure’ 
point, binoculars have saved me from having to scramble up the bank to try and determine its 
species. As the years go by,| appreciate more than ever the steps saved by my trusty binoculars.” 
ents.—Preface, Acknowledgments, Please do not Dig Native Orchids, Introduction, What 
is an orchid?, Rarity among Orchids, Looking for Wild Orchids, Special Orchid Places in the Southern 
Appalachians, Preserving Wild Orchids and Their Habitat, Using This Book, List of Native Orchid 
Genera of the Southern Appalachians, Glossary, Bibliography, and Index. —Barney Lipscomb. 


SIDA 19(2): 429. 2000 


430 SIDA 19(2) 


Teo L. Petit and Joxn P, Peat. 2000. The color Encyclopedia of Daylilies. (ISBN 0-88192- 
488-1,hbk.). Timber Press. Inc., The Haseltine Building, 133 S.W.Second Avenue, Suite 
450, Portland, OR 97204-3527, U.S.A. (503-227-2878, 503-227-3070 fax; 
www.timberpress.com). $49.95 hbk. 8 1/2" x 11", 296 pp., 1300 color photos. 


The title of this book is most descriptive and just about says it all. There is plenty of color and it is 
encyclopedic. What an undertaking and what a beautiful book the authors have given us. There 
were certainly plenty of daylilies to choose from.The authors point out that it was a daunting task 
to select which daylilies to include out of the 40,000 plus daylilies registered at completion of the 
manuscript.“Ted Petit and John Peat's simple and concise text on the history, botany, and horticul- 
ture of daylilies along with their collection of eee eSoEt One rom Suiicaas sel 
the world combine to create the most yl No 
other aon so completely catalogs the world of daylilies.” 

Contents.—Foreword; Preface and Acknowledgments; 1) The Daylily Plant; 2) History of the 
Daylily; ni nee of Daylily eng 4) Singles; 5) Eyed and Patterned Daylilies; 6) Doubles; 7) 
Spiders, Variants, and Unusual Forms; 8) Polytepal Daylilies; 9) Small and Miniature Daylilies; 10) 
Hybridizing; 11) Cultivation; 12) Daylilies of the Future; Appendi A, Des erie srier 
dix B, Sources for Daylilies; Bibliography; and Index of Picured Cult 


Peter GOLDBLATT and JOHN Manninc. 2000. Wildflowers of the Fairest Cape. (ISBN 0-620- 
24787-8, pbk.).Red Roof Design cc, Cape Town, SOUTH AFRICA. Available from Tim- 
ber Press. Inc., The Haseltine Building, 133 S.W. Second Avenue, Suite 450, Portland, 
OR 97204-3527,U.S.A.(503-227-2878,503-227-3070 fax; www.timberpress.com).$ 34.95 
hbk.,8 1/4" x 11",315 pp., 663 color photos, 1 color map 


"OH Lord, how manifold are thy works! in wisdom hast thou made them all: the earth is full of thy 
riches."—David, Psalm 104:24. And the fairest cape (Cape of Good Hope) has its share of botanical 
riches! No wonder it is referred to as one of the botanically richest areas of the earth.| don’t think 
anyone will argue that point. The two eminent botanists, Goldblatt (Missouri Botancial Garden, St. 
Louis) and Manning (National Botanical Institute, South Africa) detail 652 of the most common or 
conspicuous wildflowers to be found there. Many of the botanical wonders there are presently in 
cultivation. One visit to South Africa or one look at this book and you too will gasp at the beauty. 
Well done thou good and faithful servants! 
Contents.—Preface; The Cape Floral Kingdom; Introduction; Climate and Rainfall; Fynbos and 
Renosterveld; Wildflowers Routes: The Cape Peninsula, The West Coast, The Bokkeveld and Hantam, 
The Olifants River Valley and the Lambert's Bay Coast, The Mountain Passes, The South Coast; How 
to use this Book; Wildflower Schedule; MINETONEE Route Map Quick Guide to the Families; Illustra- 
tions; Descriptions; Glossary; and Index.—Barney Lipscomb. 


SIDA 19(2): 430. 2000 


BOOK NOTICES 431 


Bit Matcotm and Nancy Matcoim. 2000. Mosses and Other Bryophytes. An Illustrated 
Glossary. (ISBN 0-473-06730-7,hbk.).Micro-Optics Press, Box 320, Nelson, NEW ZEALAND. 
Available from Timber Press. Inc., The Haseltine Building, 133 S.W. Second Avenue, 
Suite 450, Portland, OR 97204-3527, U.S.A. (503-227-2878, 503-227-3070 fax; 
www.timberpress.com). $39.95 hbk.,5 7/8" x 8 1/4", 220 pp., 968 color photos, 22 
line drawings. 


This color glossary covers mosses, liverworts, and hornworts. According to the authors,"The three 
groups were chosen because even though they're not closely related, their structure and life cycles 
are similar enough that a term which applies to one often applies to the other two as well.” Nearly 

1000 photos—representing nearly 400 species—were taken at various levels of magnification, 
Hes wien a microscope. A oes bar is - Provided wine aoe prioto caption. The authors have in- 
g the same or similar mean- 
ings, and cross-referenced terms used elsewhere in n the glossary. Terns) in the glossary are printed 
in boldface. Example: mucronate — ending abruptly in a short point (mucro) (compare with 
apiculate, ending in a point somewhat longer than a mucro, and cuspidate, ending in an even 
longer point that's also stouter). More than one illustration is provided for terms that describe highly 
variable structures such as the spines and papillae of leaves, or brood bodies and leaf cell lumina. 
The glossary would definitely be useful to students and teachers as there are several illustrations of 
bryophytes that often are studied in the laboratory. 

ntents.—Introduction; Mosses and Other Bryophytes an Illustrated Glossary; Further Read- 

ing; and Illustration Index.—Barney Lipscomb. 


W. ArtHur WHisTLeR. 2000. Tropical Ornamentals: A Guide. (ISBN 0-88192-475-X, pbk.; 0- 
88192-448-2, hbk.). Timber Press. Inc., The Haseltine Building, 133 S.W. Second Av- 
enue. suite 450. Portland, OR 97204-3527, U.S.A. (503-227-2878, 503-227-3070 fax; 

com). $34.95 pbk., $59.95 hbk. 542 pp. 458 color photos, 50 line 


ane a xo. 

There is no doubt this book is about mostly tropical ornamentals and a few others. Many of the 
n tropical ornamentals are found in cultivation throughout the world and they are 

pesueale: tee here. There are some oe covered in me book Male are Hot MOpICah put 


they are cultivated in tropical areas Nardi lla, Heli } Is 
Solidago nemoralis, etc.). Ua 400 plus plants treated | in this book d in alphabetical order 
by scientific name and not common name. You can find the common name in the index. This of 


course allows closely ae plants to be placed together. About two-thirds of the plants are in 20 
plant families and are accounted for in an appendix, Twenty Common Plant Families. Each plant is 


ing characteristics; and Description. It really is a guide to the plants most frequently encoun- 
tered in gardens of the tropics. The photos are splendid. This is an excellent reference on tropical 
plants in cultivation. 

nts.—Preface, Introduction, Organization of the Information in Tropical Ornamentals, The 
Tropical Ornamental Plants, Wwenty Common Plant Families, Identification Key, Glossary, and In- 


dq D Pee L 
UDA. DUTTICY CIPOCUTTIU, 


SIDA 19(2): 431. 2000 


432 SIDA 19(2) 


CHRISTOPHER LLoyD. 2000. Christopher Lloyd’s Garden Flowers: Perennials, Bulbs, Grasses, 
Ferns. (ISBN 0-88192-492-X, hbk.;0-88 192-448-2, hbk.). Timber Press.Inc., The Haseltine 
Building, 133 S.W.Second Avenue. Suite 450. eee OR 97204-3527, U.S.A. (503- 
227-2878, 503-227-3070 f. m).$39.95 pbk.CAN 54.95 hbk.448 
pp.69 color ee 50 line ao 7] 1/2" x 10" 


This is aone man band g from Acanthus to Zigadenus. Interestingly enough, the 
author says,All the “nee included ave come within my experience during a long life of garden- 
ing, and most | have grown myself at some time.” So, | guess if you don't find your favorite plant in 
the reference, then | suppose it is one not grown by the author; surely that will include a lot of 
plants. However, if you are lucky and consult just one of the plants in this book you will perhaps get 
everything the author has learnt, thought, seen, tried, likes, or regrets about the plant, and that 
could be your lucky day! It coon is one man’s perspective, but what a perspective considering it 
is stale Lloyd's lifetime st 
Contents.—Introduction, ae [ca.372], Glossary, Sources of Photographs,and Index.—Barney 


ee 


Rosert H. MoHLENBROCK. 1999. The Illustrated Flora of Illinois: Sed Carex. (ISBN 0-8093- 
2074-6, hbk.). Southern Illinois University Press, RO. Box 3697, Carbondale, IL 62902- 
3697,US.A.(61 18-453-6633,618-453-1221 fax; danseit@siu.edu). $59.95 hbk. 328 pp. 
159 line drawings, 159 maps 


Wy 4 fC] rani q i | 
1a 


Wow! This is volume 14 of th sixth and last volume devoted to 
monocots. The 159 detailed ilnstrations By Paul Nelson are excellent and will aid any person in the 
identification of these species. A key to all the species is included. The usual stuff is included with 
each species: Genus species, place of publication, synonymy, description, common name, habitat, 
range, and Illinois distribution ~ map of Illinois); this is followed by a discussion of the nomen- 
oT and habitat of the speci 

m the dustjacket.—" ee more than three-fourths of the species of Carex in Illinois are 
ae of wetlands, an understanding of the genus is critical for those working in wetlands. 
Amateur ane pole geal pans well find the information extremely valuable, as will environ- 


mental and arden clubs, farm bureaus, home extension groups, scout orga- 
nizations, and aches! fibres Persons bear in natural areas programs and in rare and endan- 
gered species programs and those working on environmental impact assessments and wildlife 


R 


management projects will also and the information pertinent.’—Barney Lipscomb. 


S.T.Runket and D.M. Roosa. 1999.Wildflowers and Other Plants of lowa Wetlands. (ISBN 
0-8138-2174-6, pbk.).lowa State University Press, 2121 South State Avenue, Ames, 
IA 50014, U.S.A. (Orders: 1-800-862-6657; www.isupress.edu).$24.95 pbk.372 pp.Color 
photos. 


From the Preface-—‘With the increasing awareness of the importance of wetlands, a book was 
needed to help the non-specialist understand how wetlands are classified and to aid in the identi- 
fication of wetland plants. There are wonderful taxonomic books available, however, they are often 
so technical that all but professional taxonomists become discouraged. These manuals are often 
regional in scope and contain many plants that do not occur in the local area. This tends to frus- 


SIDA 19(2): 432. 2000 


BOOK NOTICES 433 


trate many users, particularly beginners.We think wetlands are grand places.We want you to enjoy 
them as we do. Hence, we offer this book as a starting place for those who wish to learn more 
about lowa’s wetlands and wetland plants. Plant descriptions are presented by habitat (terrestrial 
or aquatic), then refined by habit (e. 7 floating or eae or oy taxonomic giouR (e. g., ferns 
and allies or trees and shrubs). Common names vary throughou we have included 
those in frequent use, although other y be used regionally Sent We have also included the 
plant's Latin name, along with the a 


Ronatb J. Tayior. 1998. Desert Wildflowers of North America. (ISBN 0-87842-376-1, pbk.). 
untain Press Publishing Company, P.O. Box 2399, Missoula, MT 59806, U.S.A. (406- 
728-1 anced pbk. 349 pp.500 + color photos, line drawings, maps. 


From the back cover.—"Each spring, when rainfall and temperatures combine in just the right way, 
hundreds af wildfiowerd transform the deserts of North America into a sea of vibrant color. With 
descriptions and photographs of more than 500 species of flowering plants, this full-color filed 
guide leads casual visitors and certified desert rats alike through the flora of the blooming desert. 
The book includes an overview of desert ecology, a simplified botanical key, and an illistatee 
glossary that will help even novice wildflower fans denny dest eae in eonlige ence.” 
Contents.—Preface; Introduction; Desert Wildfl American Desert 
Wildflowers to Family; Plant Anatomy Illustrations; Glossary [illustrated]; Selected References; and 
Index. Desert Wildflowers of North America covers the Mojave, Sonoran, Painted, Chihuahuan, and 
Great Basin deserts. This book was planned and written for the amateur botanist and anyone who 
appreciates wildflowers, particularly in desert environments. The plants are arranged alphabeti- 
cally by family, that is the common name of the family, i.e. Acanthus Family, Allthorn Family, Ama- 
ranth Family, Barberry Family, etc. The book does n not cover or list plants on a desert-by-desert basis, 
eee times, nor does it treat the M t of the Sonoran and Chihuahuan 
erts. Because there is overlap in the desert flora of the United States and Mexico, it certainly 


i 


sear be useful in Mexico.—Barney Lipscomb. 


£, i +h VA 


JEAN ANorEWS. 1999. The Pepper Trail: History World. (ISBN 
1-57441-070-9, hbk.). Dnivceny of North Texas ‘Press, P.O. Box 311336, Denton, TX 
76203-1336,U.S.A. (Orders: 800-826-891 1,940-565-4590 fax).$50.00 hbk. 264 pp.57 
color illustrations, 7 maps, 7" x 12" 


From the dustjacket.—"Veteran afficionados and newcomers on the pepper trail will enjoy this stun 
ning addition to Jean Andrew's classic work on peppers. Andrews, who has been called ‘the first 
lady of chili pepe “the Eg DemOWNers of ue ent world,’ as well as her own registered trademark 
‘The Pepper Lady,’f movements of capsicums along the spice roads, 
through much of turkey and the Middle East, Ce and Monsoon Asia (India, Nepal, Bhutan, Sri 
lanka, Thailand, and mIclonIeS)a) plus the Senden and Hunan piovinee in China and the Silk Route. 
This latest offering of And h incl | facts, such as the etymol- 
ogy of the word ‘cayenne.” 

“The first spice to be used by may, peppers are currently ‘hot’ in Mexico, Guatemala, much of 
the Caribbean, most of Africa, parts of South America, India, Bhutan, Malyasia, the United States— 
Louisiana, Texas, and the Southwest—plus Korea.” 


SIDA 19(2): 433. 2000 


434 SIDA 19(2) 


“A chapter on what makes a pepper a pepper includes detailed descriptions and illustrations 
of twenty-seven separate varieties of the capsicum, as well as miscellaneous cultivars and detailed 
directions on she with fresh and dried peppers, including how to choose and use them and 
how to ca acl em.” 

Con aise Part I, The Pepper: How Our Food Got Hot; Part Il, Preparation & Reci- 
pes; ee epee, Subject Index; and Recipe Index. 

Here is everything you every wanted to know about beds and Maybe then some! Check 
out the recipes! | stair cae Snr al but many ale listed for the followin g dishes: Soups & Salads; 
Meat, Fowl, and Seaft ; Desserts; and preseives and Condiments. | just 
can't Hiscine ou Pappet Jelly, especially when | was raised on Wild Plum Jelly, but who knows? 


Barney L ips 


Contey K.McMuiten. 1999. Flowering Plants of the Galapagos. (/SBN 0-8014-8621-1, pbk.; 
0-8914-3710-5,hbk.).Cornell University Press, Sage House, 512 E. State Street, Ithaca, 
NY 14850, U.S.A. (Orders: 800-826-891 1, 940-565-4590 fax). $59.95 hbk., $29.95 pbk. 
370 pp. 266 color photos, 1 map, 6" x 9" 


From the back cover—“The Galapagos are home to a wide-ranging assortment of unusual plants 
and animals. The islands became famous as the site of Charles Darwin's research leading to his 
theory of evolution by natural selection, and their magnificent flora and fauna continue to draw 
visitors from around the world. Based on the author's sixteen years of fieldwork and featuring his 
exceptional photography, Flowering st of the Galdpagos is the first accessible and in-depth, yet 
compact, guide to the gia life of the 
itents.—Foreword; Preface; hele eee Galapagos Islands Map; How to Use This 
Guide, Illustrated Plant Terms; A Brief Introduction to the Galapagos Islands; Plant Descriptions, Plant 
Key eee Literature Cited; Appendix 1:List of Galapagos Islands Flowering Plants Treated in the 
2: Selected Visitor Sites and Flowering Plants Likely to Be Encountered; and Index 
“Chey than just flipping pages through the nice photographs to identify your plant inthaad i 
one must first determine and select the appropri rowth form. The 390 species are arranged in 
the book in one of five groups, i.e. tree shrub herb vine, and cactus. Each of the five groups is then 
divided into groups based on how a plant's leaves are typically arranged on its stems. Options 
include alternate, opposite, WAGHED, clusteted and Baral Next, each of these categories is subdi- 


ided based on flower color. The choices are whit w or orange, pink, red or purple, blue, green 
and brown. Once you have made your selection, ‘there is a page number that takes you to the 
appropriate subsection."Once tion is located, the reader simply has to thumb 


through the photographs until he ¢ or she finds one that matches the specimen in question.”— 
Barney Lipscomb. 


SIDA 19(2): 434. 2000 


BOOK NOTICES 435 


Witiam A.Weser (Editor).2000. The American Cockerell: A Naturalist’s Life, 1866-1948. 
(ISBN 0-87081-544-X, hbk.). University Press of Colorado, 5589 Arapahoe Avenue, 
Suite 206C, Boulder, CO 30303, U.S.A. Distributed by: University of Oklahoma Press, 
Book Distribution Center, 4100 28" Ave., N.W. Norman, OK 73069-8218, U.S.A. (Or- 
ders:800-627-7377, 800-735-0476 fax).$29.95 hbk.352 pp.7 b/w photos, line drawings. 


From the dustjacket.—"In The American Cockerell: A Naturalist’s Life, 1866-1948, botanist William A. 
Weber pulls together pieces of the life of T.D.A.'Theo’ Cockerell, a man who was an internationally 
known scientist, a prolific writer, and a highly regarded teacher at the University of Colorado in 
Boulder. The elder brother of the noted scholar Sir Sydney Cockerell, Theo labored in relative ob- 
scurity in America while his brothers and their families were basking in the limelight of smart Brit- 
ish society.” 

"Despite his alienation from his elite background, he nevertheless became a great teacher; a 
mentor,a kindly artist and writer of rhymes for children, and the greatest specialist on bees in the 
world. His contribution to the understanding of wild bees is monumental—he catalogued over 
900 species in Colorado alone, and he assiduously collected them wherever he traveled. By 1938 
he had published the names and descriptions of 5,480 new species and subspecies. Despite his 
accomplishments in entomology, however, T.D.A. Cockerell resisted specialization. He was also an 
early supporter of women’s rights, a Morrisian socialist, an avid reader; an auuiol of almost 4,000 
fate scientific papers, book reviewers, and discussion of social issues.” 

ced together from T.D.A‘'s little known autobiographical writings, The American Cockerell 
dene: this extraordinary individual's tremendous breadth of interest, competence, and 
talent.” 

Contents.—Foreword by Norma LeVeque; Joseph A. Ewan on Cockerell; Acknowledgments; 
Cockerell Chronology; Autobiographical Papers; Philosophical Papers; Academic Matters; Miscella- 
neous; Life and Habits of Bumblebees; Postcript; Biographies, Obituaries, and Notices; Endnotes; 
and Index 


WH. Fisher. 2000. Rain Forest Exchanges: Industry and Community on an Amazonian 
Frontier. (ISBN 1-56098-983-1, pbk.). Smithsonian Institution Press,470 L’Enfant Plaza, 
Suite 7100, Washington, D.C. 20560-0950, U.S.A. (Orders: PO Box 960, Herndon, Va 
20172-0960; 1-800-782-4612).$19.95 pbk. 222 pp. 11 figures, 4 maps. 


From the back cover—Drawing on both historical sources and indigenous informants, William H. 
Fisher argues that decisions to cooperate with frontier industries are best understood by taking 
into account the power of native social systems to shape the acquisition oF trade goods. Charting 
the history, politics, economics, and ecology of the regions, he tells how subsistence practices such 
as hunting and gardening have been altered be sedentarization, fia villagers interact with In- 
dian-agency and extractive-firm personnel,and how notions of barter or sale only loosely describe 
is aNeie of goods that take pace in the yee In Rain Forest Exchanges, Fisher contends that 


eo] 
| 


remain probl ic unless, in addition to she tamance 
eel and economic organization are recognized.—Review forthcoming in Sida 1 9(3), 2001. 


Tagg 1 yt , +h rt 


SIDA 19(2): 435. 2000 


436 SIDA 19(2) 


W.K. CHAPMAN, V.A, CHAPMAN, A.E., Bessette, A.R. Bessette, and D.R. Pens. 1998. Wildflowers of 
New York in Color. (ISBN 0-8156-0470-X, pbk). Syracuse a Press, Syracuse, 
NY 13244-5150, U.S.A. $24.95 pbk. 164 pp. 350 + color phot 


From the back cover.—Wildflowers of New York in Color is a field guide that will give nature enthusi- 
asts instant access to the diverse and beautiful flora of the state. It contains over 350 photographs 
and concise descriptions, written in easy-to-follow, nontechnical language. The color illustrations 
have been selected for their scientific accuracy as well as their aesthetic quality. This field guide is 
keyed in a manner that easily leads the reader to major groups based on flower color and other 
physical characteristics. Wildflowers of New York in Color contains descriptions of both commonly 
encountered and rarer, protected species. Included are color illustrations and descriptions of spe- 
cies not seen in other field guides. Nomenclature has been uCaaieS to reflect current usage. This 
book will be an indispensable companion for nature | y interested in the wildflow- 
ers of New Yor 


D.J.Raynat and D.J.Leoroto. 1999. Landowner’s Guide to State-Protected Plants of For- 
ests in New York. (ISBN 0-9670681-0-X, pbk). State University of New York, College 
of Environmental Science and Forestry, Syracuse, NY,U.S.A.$19.95 pbk. 92 pp. Color 

photos 


From the Introduction.—The purpose of this b he key diagnostic characteristics of 
the protected plants that are found in forests of New York ae using Puptogab isa ane ee non- 

technical descriptions of the plants and their Joes A ates! vallieny of plant 
in New York State. Of ') include salt and 
freshwater marshes, bogs, shrub thickets, and {bine meadows, Closed eon or fotos! communi- 
ties include southern oak-pine, oak hickory, northern hardwoods, elm-ash- Mes swamp, and bo- 
real spruce-fir forests. Some of these species also occur in open canopy communities, especially 
during early stages of succession. Following information about the cenitiesdon and ecology of 


rare. These species are legally regarded as “exploitably vulnerable,” because of their beauty and 
tendency to be collected, they are protected. All of our native orchids, nearly all of the ferns, many 
species in the lily family, and other showy native species are included in this category; some of 
these species also are very rare 


Dennis W. Macee and Harry E. AHLES. 1999. Flora of the Northeast: A Manual of the Vas- 
cular Flora of NewE York. (ISBN 1-55849-189-9, hbk.). 
The a of Massachusetts Press, Box 429, Amherst, MA 01044, U.S.A. $69.95 
hbk. 1214 pp. 995 line drawings, 2433 range maps, 6" x 9", 


From the dustjacket.—“Designed as both a reference work and year-round field manual, this vol- 
ume contains more than 2,400 range maps and 995 line drawings. For botanists, naturalists, and 
students interested in an up-to-date treatment of the vascular flora of greater New England, it will 
be an invaluable resource. 
geographic scope of the work extends from the Canadian border south through Long 
Island and west to the Hudson River. The “General Keys" section contains fourteen keys that include 
such groups as aquatic plants, vines, and woody plants in winter condition. For both woody and 
herbaceous families, the keys cover flowering as well as fruiting condition 
The “Descriptive Flora” section includes keys to all of the genera and species, descriptions of 


SIDA 19(2): 436. 2000 


BOOK NOTICES 437 


the families and genera, and accounts of the individual species. The latter incorporate information 
n wetland site index, rare status, wildlife food value, food and medicinal value for humans, and 
poisonous or hallucinogenic properties. 
distributions of more than 2,400 species are presented on range maps, and book in- 
cludes line drawings of 995 species, showing diagnostic features designed to clarify descriptions 
used in the keys. Near the end of the volume there are two matrices of diagnostic characteristics, 
one for dicots and one for woody plants in winter condition. These matrices can be used to identify 
a specimen displaying a Information, or to find examples of given vegetative, flower, or fruit 
ae rao for use in tea 
e line drawings sense not for every species, and the range maps, are nice features of 
the i In some cases not all of the couplets in the keys are lettered. The latter branches of lead 
couplets—in many of the keys—are not lettered. That doesn’t make the key wrong, but sometimes 
itis a little confusing—Barney Lipscomb. 


BOOK Sree NOTICES 


Rick Darke, 2000. The Color E fO tal Grasses on CD-ROM. (ISBN 0- 
88192-479-2,PC eat Windows 95 or higher Timber Press. Inc., The Haseltine 
Building, 133 S.W. Second Avenue. Suite 450. Portland, OR 97204-3527, U.S.A. (503- 


~— 


227-2878, 503-227-3070 om).$59.95 CD-ROM.721 color photos. 
This is the CD-ROM version ss Dales s hard copys ference Color Encyc aa CO Sede ce 
In addition to grasses, the grass-like cluded on the CD, i.e. sedges, rushes, restios, an 


cat-tails; selected os are included as well. a = version i cee Git os new plants and 
200+ color photos not found in the hardcover.The photos from the CD can be printed in full color 
for use in design work, plant identification, and nursery signage. There is an Interactive USDA Zone 
Map, a useful “Search” function that allows you to track down plants and plant names, as well as 
other useful features.—Barney Lipscomb. 


David - ZAHLER and Eowarb C. Jensen. 1999. Conifers of the Pacific Northwest [on CD- 

OM.].Oregon State University College of Forestry, Forestry Media Center, 248 Peavy 

si Corvallis, OR 97331-5702,U.S.A.(541-737-4702, 541-737-3759 fax;http://osu.orst.edu/ 
Dept/fmc; forestrm@ccmail.orst.edu). Price unknown. CD-ROM. 


"The diverse topography and climate of the Pacific Northwest combine to create t variety of habi- 
tats in which conifers flourish. Nearly 30 species of conifers grow naturally here, organized into 4 
families and 13 genera. Many other species of conifers have been introduced from other parts of 
the world and thrive here with minimal care. 

The purpose of this program is to take the mystery out of identifying our most important 
native conifers—and to share a bit of their ecology and natural history. Especially in the larger 
genera it will help immensely if you first learn to distinguish one genus from another, and then 
focus your attention on the different as that occur ee each genus. 

IBM Compatible PCs; 486 Dx, SX or greater; Wind 5, Windows NT 3.51, or later; 8 MB of 
available RAM; 8- or 16-bit sound card. chron All ae with 8 MB of available RAM and 
sound ca meaeee 

educational product won a Bronze Award in the Information ee category at the 
1999 Nara Communicators in Education Critique & Awards Prog 


SIDA 19(2): 437. 2000 


438 SIDA 19(2) 


BOOKS RECEIVED 


Daniet B.Waro and Rosert T. Inc. 1997. Big Trees. The Florida Register. (ISBN 1-885258-06- 
2, pbk.). Florida Native Plant Society. $20.00 oe viii + 223 pp. 


L Deak. Vin fal 


From the Preface.—"This Florida Register doc ti d 243 non-native trees, for a total 
of 853 listed trees. For each individual tree, information is given as to its common and scientific 
name, its Measurements in terms of circumference of trunk, height, and spread of canopy, its loca- 
tion, owner, and nominator, and its rank as a champion or other status among the elite trees of 
Florida. Of these documented trees, 151 of the native species and 13 of the non-natives are Na- 
tional champions, while ie of men falives and 118 of the non-natives are Florida champions 

ontents.—Prefa yn; Inventory of Big Trees; Giants of Yesteryear; How 
They Died; Rates eprom: Largest and Sifiallest ie Ranked by Height; The “Average” Cham- 
pion, Large Vines; County Tree-Find List; Nominators and Owners; Acknowledgments; Selected Ref- 
erences; Appendices: A. What Is a Tree? B. Mechanics of the Survey. C. Memorandum of Under- 
standing. D, How to Nominate a Champion Tree. E. How to Measure a Champion Tree. F. Evaluation 
of the Formula. G. A Case of Volume Measurement; H. William Bartram’s Big Trees; and Index to 
Scientific and Common Names 


David E. Brown, FRANK REICHENBACHER, and SusAN E. FRANSON. 1998. A Classification of North 
erican Biotic Communities. (ISBN-87480-562-7, pbk.). University of Utah Press, 
1795 E. South Campus Drive, Suite 101, Salt Lake City, UT 84112-9402, U.S.A. (801- 
585-9786, 801-581-3365 fax). $19.95 text; $20.00 map; book and map set, $34.95. 

152 pp. 118 b/w figures (photos, maps), 8" x 12" 


From the back r—A Classification of North American Biotic Communities describes a para 
classification ster for biotic communities from the Arctic circle to Panama. It inelees? 
tographs, each illustrating an example of a specific community. Intended as a first 
ing a universal standard for inventorying habitats, this system uses existing classifications of North 
pene e ee environments. 
orth American Biotic Communities” =e provides a dramatic and useful illustration of 

the oe communities delineated in the volum 

ntents.—List of Figures; List of Tables; noweaaene Introduction; ade cee le 
cal ena 2) The Classification System; 3) The Biotic Communities of North p; Plates; 
Literature Cited; and Literature Consulted 


tahlich 


TERESA CABRERA CacHON. 1999. Orquideas de Chiapas. (ISBN 968-5025-44-4, hbk.). Consejo 
Estatal para la Cultura y las Artes de Chiapas, Polyforum Mesoamericano, Calzada 
Andrés Serrra Rojas s/n.,Tuxtla Gutiérez, Chiapas, CP 29040, MEXICO. Approx. $74.00 
hbk. 196 pp. Color photos, 9" x 12". 


A+ ye | ae op che D + A | | 


Content nts. —P ICO I itacion; i Prefac | , Par ioral 
de la flor; Partes de la columna; Clasificacién; ane a las especies; eeadennas de formas s; Mapa de 
Regiones Fisiograficas del Estado; Glosario; Bibliografia; Lista de Especies de la familia Orchidaceae 
para Chipas; and Indice de nombres cientificos. 


SIDA 19(2): 438. 2000 


BOOKS RECEIVED 439 


Daphne Gail Fautin, Douctas J. Furuyma, and Frances C. James (editors). 1999. Annual Review 
of Ecology and Systematics. Volume 30. (ISBN 0-8243-1430-1, hbk.; ISSN 0066- 
4167).Annual Reviews Inc.,4139 El Camino Way, PO. Box 10139, Palo Alto, CA 94303- 
0139,U.S.A.(650-493-4400, 800-523-8635,650-424-0910 fax;www.AnnualReviews.org). 
$120.00 hbk. 641 pp., b/w illustrations and one color plate. 


Annual Review of Ecology and Systematics for 1999 is packed once again with a mix of articles on 

ecology and systematics. Volume 30 has a total of 21 articles followed by a Subject Index, Cumula- 

tive a of Contributing Authors, and a Cumulative Index of Chapter Titles, Volumes 26-30. 

Content 

The ae mand Early Evolution of Turtles 

Uses of Evolutionary Theory in the Human Genome BORG 

Streams in Mediterranean Climate Regions:A influences and Biotic Responses to Predictable 
Seasonal Events 

Choosing the Appropriate Scale of Reserves for Conservation 

Conspecific Sperm and Pollen Precedence and ao 

Global Amphibian Declines: A Problem in Applied Ec 

Using Phylogenetic Approaches for the Analysis of a Breeding System Evolutio 

Evolution of ee rsity in Warning Color and Mimicry: Polymorphisms, shifting: Balance, and 


peciatio 

ee atee of wane with Bacterial Symbionts: Insights From the Squid-Vibrio Associations 

The Relationship Between Productivity and Species Richness 

ee of ee on ae e Polymorphisms 

Polymorphisms in Systematic ae Comparative Biolo 

eRe Boog Coupling in Streams: The Pervasive ices of Flow on al a 

Astrobio xploring the Origins, Evolution, and Distribution of Life in the U 

Evolution es . Eastern Asian and Eastern North American Disjunct Dk baton: in Flowering 

Plan 

Full of Sound and Fury: The Recent History of Ancient DNA 

Do Plant Populations Purge Their Genetic Load? Effects of Population Size and Mating History on 
Inbreeding 

Historical Extinctions in the $ 

Gene Flow and Introgression oi Domesticated Plants into Their Wild Relatives 

Resistance of Hybrid Plants and sues to Herbivores, Pathogens, and Parasites 

Evolutionary Computation: An Overv 


BARBARA = sane 2000. The Berry Grower’s Companion. (|SBN 0-881 92-489-X, hbk). Timber 
s.Inc., The Haseltine Building, 133 S.W.Second Avenue. Suite 450. ay = 
97 904- 3527,U.S.A.(503-227-2878,503-227-3070 f 
hbk. 308 pp. 40 color photos, 25 b/w illustrations 17 tables, 6" x 3". 


Table of Contents. —Preface; EKO MeO SS UMeauction 1) General eae 2) Berries in the 
Landscape; 3) Strawberries; 4) Brambles; 5) Blueberries; 6) Grapes; 7) Minor Crops Appendix: North 


American Nursery Sources for Berry Plant and Other Resources: Plant Name 
Index; and Subject Index.—Review forthcoming i in Sida 19(3), 2001. 


SIDA 19(2): 439, 2000 


440 SIDA 19(2) 


G, Terry SHarrer. 2000. A Kind of Fate: Agricultural Change in Virginia, 1861-1920. (/SBN 
-8138-2569-5, hbk.).lowa State University Press, 2121 South State Street, Ames, |A 
50014-8300, U.S.A. (Orders: 800-862-6657, 515-292-0155; www.isupress.edu). $49.95 

pbk. 256 pp., b/w photos, 6" x 9" 


From the dustjacket.—‘A Kind of Fate: Agricultural Change in Virginia, 1861-1920 addresses how mod- 
ern agriculture in Virginia developed as a result of widening social acceptance of government 
intervention in agricultural affairs: an infusion of capital from increasing commodity prices, land 
values, and credit; and biological research to combat diseases. In addition to descriptive quotes 
from the period, poignant photographs capture the essence of these farmers’ daily trials and tri- 
phs.” 


um 

Contents.—Acknowledgments; Introduction; 1) Swept Away: Disease: Epidemics and Epizootics, 
A New Order of Things, Untenable Country; 2) New Needs, Old Ideas: The Dogma of “Soil Fertility,” 
Crops and Crises; 3) Toil and Trouble: Life in the Country, Debt, Taxes, and Despair; 4) Professing 
Change: Growing Knowledge: The Virginia Agricultural Experiment Station, Reaching the Farmers; 
5) The New Farming: Drive for Production, Dairying: Progressive Exemplar, Capital and Credit; 6) 
Reforming Fate: Farmers and Tenants, The Rural Life Movement in Virginia, End of an Era;Conclusion: 
Two Generations; Bibliography; and Index. 


J.K. Morton and Joan M. Venn. 2000. The Flora of Manitoulin heal and the idan dneee 
Islands of Lake Huron, Georgi SSN 
0317-3348, pbk). University of Waterloo Biology Series, eioae Biology ris 
Department of Biology, University of Waterloo, Waterloo, ON, CANADA N2L 3G1 (51 
888-4567, ext. 3751; 519-746-0614 fax; jvenn@sciborg.uwaterloo.ca). $37.45 a 
bound (Canada); $37.50 spiral bound (outside Canada): $50.83 hbk. ae. $50.00 
hbk. (outside Canada). 374 pp., 124 color photos, 997 range maps, 7" x 


“Manitoulin Island and the many smaller islands in the surrounding waters of Georgian Bay, Lake 
Huron and the North Channel, in the Great Lakes Region of Canada, is an area of unique biological 
interest with a remarkably rich flora (about ~ of the total vascular flora of Canada).” 

The Flora accounts for 1350 kinds of vascular plants. The 997 distribution maps are computer 
drawn from a database of over 66,000 records from the region."Introductory chapters deal with 
the geology, climate, post-glacial history, vegetation, conservation and history of botanical explo- 
ration in the region."The 124 colored illustrations are real nice and represent the more interesting 
and beautiful plants of the region. 

xt and the maps have been completely revised in the 3" edition; the color figures are un- 
changed from the 2 edition. There is no family key nor do families have keys to genera and gen- 


L 


era to species; an occasional key is provided to species.—Barney Lipscomb. 


SIDA 19(2): 440. 2000 


BOOKS RECEIVED 441 


Ronato M. Lanner. 2000. Conifers of California. (ISBN 09628505-3-5, pbk.;09628505-4-3, 
hbk.). Cachuma Press, PO. Box 560, Los Olivos, CA 93441, U.S.A. (805-688-0413; 
cachuma@silcom.com). $24.95 pbk., $36.95 hbk. 288 pp., 54 original full-page wa- 
tercolor illustrations, 7" x 9" 


From the cover.—"Conifers of California is the first book devoted to all of the state's conifers. Its au- 
thor, Ronald M.Lanner (professor emeritus, Utah State University) h plored, studied, and taught 
about forest trees in the West for 40 years. In Conifers of California he sh his expertise and intro- 
duces each of Ciliigmias cone-bearers in an engaging text that serves as both natural ree and 
field guide. Lanner’s narratives are accompanied by detailed identification information, watercolor 
botanic illustrations by the late Eugene O. Murman, color photographs of each species by ne 
known landscape photographers, and distribution maps 

Contents. —Preface; Acknowledgments; Eugene O. Murman, the Artist; The Names of Trees; 
Cones; Pinaceae, The Pine Family; Cupressaceae, The Cypress Family; Taxodiaceae, The Baldcypress 
Family; Taxaceae, The Yew Family; Appendices: A: California's Soft Pines and Hard Pines; B: Conifer 
Hybrids in California; C: A Key to the Genera Based on Characters of Mature; D: A Key to the Genera 
Based on Leaf Chatectels Born Lower Crown Branches; E: Alphabetical List of Conifers Growing 
Naturally Within Califort tated Bibliography; and Index. 

| must say une watercolor botanical illustrations are mouth watering! They are beautiful. And 
the photography is stunning and shows off the fascinating and often picturesque California land- 
scape. The book is beautifully designed and would make a wonderful coffee table addition any- 
where.—Barney Lipscomb 


Deni Bown. 2000. Aroids: Plants of the Arum Family. Second Edition. (ISBN 0-88192- 
485-7, hbk). Timber Press. Inc., The Haseltine Building, 133 S.W.Second Avenue. Suite 
450. Portland, OR 97204-3527, U.S.A. (503-227-2878, 503-227-3070 fax; www. 
timberpress.com). $34.95 hbk. 468 pp. 108 color photos, 50 line drawings, 6" x 9". 


Table of Contents. —Foreword to Second Edition; Foreword to the First Edition; Preface; Acknowl- 
SoG mens Introduction; 1) Vellguons: ona APNG, 2) Of Tails and traps and the Underworld; 3) 


rs;4) Aquatics and A ) A Place in the Sun;6) In the Shadows; 7) Towards the 
Light; 8) The Titans; 9) An Acquired Taste; 10) Acids and ea sae Cultivation; Checklist of 
Aroid Genera; Glossary; References; and Index.—Review forthcoming | a 19(3), 2001. 


Dorotuy J. CatLaway and M. Bretr Cattaway (Editors). 2000. Breeding Ornamental Plants. 
(ISBN 0-88192-482-2, hbk.). Timber Press. Inc., The Haseltine Building, 133 S.W. Sec- 
ond Avenue. Suite 450. Portland, OR 97204-3527,U.S.A.(503-227-2878, 503-227-3070 
fax; www.timberpress.com).$34.95 hbk. 359 pp.88 color photos, 10 b/w photos, 26 
line drawings, 26 tables, 6" x 9" 
Preface; PER NOWwlece met 1) Oonetes and its SAD DICoHONS 2) Fiape Ereeeliors apace) Malet 
3) Breeding D 5) lostas; 
7) Breeding Ornamental Aroids; 8) Breeding “AfricanViolets; 9) Breeding Gesneriads; 10) Breeding 
Amaryllis; 11) Breeding Penstemon; 12) Breeding Rhododendrons and Azaleas; 13) Breeding Kalmia: 
Mountain Laurel and its relatives; 14) Breeding Camellias; 15) Breeding Lilacs: Plant of History, Plant 
for Tomorrow; 16) Breeding Magnolias; 17) Breeding Oaks: A New Frontier; Appendix: Sources of 
Plant Breeding Supplies; Glossary; Notes on Contributors; and Index of Plant Names.—Review forth- 
coming in Sida 19(3), 2001. 


SIDA 19(2): 441. 2000 


442 SIDA 19(2) 


Kay YATSKIEVYCH. 2000. Field Guid 1j Wildlf (ISBN 0-253-21420-3, pbk,). Indiana 
University Press, 601 N. Morton Street, Bloomington, IN 47404, U.S.A. (Orders: 800- 
842-6796, 812-855-8507 fax).$17.95 pbk. 372 pp.640 color photos, line drawings, 


" x 3" 


Contents.—Preface; Acknowledgments; Introduction: Included species, Species numbers, Photo- 
graphs, Drawings, Scientific names, Authors of scientific names, Vernacular names, Measurements 
of plant parts; Main paragraph of species entry: Sizes of plants and flowers, Distinguishing charac- 
teristics, Habitats and localities, Blooming dates, Status in the state, Federal status; Glossary; What is 
a flower?: Aster family terminology; Flower ee Photos and text; Selected references; and Index. 
This is a nicely done wildflower guide; Hoosiers should be proud.—Review forthcoming in 
Sida 19(3), 2001 


ALAN E. Bessette, WILLIAM C. Roopy, and Arveen R. Bessette. 2000. North American Boletes: A 
Color Guide to the Fleshy Pored Mushrooms. (ISBN 0-8156-0588-9, hbk.). Syra- 
cuse sen ae Press, Syracuse, NY 13244-5160,U.S.A.$95.00 hbk. 396 pp.450 color 
photos, 7 


Contents.—Preface; Acknowledgments; North a fees in Brief; Introduction to the 
Boletes; Typical Bolete Fruiting Body: Macroscopic Features; How to Use This Book; How to Identify 
Boletes; Field Key to ne: ge eaalneda . cate Gener any epee Undescribed Boletes, 
Hypomyces, and P to Common Names; and 
Index to Genera Species. -Reven fonpcoming! in Sida 19(3), 2001. 


Git Nevson. 2000. The Ferns of Florida. A Reference and Field Guide. (ISBN-1-56164-197- 
9, pbk., 1-56164-193-6, hbk.). Pineapple Press, Inc., PO. Box 3899, Sarasota, FL 34230, 
U.S.A. (www.pineapplepress.com). $27.95 hbk. 208 pp. 200 + color photos, 6" x 9". 


From the back cover—"At least 163 kinds of ferns have been recorded in Florida. From northern 
species whose southern ranges extend into the rolling uplands of the state’s northern tier to tropi- 
cal treasures that spill over the state's southern tip, Florida is without parallel in fern diversity and 
number. No other like-sized region of the continental United States can boast such an expanded 
assortmen 

s field guide is the first in 25 years to treat this amazing variety of ferns. Following in the 
sane of John K. Small’s 1931 Ferns of Florida, and Robert Long and Olga Lakela’s 1976 edition of 
the same name, this new volume catalogs and illustrates all of Florida's native and naturalized fern 
species. Color plates feature more than 200 images, some of which include rare species never be- 
fore illustrated in color. Rare and hard-to-find species such as some of our spleenworts and filmy 
ferns are included, as well as such common species as the cinnamon fern pictured on the front 
cover. 

Clearly written detailed descriptions, along with helpful identification tips and a selection of 
botanical keys all of which are based on field observations help readers identify plants in the wild. 
Also included are notes on each species’ growth form and habitat, as well as general remarks about 
its botanical and common names, unique characteristics, garden use, and history in Florida. 

ther important features of the book include a complete glossary, a section on the fern life 
cycle and structure, a history of fern study in Florida, a discussion of some of the state’s best natural 
ferneries, and an extensive bibliography."—Review forthcoming in Sida 19(3), 2001 


SIDA 19(2): 442. 2000 


BOOKS RECEIVED 443 


Ken E. Rocers. 2000. The Magnificent Mesquite. (ISBN 0-292-77105-3, hbk.). University of 
Texas Press, RO. Box 7819, Austin, TX 78713-7819, U.S.A. (512-47 1-4032).$24.95 hbk. 
167 pp., 18 color photos, 2 line drawings, 5 maps, 12 tables, 5" x 9" 


From the dustjacket—The mesquite tree has deep roots in the American Southwest, literally and 
figuratively. A reliable source of food and shelter even in the severest droughts, it sustained Ameri- 
can Indians for centuries. Today, mesquite is widely popular for barbecuing, woodworking, furni- 
ture making, flooring, sculpture, jewelry, and food products ranging from honey to jelly and syrup. 
Even ranchers, who one fought to eradicate mesquite s an invasive weed, have come to value its 
multiple uses on bi managed rangeland. 

In this b ritten especially for a general readership, one of the world’s leading authorities 
on mesquite presents a wealth of information about its natural history and commercial, agricul- 
tural, and woodworking uses. Ken Rogers describes the life cycle, species, and surprisingly wide 
distribution of the mesquite, which is native or naturalized not only in the Southwest and Mexico, 
but also in India, Africa, Australia, South America, and Hawaii.He discusses the many consumer and 
woodworker uses of mesquite at length—even giving instructions for laying a aoe Wee 
floor and making mesquite bean jelly. He also looks into the ways that people are using mesquite 
in nature, from rangeland management in the Southwest to desertification prevention in arid coun- 
tries. Color photographs and maps complement the very readable text 

Contents.—Introduction; Mesquite: What Is It”; Predators of Mesquite; Mesquite in Texas And 
the SOP EIWESTEN United States; The Uses of Mesquite; The Mesquite in Verse; Worldwide Aspects 
of Prosopis; Texas Ebony—Mesquite's Astonishing Cousin; Mesquite’s Future; Appendix |. The Genus 
Prosopis amily i cumainesie) ieaeeee and Its Species; Appendix 2. Sources of Information; Ref- 
erences and Further Reading; and Index.—Review forthcoming in Sida 19(3), 2001. 


JoHN J. Wiens. 2000. Phylogenetic Analysis of Morphological Data. (ISBN 1-56098-816- 
9, pbk.; 1-56098-841-X, hbk.). Smithsonian Institution Press, PO Box 960, Herndon, 
VA 20172-0960, U.S.A. (Orders:800-782-4612).$26.95 pbk.,$49.95 hbk. 220 pp. Black/ 
white Lapune eo 


ntents.—1)M t tics:Conflicts, Artifacts, and Misconceptions; 
2) Character Selection aid the Methodology of ene Phylogenetics; 3) Discovery of Phy- 
logenetic Characters In Morphometric Data; 4) The Usefulness of Ontongeny in Interpreting Mor- 
phological Characters; 2) Coding Morphological Variation Within Species and Higher Taxa for Phy- 
logenetic Analysis;6) Hybridization and Phylogenetics; Special Insights From Morphology; 7) Using 
Stratigraphic infownation in Phylogenetics; 8) Logical Problems Associated With Including and Ex- 
cluding Characters During Tree Reconstruction and Their Implications for the Study of Morpho- 
logical Character Evolution; and Index.—Review forthcoming in Sida 19(3), 2001. 


SIDA 19(2): 443. 2000 


 —> 


“You don't have to be a botanist to use it! All the tools you need 


to teach plant taxonomy, family characters, classification, etc. 


Excellent introductory materials on the history and geology of 
North Central Texas. Great for identifying indigenous plants in 
an ecosystem! Essential for student wildflower collection 
projects! Use this one book instead of five, it has it all.” 
—Sharon Foster, 7th grade teacher, 

Fort Worth Country Day School 


“The best available, user-friendly source of information about 
Our native and naturalized plants. This book should be in the 


library of every gardener, landscape architect, horticulturist 


urban planner, farmer, rancher, and wildflower enthusiast.” 
—Howard Garrett, The Dirt Doctor 


“INCREDIBLE, FANTASTIC, WONDERFUL, BEAUTIFUL, ETC. WOW! 
To be more specific: COMPREHENSIVE, ENCYCLOPEDIC, 
INNOVATIVE. With every species illustrated - it’s a new 
departure in Texas floras, a breath of fresh air.” 

—Marshall Johnston, Co-author, 

Manual of the Vascular Plants of Texas 


“The Illustrated Flora on North Central Texas is a magnificent work. 
This abundantly illustrated volume will not only be of use to 
those people wanting to identify Texan plants, it will be a 
model of how a local flora should be prepared.” 

—Sir Ghillean Prance FRS 


Director, Royal Botanic Gardens, Kew 


THEY ARE SAYING ABQUT.... 


Nets, 
q ust 
3 : io 


=~ 
_ 


SMa ER's 
RATED 
RA OE 


CAA bE hie WHMIS FT 


SHINNERS & MAHLER’S 
ILLUSTRATED 
FLORA OF 
NORTH CENTRAL 
TEXAS 


Now IN ITS SECOND PRINTING ! 


DETAILED BROCHURE WITH ORDER FORM 
AVAILABLE BY REQUEST ON THIS GROUND 
BREAKING NEW FLORA. 


CALL BRIT Press at 
817 332-4441 or 
EMAIL! SIDA@BRIT.ORG 
FOR HOW TO ORDER. 


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Ee: ‘ cr i wmpomeect os a 
ic Sane 


pred 


ISSN 0036-1488 


SOURCE OF 


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Barney L. Lipscomb, Editor 
Botanical Research Institute of Texas 
509 Pecan Street 

Fort Worth, Texas 76102-4060, USA 
817 332-4441 / 817 332-4112 FAX 


Electronic mail: sida@brit.org 


Home page at the URL: http://www.brit.org/sida/ 


SIDA CONTRIBUTIONS TO 


Bsr. 


Botany was founded John W. Thieret, Associate Editor 


by Lloyd H. Dept. of Biological Sciences 
: Northern Kentucky Universit 
Shinners, above left _ i ea a 
Highland Heights, KY 41099 USA 
in 1962. Inherited 
by William F. Félix Llamas, Contributing Spanish Editor 
Mahler, Director Dpto. de Botanica, Facultad de Biologia 
‘ Universidad de Leon 
Emeritus of BRIT, ae 
E-2471 Leon, SPAIN 
above right, in 
aay atieesinice Guidelines for contributors are available upon request 
1993 it has been and on our SIDA home page as well as the last page 
of each issue. 
published by BRIT. Subscriptions for year 2001: 
$27. Individual 
550. USA Institutions 
$60. Outside USA 


numbers issued twice a year 


O©SIDA, CONTRIBUTIONS TO BOTANY 
VOLUME 19, NUMBER 3, PAGES 445-766. 
23 Aucust 2001 

CopyriGHT 2001 


BOTANICAL RESEARCH INSTITUTE OF TEXAS 


PRINTED IN THE UNITED STATES OF AMERICA 


ISSN 0036-1488 


TABLE OF CONTENTS 


FLORISTIC DOCUMENTATION IMPERATIVES: SOME CONCLUSIONS FRON TEMPO SURVEYS IN PAPUA NEW GUINEA 
W. TAKEUCHI and M. GOLMAN—445 


A LEAF BLADE ANATOMICAL SURVEY OF MUHLENBERGIA (POACEAE: MUHLENBERGIINAE) 
PAUL M. PETERSON and YOLANDA HERRERA-ARRIETA—469 


PSEUDOGNAPHALIUM AUSTROTEXANUM (ASTERACEAE: GNAPHALIEAE), A NEW SPECIES FROM SOUTHEASTERN TEXAS 
AND ADJACENT MEXICO 
Guy L. Nesom—50 


TAXONOMIC NOTES ON KEYSSERIA AND PYTINICARPA (ASTERACEAE: ASTEREAE, LAGENIFERINAE) 
Guy L. Nesom—513 


A NEW SPECIES OF SACCOGLOSSUM (ORCHIDACEAE) FROM THE HANS MEYER RANGE, NEW IRELAND, PAPUA NEW GUINEA 
N.H.S. Howcrort—519 


NOMENCLATURAL CHANGES IN PENNISETUM (POACEAE: PANICEAE) 
JOSEPH K. WiprFF—523 


REEVALUATION OF AYLACOPHORA AND (ASTERACEAE: ASTEREAE) 
ost M. BONIFACINO and GISELA Gancue = 5a). 


GEOGRAPHIC VARIATION AND TAXONOMY OF NORTH AMERICAN SPECIES OF MIRABILIS, SECTION OXYBAPHOIDES (NYCTAGINACEAE) 
RICHARD SPELLENBERG and SerGio R. RODRIGUEZ TUERINA—539 


A REVISION OF SALVIA SECTION HETEROSPHACE (LAMIACEAE) IN WESTERN NORTH AMERICA 
Jay B. Wacker and WAYNE J. ELISENS—571 


TAXONOMY OF STENARIA (RUBIACEAE: HEDYOTIDEAE), A NEW GENUS INCLUDING HEDYOTIS NIGRICANS 
EDWARD E. TERRELL—591 


NOTES ON VARIATION IN PSEUDOGNAPHALIUM OBTUSIFOLIUM (ASTERACEAE: GNAPHALIEAE) 
Guy L. NESoM—615 


BIOLOGICAL STATUS OF ARGYTHAMNIA LAEVIS (EUPHORBIACEAE) 
B.L. TURNER—621 


VALIDATION OF THE NAME OROBANCHE LUDOVICIANA SUBSP. MULTIFLORA (OROBANCHACEAE) 
HEATHER L. WHITE and Watter C. HOLMES—623 


AN ANOMALOUS POPULATION OF ASTER (ASTERACEAE: ASTEREAE) SENSU LATO IN MICHIGAN 
Guy L. Nesom—625 


BIBLIOGRAPHICAL NOTES ON THE PUBLICATION OF WOOLWARD’S MONOGRAPH ON THE GENUS MASDEVALLIA (ORCHIDACEAE) 
Guido J. BRAEM—633 


DOCUMENTED CHROMOSOME NUMBERS 2001:1.CHROMOSOME NUMBER OF LUPINUS HAVARDII (FABACEAE) 
B.L. TURNER and A.M. PoweLL—639 


FLorisTics 


GNAPHALIUM EXILIFOLIUM (ASTERACEAE: GNAPHALIEAE) IN COLORADO AND SOUTH DAKOTA 
Guy L. Nesom—641 


THE VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 
Mac H. ALForD—645 


NOTES ON THE INCREASING PROPORTION OF NON-NATIVE ANGIOSPERMS IN THE MISSOURI FLORA, WITH REPORTS OF THREE 
NEW GENERA FOR THE STATE 
GEORGE YATSKIEVYCH and JAY A. RAVEILL—701 


GLOCHIDION PUBERUM (EUPHORBIACEAE) NATURALIZED IN SOUTHERN ALABAMA 
MiriAM L. FEARN and Lowe tt E. UrsAtscH—711 


RADIATE AND ERADIATE INDIVIDUALS IN GRINDELIA NUDA (ASTERACEAE) 
RT. HARM 


CUPRESSUS ARIZONICA (CUPRESSACEAE) NEW TO THE DAVIS MOUNTAINS OF WEST TEXAS 
JOHN P. KaRGES and James C. ZECH—719 


DIPSACUS FULLONUM (DIPSACACEAE) AND VERBESINA WALTERI (ASTERACEAE), NEW TO TEXAS 
JASON R. SINGHURST and Water C. HOLMES—723 


CAREX ABSCONDITA (CYPERACEAE: CAREYANAE) IN LOUISIANA 
David J. ROSEN—727 


/D R 


PREVIOUSLY UNREPORTED FROM LOUISIANA 
ea ROSEN, STANLEY D. JONES, and JosePH K. WiprF—731 


A SCUTELLARIA (LAMIACEAE) NEW TO NORTH CAROLINA AND A KEY TO THE SMALL-FLOWERED CAROLINA CONGENERS 
ALEXANDER KRINGS and JOSEPH C. NEAL—735 


LESPEDEZA CUNEATA (FABACEAE), A FIRST RECORD OF ITS OCCURRENCE IN MEXICO 


EDUARDO ESTRADA C. and CARMEN YEN M.—741 


EMOIR 
Rupert C. BARNEBY (OCTOBER 6, 191 1—December 5, 2000). A MEMOIR 
RICHARD SPELLENBERG—745 


BOOK REVIEWS AND NOTICES 512, 522, 590, 620, 638, 642, 644,700, 710,714,718, 722, 726, 730, 740, 744, 752 


Index to new names and new combinations in Sida 19(3), 2001. 


DITAXIS HUMILIS FORMA LAEVIS (A. GRAY EX TorREY) B.L. TURNER, FORMA NOVA.—622 
KEYSSERIA LAUTERBACH SECT. SANDWICACTIS NESOM, SECT. NOV.— 514 
MIRABILIS LAEVIS VAR. CRASSIFOLIA (CHOISY) SPELLENB., COMB. NOV.—549 
MInABIEIS LAEVIS VAR. VILLOSA (KELLOGG) SPELLENB., COMB. NOV,—551 
NuTT. (NuTT.) COLLINS EX H.L. WHITE & W.C. HOLMES, STAT. NOV,—623 

PENNISETUM PENNISETIFORME (HOCHS. & STEUD. EX STEUD.) WiIPFF, COMB. NOV,—527 
PENNISETUM SETIGERUM (VAHL) WIPFF, COMB. NOV.—526 
PENNISETUM SOMALENSIS (CLAYTON) WiIPFF, COMB. NOV.—527 
PSEUDOGNARHATION AUSTROTEXANUM NESOM, SP. NOV.—507 

AICRADENIUM (WEATHERBY) NESOM, COMB. ET STAT. NOV.—618 
comin. PICKERINGII (A. GRAY) NESOM, COMB. NOV.—5 16 
SACCOGLOSSUM TAKEUCHI] HOWCROFT, SP. NOV.— 


STENARIA BUTTERWICKIAE (TERRELL) TERRELL, COMB. NOV—595 

STENARIA MULLERAE (FOSB.) TERRELL, COMB. NOV.—597 

STENARIA MULLERAE VAR. POOLEANA (B.L. TURNER) TERRELL, COMB. ET STAT. NOV.—598 
STENARIA NIGRICANS (LAM.) TERRELL, COMB. NOV.— 

STENARIA NIGRICANS VAR. BREVIFLORA TERRELL, VAR. NOV.—605 

STENARIA NIGRICANS VAR. FLORIDANA (STANDL.) TERRELL, COMB. NOV.—605 

STENARIA NIGRICANS VAR. GYPSOPHILA (B.L. TURNER) TERRELL, COMB. NOV.—606 
STENARIA RUPICOLA (GREENMAN) TERRELL, COMB, NOV.— 

STENARIA UMBRATILIS (B.L. ROBINSON) TERRELL, COMB. NOV.—610 

STENARIA UMBRATILIS VAR. BREVIPEDICELLATA TERRELL, VAR. NOV.—61 1 


¥ 


FLORISTIC DOCUMENTATION IMPERATIVES: 
SOME CONCLUSIONS FROM CONTEMPORARY 
SURVEYS IN PAPUA NEW GUINEA 


W. Takeuchi M.Golman 
Botanical Research Institute of Texas (BRIT) Divisional planader 
c/o Lae Nanci Herbarium gua Papua New Guinea Forest Authority 
Papua New Guinea Res stitute PNG Nati ona Fors ae 
PO. Box 314, Lae, iirc Province 411 PO. Box 5055, Boroko 
PAPUA NEW GUINEA National ne itol District 
wtakeuchi@global.net.pg PAPUA GUINEA 
mgolman@datec.com.pg 
ABSTRACT 
favecennyeais Par i I J-assessment | 1 i ingl 
f] ic esti f site value. Although some of the results have been widel y 


dissonant’. there ner been few attempts at overall synthesis despite the obvious need for such 
review. A commento: on botanical aon and epcumenvaton curing ee past decade is pro- 
f,,1£57] 


vided, with particular f : of goals 
implicit in comprehensive bioinventory. 


TOK IGO PAS (MELANESIAN TOK PISIN) 


Eongna nau tasol, tein teat luksave na ise parm aut ibia ame Sait moa long ol bus na diwai 
pnaibin kamap long planti 


hap mabaue ibin igat wok oe esol i kamap long aiepela wok lone on haumas bus na diwai 
istap long Papuasia. Dispela em i ripot o toksave long ol wok painim aut wei ikamap long ten pela yia 
igo pinis na luksave long wok ibai kamap bihain taim long wei bilong painim aut ol diwai na bus. 


ABSTRACT 
Mi, MFM RID 5S OMMOBSECHMTSLOOAKELT. GDEVET 
ERAYbKARCKDTAHADONANTT VT REOSEMLCEV, BODORS 
HARBROR<KRRENTNS. CHOOMROBEM., HROMAORRICH 
BRIBH CBAC EISRSDPCHSICODDDHSY, ENKHOBALEBSNTE 
i. BRM. REOMRAKCHRE ORO TAZMRERSICHF ONT 
WSDEDDPORMICRASH TEMS, HKIOFM ORME mt REN IC RT 
Smee. 


INTRODUCTION 


]: 


In spite of its status asa global center for | ification, Papua New Guinea 
(PNG) has the dubious distinction of being one of Malesia’s most inadequately 
surveyed nations. Stevens (1989) had defined the relatively well-collected areas 
in PNG as having a collections density of 50-100 specimens per 100 sq km. But 


SIDA 19(3): 445 — 468. 2001 


446 BRIT.ORG/SIDA 19(3) 


even by this easy measure, the well-collected localities identified by his criterion 
were primarily confined toa few principal drainages and high mountains. Re- 
cent analysis shows that the background level of collection density averages 
less than 25 per 100 sq km over New Guinea as a whole (Conn 1994), a rate 
substantially below the standard regarded by Stevens as a minimal baseline. 
Within the Malesian region, only the Celebes and Sumatra have comparably 
low collection indices (Stevens 1989). Although Steenis (1950) had estimated 
that 50 years of coordinated exploration would be required to document the 
New Guinea flora, the rate of documentation has not improved since that as- 
sessment was made, and has in fact dramatically declined within the last 30 
years (cf. Conn 1994). If rational conservation strategies and land-use policies 
are to be devised and implemented by indigenous agencies, the present trends 
in documentation must be reversed. The urgency for corrective action is espe- 
cially acute when the target country represents a biodiversity hotspot within 
which social and economic changes are expected to intensify. 

Papua New Guinea's population growth rate of 2.3% is now one of the high- 
est in the Pacific region. From a present base of 4.7-5.0 million, the number of 
people is expected to double by 2024 (Gumoi & Sekhran 1995). The demographic 
projecti pecially consequential because an estimated 84% of the popu- 
lation is rural, and all such households are dependent to some degree on slash 
and burn agriculture (ibid). Currently, an estimated 200,000 hectares of land 
are cleared annually for subsistence, from which 20,000-30,000 hectares rep- 
resent natural-growth forest permanently removed by various clear-cut opera- 
tions including industrial logging (Filer 1995). 

With the anticipated increases in human population, subsistence activi- 
ties are expected to progressively mimic the effects of forest clear-felling as fal- 
low cycles are accelerated (Louman & Nicholls 1995). When PNG enters the 
steep part of the predicted population growth curve, serious environmental 
impacts are likely to occur. There are already mounting indications of impend- 
ing failure in the subsistence systems of several provincial areas due to intensi- 
fication of cropping rotations (Levett & Bala 1995). 

At present, PNG still retains 70% of its primary forest cover (McAlpine & 
Quigley 1998) and is one of only four tropical countries with extensive tracts of 
original vegetation (Suzuki 1993). This remarkable state of preservation has 
unfortunately fostered a complacency among science professionals by encour- 
aging an expectation that current inadequacies in documentation can always 
be reversed by future action. The underlying assumptions are not likely to per- 
sist however. Time will start running out sometime during the next generation. 

The poor state of floristic documentation has many and significant mani- 
festations in Papuasian botany. A disproportionate number of taxa are known 
from single collections, and taxonomic knowledge even at family level is often 
highly superficial Johns 1993). Virtually nothing can be said of the basic biology 


FLORISTIC 447 


and populational variation for many of the most important Papuasian plant 
groups (Stevens 1989). While it is generally conceded that certain mountainous 
areas are hotspots for floristic end it is impossible to identify low-elevation 
centers with the scanty data at hand. Some consequences of the past emphasis 
on high elevation exploration can be seen in recent results from The Nature 
Conservancy's lowland survey of Josephstaal, where 8 species were newly de- 
scribed from accessible sites near the principal national highway in Madang 
Province (Huynh 2000; Takeuchi 2000a, 2001). Lowland environments are pre- 
ferred venues for economic development, so there is considerable potential for 
floristic losses in this zone. Continued neglect of botanical documentation in 
lowland habitats can have severe consequences in any future intensification of 
commercial logging, considering that over 50% of PNG’s loggable forests (as 
defined by industry standards) are in the lowlands (Louman & Nicholls 1995). 
Of the forest blocks currently representing merchantable stands, 67% also occur 
in medium-crowned lowland hill forest (sensu Paijmans 1975; or forest category 
Hm on Hammermaster & Saunders 1995), a vegetation type which is probably 
Papuasia’s richest floristic formation (Llouman @ Nicholls 1995). 

Several competent observers have enumerated localities of particular value 
and urgency for exploratory survey within PNG Johns 1993; Steenis 1950; 
Stevens 1989). Acommon thread extending through all these recommendations 
is that the uaa status of highlighted areas has hardly improved since 


the time of the S tary. Very little has changed with respect to the 
quantity and quality of botanical data over the past several decades. The ak 
of substantive progress adversely tst d ecological 


and ultimately impedes eee management and development within the 
areas in question. 


SOCIAL AND INFRASTRUCTURAL CONSIDERATIONS 


In PNG, any activity requiring access to natural resources must include consid- 
eration of the traditional land tenure system. An estimated 97% of the country 
is under customary ownership, subject to complex systems of usage rights and 
social relationships which are themselves superimposed over a multitude of 
cultural-linguistic traditions (Crocombe 1974; Holzknecht 1995). Alienated land 
and properties otherwise under governmental control are virtually nonexist- 
ent. Irrespective of endorsements from external agencies, the final arbiters in 
land-use issues are the village clans and landowners of specific forest blocks 
(ie. the ‘papa graun’). 

Because so many prerogatives reside with local villagers, direct negotiations 
with the customary tenants are mandatory for any scientific program. This is 
not so easily done when an investigation’s principals are overseas-based. Due 
to the proliferation of advocacy groups with environmentalist agendas, many 
landowners have also become conditioned to regard extractive activities on their 


448 BRIT.ORG/SIDA 19(3) 


land ipartienian’y by foreigners) with considerable suspicion. This situation 


applies to bioprospecting. Survey operators do well to avoid prospec- 
tive collecting altogether, because of objections that have been raised in relation 
to such activities. Any contemplated project would be required to explain its 
activities to an oftentimes skeptical and uninformed audience. Having a func- 
tional knowledge of the lingua franca (Melanesian tok pisin) is essential. 

A significant consequence of the primacy of customary rights in Papuasia 
is that certain Western mechanisms for permit issuance and resource access are 
culturally irrelevant in PNG. There is for example, no such requirement asa ‘plant 
collecting permit. Even if such permits were established by government agen- 
cies, they could never be enforced at the local level where botanical collecting 
actually occurs. Only the customary landowners can grant approval for removal 
of materials from their territory. Paradoxically, while this eliminates much of 
the bureaucracy characteristic of Eurocentric management systems, survey 
operations are often rendered more complicated and unpredictable, because the 
activities are entirely subject to the whims of individual landowners. 

Modern commentators have been unanimous in expressing a need for de- 
veloping local capacity as a prerequisite for longterm assessment and manage- 
ment of PNG’s biodiversity (Beehler 1993; Conn 1994; Damas 1998; Johns 1993; 
Sekhran & Miller 1995). Public sector agencies in PNG are subject to unpredictable 
changes in government support due to shifts in political direction. Based on pre- 
vious trends, it is very unlikely that vital commitments to science capacity 
building will occur through in-country funding. Collaborative studies are a 
potentially effective means for improving internal capabilities when appropriate 
agencies are engaged as partners in research. Programs which provide’ lor partici- 
pation of qualified counterparts can make longterm contributi bioinventory, 
but the partnerships must be carefully selected and not reel convenient. 

The PNG infrastructure in science has experienced profound changes over 
the last thirty years. During the colonial administration and for a brief time after- 
wards, many of the functionsassociated with floristic survey lly invested 
in highly capable units such as Lae Herbarium (LAE) and the former Department 
of Forests. Government facilities in the 1960s and early 70s virtually monopo- 
lized plant exploration within PNG. In contrast, nongovernment organizations 
(NGOs) were conspicuous by their overall absence from activities involving 
botanical documentation. However after PNG became a sovereign state, the capa- 
bilities of national agencies for floristic work progressively declined as budgets 
were subjected toa political reordering of priorities (cf. Conn 1994). The earlier 
priorities are unlikely to be restored in the future because institutional and social 
realities have been so completely transformed. For example, there are now no 
Ph.D.-level professionals in the PNG National Forest Service, and currently only 
one Me SC: eccrine is seine, with the national herbarium. 

V ¢ vpabilities have been mitigated within 


FLORISTIC 449 


the last LO years by countervailing developments in the nongovernmental sector. 
A seminal event was the Conservation Needs Assessment (CNA, Beehler 1993), 
the first countrywide plan to define comprehensive priorities for conservation 
and research action. The CNA stimulated establishment of a multitude of Wildlife 
Management Areas (WMAs) and of associated projects based on the Integrated 
Conservation and Development (ICAD) model (cf. Saulei & Ellis 1998). Not co- 
incidentally, many of the post-CNA initiatives encompass at least in part, the 
areas prioritized by the CNA. Together with these developments, a number of 
NGO entities have assumed effective jurisdiction over the WMAsand their land- 
owner groups. Many of PNG’s prime wilderness environments are presently 
included under NGO/Management Area partnerships. The total biodiversity 
represented by such partnerships is very substantial even though the arrange- 
ments collectively comprise only ca. 8% of the PNG land area. In addition, a 
large backlog of sites is under consideration for future conservation action. 

The combined effect of these events is that the hierarchy of PNG science- 
related administration has been transformed by the creation of a new infra- 
structure. Due to the relative recency of the new arrangements, there has been 
little change in the way biological surveys are conducted in PNG. However nearly 
all the WMA-related NGOs maintain a fulltime presence within their respective 
wards, so the opportunities for achieving effective ity integration with 
surveys are now very ee The Mes typically include resident-coordi- 
nators and j iaison with landowner groups, 
critical functions that government interests can no longer provide in the remote 
areas. From an operational perspective, each of the NGO/WMA combinations 
is the equivalent of a research facility. The elements for multilateral surveys 
combining professional and village participants are thus in place, requiring only 
that the individual components be drawn together under a common plan. A 
future schedule for comprehensive bioinventories could be constructed using 
the government planning instruments on one hand and community level 
implementors composed of WMA/NGOs on the other. Linkages of this sort are 
already the basis for several contemplated operations. 


PARATAXONOMIST ENHANCEMENT OF SURVEYS 


The sheer numbers of species involved in documentation and inventory are 
overwhelming when approached from traditional perspectives in collection and 
curation. Time honored traditions in tropical exploration are no longer adequate 
to the tasks of acquiring and analyzing large collection sets. A revised approach 
to floristic documentation is clearly required, and is perhaps best adapted from 
the experiences of entomologists faced with problems similar to those in bo- 


tanical inventory. 
Through refinement of bl hodol first developed by INBio 
in Costa Rica Janzen et al. 1993), PNG-based researchers at the Parataxonomy 


450 BRIT.ORG/SIDA 19(3) 


Training Center (PTC) have devised practical solutions to the time demands 
presented by intensive sampling in rich tropical habitats (Basset et al. 2000; 
Novotny etal. 1997). Their approach has been to train local villagers (as parataxono- 
mists) in the fundamentals of collecting, specimen sorting, identification, and 
computer-based data management. The few international professionals are pri- 
marily engaged as instructors, quality control agents, and ultimately as data 
interpreters. By focusing the intervention of individuals to the points where 
their expertise is most effective, the research process is thus streamlined and 
accelerated. The immediate product of these arrangements is that extensive 
specimen sets have been acquired and processed within time frames that would 
ordinarily require prohibitive inputs. The demonstrated success of such orga- 
nization in entomology by itself shows that the protocols can work for botany. 
Insects after all, exceed the floristic diversity by several orders of magnitude. 

Asanexample of the enhancements offered by the new procedures, within 
a period of 5 years the PTC studies in insect herbivory have collected, sorted, 
and mounted over 100,000 specimens of leaf-chewing and sap-sucking insects 
representing ca. 1,300 species (cf. Basset et al. 2000; Novotny et al. 1997). Such 
outputs considerably exceed those obtained by conventional efforts without 
parataxonomist assistance. Over 40 scholarly papers have been published by 
the research team, in stark contrast to the normal downtime between study 
inception and publication (usually 4.6 years) for the kind of eco-entomological 
inquiry being undertaken by PTC researchers (Erwin 1995). These improve- 
ments are a direct result of the use of parataxonomists in time-intensive ac- 
tions such as collecting and sorting, allowing other participants to optimize 
their own activities on cost-effective schedules. With the marked increases in 
sampling outputs, new insights have emerged which could only have arisen 
from statistically large datasets, such as are now being generated by the new 
protocols. The PTC sampling program has led toa reevaluation of insect-plant 
relationships, with wide-ranging implications for understanding the compo- 
nents of invertebrate diversity in tropical systems (Basset et al. 2000; Novotny 
et al. 1997). In an analogous manner, quantitatively boosted floristic surveys 
have the promise of spawning comparable advances in our knowledge of the 
taxonomy and ecology of the Papuasian/Malesian flora. This is especially likely 
when the existing sampling coverage for plants is so erratic and sparse. 

A significant factor in the success of the PTC operation is the fact that instruc- 
tors and students share fulltime residence in a combined laboratory-dormitory 
complex while pursuing common research objectives. Continuous interactions 
between mentor-trainers and parataxonomists instill a sense of fraternity and 
purpose which is not easily replicated by conventional projects, even though 
the latter may otherwise superficially mimic the PTC program structure and 
objectives. Preservation of the social relationships will be crucial to effective 
transferance of the parataxonomist concept to floristic survey. Similar patterns 


FLORISTIC 451 


for success were previously pioneered at the Christensen Research Institute 
(Orsak 1993) and more recently by the Village Development Trust. Botanical 
planners should note the methodological paradigms, particularly the conditions 
contributing to their effectiveness, as the implications for floristic survey are 
both timely and considerable. 

A point worth repeating is that the parataxonomist concept has been thus 
far applied primarily by entomologists. Unfortunately the botanical profession 
has been slow to recognize that many problems of biological sampling in the 
tropics are universal, and applicability of successful techniques is likely to cut 
across disciplinary lines. Especially with traditional cultures such as PNG, folk 
knowledge of plants is often more extensive than the corresponding base for 
insects (Basset et al. 2000) so plants are actually very appropriate subjects for 
parataxonomist-assisted investigation (Novotny pers. comm. 

o ce eal Pala obeny programs are aitempred within PNG, 

ts will be required in the facilities associated with 
biological en Development of local capacities for floristic survey 
is unlikely to achieve lasting results if the physical security of collections (and other 
survey products such as databases) cannot be assured inside the host country. 
This can be a problem in developing societies where funding priorities for sci- 
ence are generally low. A permanent institutional base will also be needed, and 
is best achieved through the development of organizations specifically devoted 
to parataxonomy, rather than by placing tsin preexisting herbaria 
or government institutions. With scientific facilities in the public domain, there 
is likely to be an administrative bias favoring professional staff over individuals 
without formal credentials. The continuity of parataxonomists in such envi- 
ronments would be less secure than in a mission-specific unit such as PTC. 

Steenis’s estimate of a 50-year cycle of coordinated exploration is opera- 
tionally impossible. In its reliance on outdated concepts of how such inventories 
should be achieved, it is also pecoueaubic with the social realities of contem- 
porary Papuasia. Existing funding f unlikely to support such extended 
programs of deferred realization anyway. As long as limiting factors devolve 
exclusively upon a small number of highly trained professionals, whether in- 
digenous or foreign, it is doubtful that real progress will be made toward the 
goals implicit in comprehensive survey. Only by significant expansion of the 
workforce, like that afforded by parataxonomy, will the inventory process be 
able to encompass the diversity within Papuasian forest ecosystems. Method- 
ologies which improve existing rates of floristic documentation are especially 
urgent in view of the habitat destruction which has occurred in Malesia since 
the time of van Steenis (cf. inter alios Kiew 1990). 

Future surveys should include teams composed of purpose-trained 
parataxonomists. Participation by landowners will also permit outputs to be 
amplified across the board, resulting in survey yields substantially higher than 


452 BRIT.ORG/SIDA 19(3) 


conventional expeditions. A collateral advantage of community involvement 
isthe associated opportunity for integrating traditional knowledge systems into 
the collections documentation. Ethnobotanical inquiry can be easily assimi- 
lated when local inhabitants are engaged in surveys. 

Although there is clearly a general failure of floristic documentation in 
Papuasia, little attention has been explicitly devoted to the way field operations 
are actually conducted. Yet that should be the logical starting point in any analy- 
sis, because how collections are acquired and the limitations associated with 
their acquisition, cannot help but affect everything else which follows. In any 
such examination, probably the most obvious limiting factor which would 
emerge is the ineffectiveness of existing collecting methods. 

In the early days of the PNG Forest Service, rifle fire was often used to bring 
down fertile branches from the canopy. Nowadays, in a country where high- 
powered firearms require special permits, this is no longer a viable option for a 
number of reasons, not least of which being that possession of such weapons 
would attract undesirable attention to the collecting teams. In order to obtain 
specimens from high canopies, local climbers are thus employed on nearly all 
surveys. Selected trees are also frequently cut down. Both methods are very time 


consuming however, and it is not unusual for a single collection obtained by 
such means to take a half hour or even longer. Other procedures using sling- 
shots, extensible poles, wire saws, etc. are useful only in certain situations, and 
also require a substantial amount of practice before the field assistants can 
achieve reasonable proficiency. 

The search for suitable gatherings is often lengthy just in itself. Especially 
in mature growth, where the forest biomass and collection targets are located 
far above the ground, few taxa will be within easy reach. Under prevailing con- 
ditions, a collector with several assistants can expect to obtain an average of 
only 30 taxa per day. Daily tallies tend to be higher in regrowth and montane 
vegetation because of their lower statures, but generally a botanist will not take 
more than 50 numbers even under favorable circumstances. Add to these con- 
siderations the fact that collectors in logistically difficult environments are 
often burdened by institutional quotas for multiple duplicates, and the time/ 
cost demands are increased even further. 

Efforts diverted to the preparation of duplicate samples detract from the 
documentation process. The international herbaria with significant traditions 
in Papuasian botany are few innumber, soa point of diminishing returns is quickly 
reached when distributing specimens. Collections consisting of numerous du- 
plicates are very inefficient in terms of the costs in obtaining those duplicates. 
Floristic inventory is better served by securing small sets G duplicates) of dif- 
ferent conspecific numbers rather than by obtaining single numbers with many 
duplicates. On the former procedure population variation can be effectively 
assessed, while the latter procedure contributes little. 


FLORISTIC 453 


The reasons for the inadequate documentation of the Papuasian flora can 
be entirely understood at the most immediate and basic operational level: that 
of the individual collector laboring in the bush. No matter how much individual 
effort is expended, the per capita outputs are not going to increase to an extent 
necessary to reverse the current trends in botanical inventory. Since personal 
yields are not amenable to improvement, the common sense alternative is to ex- 
pand the workforce. The most practical and socially realistic means of achieving 
this in traditional societies is through the eee of parataxonomists. 

An unfortunate fact 0 veysis that f assisted 
outputs in themselves cannot achieve all : een pee Even when survey 
collections attain respectable volumes, many past efforts have suffered from the 
myopic attitude that the botanical gatherings are an end in themselves. 
Oftentimes there has been no attempt to disseminate findings, or even toassemble 
the results into any kind of usable form. These omissions discourage rational- 
ization of resource management within the surveyed areas, which in practical 
terms is probably the most important downstream product from biosurveys. 

Perhaps the best example of the preceding circumstance is provided by 
current developments in the April-Salumei region of East Sepik Province. No 
other classical locality in Papuasia is of such critical historical-biotic value, 
owing to the fact that nearly all of Ledermann’s interior sites from the 1912-13 
Kaiserin-Augusta (Sepik) Expedition fall within this tract (cf. Veldkamp et al. 
1988). Although key localities in the Hunstein Range were revisited by CSIRO 
botanists Hoogland and Craven in 1966, and by a National Geographic sponsored 
contingent in 1989, there is still no compilation of surviving specimens from the 
now mostly-destroyed Ledermann sets, nor any published compendium from 
the subsequent Hunstein expeditions! In the meantime, scores of significant 
discoveries have been recognized from the newer surveys, including 6 species 
of Freycinetia from the 1989 expedition alone (Huynh 1999). The recovery of 
the endemic genus Sepikea, formerly known only from an illustration in 
Schlechter (1923), has also occurred in recent years (B.L. Burtt, pers. comm.). 
These developments go unnoticed by resource planners because the discoveries 
are reported in technical journals which are inaccessible to government agencies 
or are discussed only within a small circle of botanical collaborators. The April- 
Salumei tract is currently a focal point of contention between conservation, 
landowner, logging, and mining interests (cf. Bakker 1994; Filer & Sekhran 1998), 
and a variety of future land uses is now under planning consideration. The re- 
sults of uninformed action ina locality with such unique biotic and scientific- 
historical values are potentially devastating. 

It is thus an imperative that liaising mechanisms be erected for ensuring 
that surveys connect directly with the agencies responsible for priority-setting 


'There is an unpublished report (Sohmer et al. 1991) but it contains many misidentifications. 


454 BRIT.ORG/SIDA 19(3) 


in wilderness territories. It is not sufficient to establish links between universities, 
herbaria, or NGOs with community-level jurisdictions over survey sites. These 
entities do not possess the statutory powers for determining policies and priorities 
in the resource operations (e.g., forestry and mining) which have the greatest 
potential impacts on the environment. Without critical inputs to the planning 
facilities in government, surveys may end up as mere information-gathering 
exercises from habitats which subsequently disappear. At a minimum, floristic 
inventories should be consciously directed to the Forest Planning 
Division of the PNG Forest Authority, and to the Nature Conservation Division 
of the Office of Environment and Conservation. 


THE RELATIONSHIP TO COMMUNITY AND FLORISTIC PATTERNS 


In the last decade, a substantial amount of data has been acquired from many 
of PNG’s forest environments. The greater part of this work has been conducted 
under the auspices of NGOs operating within their respective conservation areas. 
A discouraging aspect of many such studies is that they are either never for- 
mally published, or otherwise appear in publications of limited readership and 
distribution. In the following discussion, several distinct but interrelated issues 
are considered, in some cases drawing upon data which are available locally, 
but not readily accessible to the wider scientific community. The commentary 
addresses selected topics in 1) morphospecies enumeration, 2) floristic rich- 
ness on environmental gradients, and 3) the relationship between collections 
in Lae Herbarium to timber concessional activity. 

1) Morphospecies enumeration.—A|though the size of its flora is of con- 
siderable general interest, there is little consensus on the number of plant species 
within Papuasia (cf. Collins et al. 1991; Frodin 1984; Good 1960; Hoft 1992; Johns 
1993; Womersley 1978). The only sure way of gauging the total floristic inven- 
tory for PNG is through systematic revision, but it will be many generations 
before a Flora Malesiana-style compilation can be concluded (Geesink 1990). 

In spite of such concerns, the locality-specific inventories are the most 
important ones for planning purposes. While flora-wide summaries may be of 
broad conceptual interest, they do not provide the sort of information which is 
relevant at operational levels. Management actions are typically evaluated and 
implemented for specific localities, and this requires detailed information on 
the floristic profiles of individual tracts. Even if many plant families will re- 
main unrevised through the foreseeable future, the enumeration of Papuasian 
morphospecies and the determination of their spatial distributions is an achiev- 
able objective. Knowledge of the local components of floristic diversity is a ba- 
sis for rationalized priority-setting and decision-making at the level of agency 
implementation. And ultimately it is the conservation of local diversity which 
is the foundation for future floristic inquiry even though the larger regional 
patterns provide the basis for placing the local knowledge in context. 


=“ 


FLORISTIC 455 


Species checklists are thus of greater practical significance for planning 
and conservation than is generally conceded, if they are used in conjunction 
with other information sources. However, unless the compilations accurately 
reflect current taxonomy and are periodically reviewed and adjusted, they can be 
an actual disservice to government planners and managers. In this connection, 
attempts to automate the process of plant identification through interactive 
keys have promise as a heuristic and practical tool, although the methodology 
is likely to be constrained by limited access to 
tries such as PNG 

2. Floristic richness on environmental gradients.—The total number of 
Papuasian plant morphospecies enclosed within individual territories, on 
elevational or horizontal environmental gradients, is also still unknown. Recent 
intensive surveys at Crater Mt. have produced the highest single-locality census 
thus far achieved in PNG (ie. 1,200 mot pl Lospec ies: Takeuchi 1999, 2000b) but the 
counts are not comprehensive. Inventories intending to evaluate total floristic 
content within environmentally variable tracts must eventually quantify species 
richness between habitats. A problem for any manager working with limited 
resources is the question of how to apportion survey effort through time and 
space toa targeted flora. 

In the Neotropics the answer to this issue would be relatively straightfor- 
ward. The number of tree species is inversely related to elevation, with the high- 
est counts in the lowlands and with richness falling progressively with altitude 
(Gentry 1988). When the emphasis is on evaluating the tree flora (as is usually the 
case with forestry operations), surveys would produce maximized returns by 
concentrating at low elevations. A salient qualification however, is that 
neotropical richness seems to be highest in the montane zone (cf. Henderson et 
al. 1991) if all plants (including nontree species) are considered, so the optimal 
sampling plan for surveys would be dependent on the objectives. 

In Papuasian habitats floristic richness patterns are more equivocal and 
complex than those reported by Gentry (1988) and are generally consistent with 
Henderson et al. (1991) if the nontree component is considered. On New Ireland, 
tree counts have been reported as declining monotonically with elevation in 
the manner of the Neotropics (Takeuchi @ Wiakabu 1997). However this result 
has not been replicated and may be an artefact of widely separated Sampling 
stations. Foster (1997), by employing a continuous line pli 
concluded that the species richness curve for New Ireland hasa anid: elevation 
bulge, being highest at 750 m and diminishing above and below that level. 
Kulang et al. (1997) reported maximum species totals at 1,000 m elevation along 
an altitudinal sequence in Madang Province. In a similar study, species diver- 
sity from forest habitats on New Britain attained maximum values between 
600-800 m (Balun et al. 1996). Previously, the 1995 Bismarck-Ramu survey had 
determined the 600 m level as being floristically richest among examined sites 


aes poe 
P resources WI1thin coun- 


g methodology, 


456 BRIT.ORG/SIDA 19(3) 


(Hedemark et al. 1997). Although the lowland-montane ecotone has not been 
critically evaluated in Papuasia, there are converging indications that the point 
of highest floristic development lies somewhere in or near that transition. An 
obvious implication for local conservation initiatives is that this interval should 
thus serve as the botanical core for protected areas. Future surveys can increase 
their efficiency by allocating more time to the low montane ecotone instead of 
attempting equal coverage of all habitats on an altitudinal gradient. The lowland 
rainforest and the high montane forests are apparently less diverse, so survey 
effort should be allocated accordingly. 

The substantial variation (600-1,000 m) in the elevations of beta diversity 
maxima suggests influence of local factors. From the differences between sites it 
is also apparent that the richest communities can be identified only by actual 
survey, and not by ee oes from ou epiane in other areas. Site-spe- 
cific vegetational histories, cli Massernerhebung 
(cf. Grubb & Stevens 1985), substrate distinctions, sampling methodology, etc., 
are factors probably responsible for the contrasts between locations. 

Even while the argument from raw numbers seems clear enough, the situ- 
ation is still obscured by considerations of quality. Plant species do not have 
the same value, at least from conservation perspectives. The local endemic is 
understandably valued more than widely distributed taxa, so richness is only 
one aspect of site assessment. Due to difficulties in generating the required data, 
there are no published accounts comparing endemism between different 
Papuasian environments. This is an obvious lacuna which should be considered 
in future inventories. Although the unrevised status of many plant families isa 
serious constraint, preliminary estimates could be obtained by using the taxa 
covered in modern treatments. 

It is also appropriate to note that while previous efforts have attempted to 
quantify relationships between beta diversity and elevation, the critical variable 
is not elevation but the forest classification unit. In whatever manner the forest / 
community type is defined, whether by species composition, physiognomy, or 
a combination of both, it is really the forest type which underpins the richness 
variation. Elevation isan obvious environmental control but its effects are mani- 
fested through the vegetation formation or ‘life zone, and it is certainly not the 
only controlling factor. Knowing which forest formations have the highest number 
of species, in both absolute and proportional terms, is thus a more meaningful 
focus for inquiry than any site-specific relationship between richness and el- 
evation per se. This is especially true because the same species and/or vegetation 
types often have different elevational ranges at different locations, which is at 
least partly responsible for the inconsistent results obtained by current inves- 
tigations on elevation-dependent richness. On anecdotal grounds, it is apparent 
for example, that many plant taxa have anomalous low-elevational occurrences 


TAKEUCHI FLORISTIC 457 


on ultrabasics. There are also the Massernerhebung induced permutations, to 
cite another obvious influence (cf. the ‘melange effect’ in Grubb & Stevens 1985). 

Establishing correlations between forest units and floristic diversity is com- 
plicated by the existence of several contrasting systems for forest classification 
in Papuasia (i.e, Hammermaster & Saunders 1995; Johns 1977; Paijmans 1975, 
1976; Saunders 1993). The first scheme however, is now supported by a GIS (Geo- 
graphic Information System) augmented by transparency overlays for the pre- 
existing 1:500,000 scale topographic maps (Australian Survey Corps) and also 
with separately issued 1:100,000 scale maps showing the various forest units 
for all of PNG. It thus provides a very useful foundation for planning and execut- 
ing the surveys needed to resolve outstanding issues. 

3. The relationship between collections in Lae Herbarium to timber 
concessional activity.—Recent summaries (Balgooy et al. 1996; Welzen 1997) 
from the Flora Malesiana suggest that New Guinea has the highest rates of flo- 
ristic endemism in Malesia. Based on current revisions, most of the endemic 
species appear to be concentrated in montane habitats. Using selected genera, 
Heads (in press) arrives at some of the same conclusions regarding relation- 
ships between montane environments and endemism. 

Collection densities in Papuasia are very clearly skewed in favor of mon- 
tane areas (Conn 1994), so to a certain extent phytogeographic summaries will 
be affected by the sampling inequalities. The greater part of the LAE holdings 
(and thus the overseas duplicates resulting from them) originate from the 
Mamose? region and the Highlands (cf. Fig. 1), so it is inevitable that a certain 
bias has been introduced into distributional summaries derived from such a 
foundation. The magnitude of such biases can be inferred from the geographi- 
cal unevenness of collections in the Lae Herbarium, as summarized in Table 1. 

The core of the PNG national collections in botany consists of the NGF 
and LAE series specimens (1-49,999 on the New Guinea Force numeration and 
thereafter with higher numbers on the LAE sequence). The institutional sets 
currently end at ca. 85,000. Both the NGF and LAE sequences have many blank 
intervals consisting of number blocks which were assigned to past collectors 
but never actually used. A substantial amount of material was also rejected or 
destroyed after the collections had been recorded. Back-numbering of contem- 
porary gatherings has been employed in an attempt to [ill in these gaps but 
there are still much fewer collections than indicated by the institutional num- 
ber. Table lis thus based on a manual count of archived labels physically repre- 
sented by corresponding specimens in the herbarium. 

The NGF and LAE series constitute approximately 25% of the estimated 
275,000-300,000 specimens in the national herbarium, though it should be 


-Mamose region is the admini i i d of West Sepik, East Sepik, Madang, and Morobe Provinces. 


MANUS 2 © we 
— ea a o 
= ss. ae & 
eee SS eC 
WEST “ 2 : . 
. ? NEW IRELAND Na 6 
et X 


Islands Region 
cee |) 


an 
ot 
) 
C 
ie 


J ) 
vo. lag 
eee ae — NEW BRITAIN sew enean,_ \ 


f ® J : e 
e ( SOUTHERN EASTERN \ 
) HIGHLANDS (_HIGHLANDS MOROBE | BOUGAINVILLE 
‘ ee 
a > 
ral 
WESTERN 
\ we 
») 
SS Papua }- 
~ me 7 Sy 
a te ae Q NW 
ft 
a) 
ae Pa a= 
MILNE BAY 
on 
sort ao 
= 
Sal 
oO 
=] 
ae royce Oran ts new Guinea. mney tern Highland posed nga in pe western nearly tern Highlands P str. in the eastern part. In similar fashion the s 
Chiml Cth) IC ut strT Uiakhl 1. | g I gi 1sffinie: } Arictrall } than = 
. . © 
to the Island Arc terranes (Pigram & Davies 1987). io) 


TAKEUCHI AND GOLMAN, FLORISTIC 459 


Taste 1. Representative profile of plant collections at Lae Herbarium tabulated by province and 
region of origin (from NGF, LAE, and CSIRO series numbers). 


Province Counts % of Total Sample 
HIGHLANDS REGION 
Chimbu 8,060 8.86 
Eastern Highlands 5,837 6.42 
Enga 38 0.04 
Western Highlands 3844 423 
Subtotal Highlands Region 17,779 19.6 
ISLANDS REGION 
East New Britain 1,701 1.87 
Manus 896 0.98 
New Ireland 1,418 1.56 
North Solomons (Bougainville) 2,475 Zi 
West New Britain 4104 45] 
Subtotal Islands Region 10,594 11.6 
MAMOSE REGION 
East Sepik 3,396 oe 
Madang 3,485 3.83 
Morobe 25,509 28.04 
West Sepik 6,082 6.69 
Subtotal Mamose Region 38,472 42.3 
PAPUAN REGION 
Central 8,262 9.08 
Milne Bay 5,404 5.94 
Northern 1,520 1.6/7 
Subtotal Papuan Peninsula 15,186 16.7 
Gulf 1,666 1.83 
Western 3,691 4.06 
Southern Highlands 3,569 3.92 
Subtotal Papuan Austrocraton 8,926 9.8 
Total 90,957 100.0 


noted that the size of the total holdings is not accurately known and tends to be 
overstated because of discontinuities in the recording system. A major data re- 
covery effort would be required to collate locality information on most of the 
remaining collections. The notable exceptions are the early CSIRO sets obtained 
by L. Craven, T. Hartley, P. Heyligers, R. Hoogland, K. Paijmans, and R. Schodde. 
The LAE duplicates by these collectors are entered in herbarium logbooks and 
have been easily incorporated into Table 1. Approximately 91,000 specimen 
numbers are included in the combined tallies (Table 1). 


460 BRIT.ORG/SIDA 19(3) 


PERCENT 
Oo 


percent of region in 
timber concessions 


percent LAE collections 
from region 


percent LAE collections 
from Morobe Province 


iSLANDS PAPUAN PAPUAN HIGHLAND 
AUSTROCRATON PENINSULA 
Fic? Dal . | L ! 4 . . J +l lati Pe Pe eee sg Lhice 4 TIAL Ti re 
F Fr é JJ J 7 
Th + ¢t = rd AA L D . - } ae | rar lo¢ £ 
Mamose region. In general, the highest proportion of timber i i ith the least botanical d 
mentation, tl | traini lanni 1 


7 be J t J J 


The obvious preponderance of specimens from Morobe and adjacent prov- 
inces, and the opposing paucity of material from Western and Gulf, are apparent 
even from casual inspection of the national collections. With 28% of the collec- 
tions, Morobe Province is very disproportionately represented. The disparities 
clearly reflect the concentration of Forest Service facilities and infrastructure 
at Lae and Bulolo. A similar situation is shown by the relatively high numbers 
of specimens from Central Province, especially from the Brown River and the 
Sogeri areas, owing to their proximity to Port Moresby. 

If timber concessions are tabulated by province and region (Table 2),a general 
inverse relationship is evident between the distribution of existing concessions 
and the collections coverage of the corresponding areas (Fig. 2). The provinces 
most susceptible to logging impacts (Islands region and the Austro-geoprovince) 
have the fewest herbarium specimens as reflected in the LAE sample, while the 
best documented regions (Mamose and Highlands) have relatively little logging 
activity. This inverse relationship shows that future forest sector development 
is likely to be most intense in areas which are floristically the least known, and 


TAWCIICU FLORISTIC ” 


TABLE 2. Summary of timber concessional in Papua N bulated | d 
Province Total Concessional Area (ha) % of Province in Concessions 
HIGHLANDS REGION 
Chimbu 0 0.0% 
Eastern Highlands 0 0.0% 
nga 43 483 3.7% 
Western Highlands 83,129 9.8% 
Subtotal Highlands Region 126,612 3.4% 
ISLANDS REGION 
East New Britain 577,287 38.2% 
Manus 51,/34 24.6% 
New Ireland 564,631 58.7% 
North Solomons (Bougainville) 101,120 10.8% 
West New Britain 1,552,628 74.8% 
Subtotal Islands Region 2,847,400 50.0% 
MAMOSE REGION 
East Sepik 630,949 14.4% 
Madang 387,870 13.5% 
Morobe 2/6,/51 8.2% 
West Sepik 681,255 18.9% 
Subtotal Mamose Region 1,976,825 13.9% 
PAPUAN REGION 
Central 484778 16.2% 
Milne Bay 225,101 15.8% 
Northern 358,096 15.9% 
Subtotal Papuan Peninsula 1,067,975 16.0% 
Gulf 2,536,478 74.9% 
Southern Highlands 39,241 1.5% 
Western 2,028,312 20.7% 
Subtotal Papuan Austrocraton 4,604,031 29.2% 


may thus involve considerable biodiversity risk. 

Documentation action is especially imperative for localities where con- 
cessional activity is currently occurring or imminent (cf. Fig. 3, reproduced from 
Papua New Guinea Forest Authority 1998). Floristic data is required not only to 
rationalize the logging plans for affected areas, but also to assess the postharvest 
consequences of forest felling. The opportunity for discovering localized species 
and of recording populational variation could be otherwise irretrievably lost. 

Judging from historical patterns of collecting and future forestry needs, the 
highest-priority documentation targets should be the lowlands of Papua and the 


BRIT.ORG/SIDA 19(3) 


I N\ we 
: ; 
. \ fala 


Fic. 3.M p 


463 


FLORISTIC 


ss 
wy 
0 100 200 
Kilometers 
i 5 
1 
N' 
1949 
| 
| 
9) 
go 
5 ue ° ar 
| CV ee ~ 
NS NN J 
ea ° 
SIN en ‘a gon 
oe aS, \e a“ 
Ae ae or % f } 
ft os 
ee pe qa =O 
> 
Os, aie 
aan 


464 BRIT.ORG/SIDA 19(3) 


Austrocraton subregion, particularly Gulf Province. This does not necessarily 
mean that such environments are more speciose or significant, only that the 
flora there is comparatively less explored and at greater risk of alteration before 
it has been documented. Based on Fig. 3, New Britain should also be a priority 
target, but much of the island has been previously logged and taxonomic losses 
have presumably already occurred asa result of forest removal (oil palm plan- 
tations now cover large sections of West New Britain). In contrast, Gulf envi- 
ronments are primarily in natural growth. 


DISCUSSION 


Papuasian bioinventories involving expeditions and contingents of highly 
trained specialists are relicts from a bygone era. Institutional and social realities 
within a rapidly evolving PNG indicate the appropriateness for change, even 
though the manifestations of such need are obscured by the misconceptions 
imposed by international and cultural distance. Existing failures in documen- 
tation are certainly not attributable to a lack of collective scientific interest or 
dedication of past workers. The deficiencies are principally methodological. The 
iueies oe the last te years Me that conventional itineraries are not going 

t ti tion of critical environments within accept- 
able time frames. As long as the burdens of inventory are borne primarily by 
an elite professional corps, the documentation of PNG’s biotic richness will con- 
tinue to be an elusive objective. The human assets for survey must be applied in 
more effective ways than previous programs or the deficiencies will persist. 
Unfortunately time is running out. 

Past attempts at floristic inventory have been institutionally centralized, 
expeditionary, brief, logistically intensive, and with participation by a select 
membership. Future operations will need to become decentralized, continuous, 
participatory, and effected primarily by personnel that are preferably actually 
living at the sites being subjected to bioevaluation. Parataxonomists and the in- 
frastructure to support them, are necessary elements in achi 
The new schedules will also have to be acutely responsive to emerging grass- 
roots assertion of community ownership rights and the resulting demands for 
stakeholder participation in all activities involving customary resources. Un- 
less there is a major rethinking and overhaul of existing strategies, the objec- 
tives outlined by van Steenis half a century ago are unlikely to be realized. 


onrcamesc 


ACKNOWLEDGMENTS 


The senior author thanks Vojtech Novotny (Institute of Entomology, Czech 
Academy of Sciences) for providing access to grant proposals and supporting 
documents. Many suggestions made by Dr. Novotny were incorporated into the 
final draft. Michael Heads (University of Goroka) also shared unpublished in- 


FLORISTIC 465 


formation. Assistant Director Barney Lipscomb (Botanical Research Institute 
of Texas) extended his usual support and assistance. Colleague Hitofumi Abe 
(Ecosystem Research Group, University of Western Australia) provided the 
Japanese translation. The PNG National Forest Authority gave permission for 
reproducing data from Service files. 

We acknowledge referees PF. Stevens (Missouri Botanical Garden) and J. 
Pipoly II (Fairchild Tropical Garden) for their characteristically thorough cor- 
rection of the manuscript. 


REFERENCES 


Bakker, E. 1994. Return to Hunstein forest. Natl. Geogr. Mag. Feb:40-63. 

Batcooy, M.M.J. van, P. Hovenkamp, and P. van Weizen. 1996. Phytogeography of the Pacific - 
floristic and historical distribution patterns in plants. In: A. Keast and S. Miller, eds. The 
origin and evolution of Pacific island biotas: New Guinea to Eastern Polynesia: pat- 
terns and processes. SPB Academic Press, Amsterdam, the Netherlands. Pp 191-214. 

Bavun, L., Emrik, and L. Orsak. 1996 (unpublished). A study on plant species diversity and 
Spatial patterns in rain forest Communities from Sulka area in New Britain Island, Papua 
New Guinea. Pacific Heritage Foundation. 

Basser, Y., V. Novotny, S.E. Mitter, and R. Pyte. 2000. Quantifying biodiversity: experience with 
parataxonomists and digital photography in Papua New Guinea and Guyana. 
BioScience 50:899-908. 

Beever, B., ed. 1993. Papua New Guinea conservation needs assessment report, 2. PNG 
Dept. of Environment and Conservation, Boroko. 

Cottins, N.M.,J.A. Saver, and T.C. WHitmore, eds. 1991.The conservation atlas of tropical forests, 
Asia and the Pacific. Macmillan Press, London. 

Conn, B.J. 1994. Documentation of the flora of New Guinea. In: C.-l Peng and C.H. Chou, 
eds. Biodiversity and terrestrial ecosystems. Institute of Botany, Academia Sinica Mono- 
graph 14:123-156. 

Crocomee, R. 1974. An approach to the analysis of land tenure systems. In: H. Lundsgaard, 
ed. Land tenure in Oceania. ASAO Monograph 2, University Press of Hawaii, Honolulu. 

Damas, K. 1998. The present status of plant conservation in Papua New Guinea. In: C.-| 
Peng and PP. Lowry Il, eds. Rare, threatened, and endangered floras of Asia and the 
Pacific rim. Institute of Botany, Academia Sinica Monograph 16:171-179. 

Erwin, T.L. 1995. Measuring arthropod biodiversity in the tropical forest canopy. In: M.D. 
Lowman and N.M.Nadkarni, eds. Forest canopies. Academic Press, San Diego. Pp 109-127. 

Ficer,C. 1995. The nature of the human threat to Papua New Guinea's biodiversity endow- 
ment. In:N.Sekhran and S. Miller, eds. Papua New Guinea country study on biological 
diversity. Colorcraft Ltd, Hong Kong. Pp 187-199. 

Ficer, C. and N. SexHran. 1998. Loggers, donors and resource owners. Papua New Guinea 
National Research Institute, and International Institute for Environment and Develop- 
ment, Monograph 34, London. 


466 BRIT.ORG/SIDA 19(3) 


Foster, R. 1997 (unpublished). The forest vegetation. |n:B. Beehler,ed.A biodiversity assess 
ment of southern New Ireland, Papua New Guinea. Conservation International, Wash- 
ington, DC. Pp 46-68. 

Frooin, D.G. 1984. Guide to standard floras of the world. Cambridge University Press, 
Cambridge. 

Geesink, R. 1990.The general progress of Flora Malesiana.|n:P Baas, K. Kalkman,and R.Geesink, 
eds. The plant diversity of Malesia, proceedings of the Flora Malesiana symposium 
commemorating Prof.Dr.C.G.G.J.van Steenis. Kluwer Academic Publishers, Dordrecht, 
Boston, and London. Pp 11-16. 

Gentry, A. 1988. Changes in plant community diversity and floristic Composition on envi- 
ronmental and geographical gradients. Ann. Missouri Bot. Gard. 75:1-34. 

Goop, R.1960.On the geographical relationships of the angiosperm flora of New Guinea. 
Bull. British Mus. Nat. Hist. Bot. 2:205-226. 

Gruss, PJ.and PF. Stevens. 1985. The forests of the Fatima Basin and Mt. Kerigomna, Papua 
New Guinea: with a review of montane and subalpine rainforests in Papuasia. Australian 
Nat. Univ./Anutech, Dept. Biogeogr. & Geomorph., Research School of Pacific Studies, 
Publ. BG/5, Canberra. 

Gumol, M. and N. SekHRAN. 1995. An overview of the Papua New Guinean economy: the 
implications for conservation. |In:N.Sekhran and S. Miller,eds. Papua New Guinea country 
study on biological diversity. Colorcraft Ltd, Hong Kong. Pp 41-57. 

Hammermaster, E.T. and J.C. Saunpers. 1995. Forest resources and vegetation mapping of 
Papua New Guinea. PNGRIS Publ. 4. CSIRO and AIDAB, Canberra. 

Heaps, M.J.(in press). Regional patterns of biodiversity in New Guinea plants. Bot.J.Linn.Soc. 

Heoemark, M., S. HAMILTON, and W. TakeucHi. 1997. Report on the first Bismmarck-Ramu biological 
survey with sociological and logistical comments. PNG Dept.of Environment and Con- 
servation, Port Moresby. 

Henperson, A.,S.P. CHurcuitt,and J.L.Luteyn. 1991.Neotropical plant diversity: Are the northern 
Andes richer than the Amazon Basin? Nature 351:21-22 

Hort, R. 1992. Plants of New Guinea and the Solomon Islands. Dictionary of the genera 
and families of flowering plants and ferns. Wau Ecology Institute Handbook 13. 

HotzknecHr, H. 1995, Papua New Guinea's land tenure, land use and biodiversity conserva- 
tion. In: N. Sekhran and S. Miller, eds. Papua New Guinea country study on biological 
diversity. Colorcraft Ltd, Hong Kong. Pp 59-66. 

Huynu, K.L. 1999. The genus Freycinetia (Pandanaceae) in New Guinea (part 2). Bot. Jahrb. 
Syst. 121:149-186. 

Huynx, K.L. 2000. The genus Freycinetia (Pandanaceae) in New Guinea (part 3). Candollea 
55:299-322. 

JANZEN, D.H., W. HALLWACHS, J. JIMENEZ, and R. Gamez. 1993. The role of the parataxonomists, 
inventory managers, and taxonomists in Costa Rica's national biodiversity inventory. 
In:W.V. Reid, S.A. Laird, C.A. Meyer, R.Gamez, A. Sittenfeld, D.H. Janzen, M.A. Gollin, and C. 
Juma, eds. Biodiversity prospecting: using generic resources for sustainable develop- 
ment. World Resources Institute, Washington. Pp 223-254. 


FLORISTIC 467 


JOHNS, R.J.1977.The vegetation of Papua New Guinea. Part 1:An introduction to the veg- 
etation. PNG Office of Forests (reprinted 1984). 

JoHNs, R.J.1993. Biodiversity and conservation of the native flora of Papua New Guinea. |n: 
B. Beehler, ed. Papua New Guinea conservation needs assessment report, vol. 2. PNG 
Dept. of Environment and Conservation, Boroko. Pp 15-75. 

Kiew, R. 1990. Conservation of plants in Malaysia. In: P. Baas, K. Kalkman, and R. Geesink, eds. 
The plant diversity of Malesia, proceedings of the Flora Malesiana symposium com- 
memorating Prof. Dr.C.G.GJ. van Steenis. Kluwer Academic Publishers, Dordrecht, Boston, 
and London. Pp 313-322. 

KULANG, J., O. Gesia, and L. BALUN. 1997 (Unpublished). A study on the biological diversity of 
the Hagahai area in the Madang Province, Papua New Guinea. 

Levert, M.and A. Bata. 1995. Agriculture in Papua New Guinea. In: N. Sekhran and S. Miller, 
eds. Papua New Guinea country study on biological diversity. Colorcraft Ltd, Hong 
Kong. Pp 125-153. 

Louman, B.and S. NicHoLts. 1995. Forestry in Papua New Guinea. In:N. Sekhran and S. Miller, 
eds. Papua New Guinea country study on biological diversity. Colorcraft Ltd, Hong 
Kong. Pp 155-167. 

Mcavpine,J.and J.Quictey. 1998. Forest resources of Papua New Guinea. Summary statistics 
from the forest inventory mapping (FIM) system. Coffey MPW Pty Ltd for the Austra- 
lian Agency for International Development and the PNG National Forest Service. 

Novotny, V., Y. Basset, S. Miter, A. ALLISON, G.A. SAMUELSON, and L. Orsak. 1997. The diversity of 
tropical insect herbivores:an approach to collaborative international research in Papua 
New Guinea. Proceedings of the International Conference on Taxonomy and 
Biodiversity Conservation in East Asia. Korean Inst.for Biodiversity Research of Chonbuk 
National University (KIBIO) series 2:112-125. 

Orsak, L.J. 1993. Killing butterflies to save butterflies: a tool for tropical forest conservation 
in Papua New Guinea. News Lepidopterists Soc. 1993:71-80. 

Paumans, K. 1975. Explanatory notes to the vegetation map of Papua New Guinea. Land 
Research Series 35, CSIRO, Melbourne 

PAuUMANS, K., ed. 1976. New Guinea vegetation. CSIRO and Australian National University 
Press, Canberra. 

Papua New Guined Forest AUTHORITY. 1998. Forest resource acquisition general information. 
PNG Forest Authority. 

Picram, C.J.and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea 
orogen. J. Austr. Geol. Geoph. 10:193-211. 

Sautel, S.M. and J.-A. Eitis, eds. 1998. The Motupore Conference: ICAD practitioners’ views 
from the field. A report of the presentations of the second ICAD conference. Motupore 
Island (UPNG), Papua New Guinea 1-5 September, 1997. Dept. of Environment and 
Conservation, Papua New Guinea/United Nations Development Programme PNG/93/ 
G31 Biodiversity Conservation and Resource Management. 

Saunoers, J.C. 1993. Forest resources of Papua New Guinea. Explanatory notes to map. 
PNGRIS Publ. 2, CSIRO and AIDAB, Canberra. 


468 BRIT.ORG/SIDA 19(3) 


SCHLECHTER, R. 1923. Gesneriaceae Papuanae. Bot. Jahrb. Syst. 58:255-379. 

SeKHRAN, N. and S. Miter, eds. 1995. Papua New Guinea country study on biological diversity. 
Colorcraft Ltd, Hong Kong. 

Soumer, S.H., R. Kiapranis, A. ALuson, and W. TakeucHi. 1991 (unpublished). Report on the 
Hunstein River expedition-1989. 


STeeNIs, C.G.G.J. vAN. 1950. Desiderata for future exploration. In: MJ.van Steenis-Kruseman, 
Malaysian plant collectors and collections. Flora Malesiana ser.|, 1:cvii-cxvi, maps 2 and 
3. Noordhoff/Kolff., Djakarta (reprinted 1985, Koeltz, Koenigstein) 

STEVENS, PF. 1989. New Guinea. In: D.G. Campbell and H.D. Hammond, eds. Floristic inven- 
tory of tropical countries: the status of plant systematics, collections, and vegetation, 
plus recommendations for the future. New York Bot. Gard., New York. Pp 120-132. 

Suzuki, D. 1993. Time for a change. Allen and Unwin, St. Leonards. 

TAKEUCHI, W. 1999. New plants from Crater Mt., Papua New Guinea, and an annotated check- 
list of the species. Sida 18:961-1006. 

TAKEUCHI, W. 2000a. A floristic and ethnobotanical account of the Josephstaal Forest Man- 
agement Agreement Area, Papua New Guinea. Sida 19:1-63. 

TakeucHi, W. 2000b. Additions to the flora of Crater Mt., Papua New Guinea. Sida 19:237-247. 

TAKEUCHI, W. 2001. New and noteworthy plants from recent botanical surveys in Papua 
New Guinea, 7. Edinb. J. Bot. 58:159-172. 

TaKeucHi, W. and J.Wiakasu. 1997 (unpublished). A transect-based floristic reconnaissance 
of southern New Ireland. In: B. Beehler, ed. A biodiversity assessment of southern New 
Ireland, Papua New Guinea. Conservation International, Washington, DC. Pp 69-81. 

VeoKAmP, J.F,W. Vink, and D.G. Fron. 1988. X1.Ledermann’s and some other German localities 
in Papua New Guinea. Flora Malesiana Bull. 10:32-38. 

WeELZEN, PC. VAN. 1997. Increased speciation in New Guinea:tectonic causes? In:J. Dransfield, 
M.J.E. Coode, and D.A. Simpson, eds. Plant diversity in Malesia Ill. Proceedings of 
the Third International Flora Malesiana Symposium 1995. Royal Botanic Garden, Kew. 
Pp 363-387, 

Wowersiey, J.S., ed. 1978. Handbooks of the flora of Papua New Guinea 1. Melbourne 
University Press. 


A LEAF BLADE ANATOMICAL SURVEY OF MUHLENBERGIA 
(POACEAE: MUHLENBERGIINAE) 


Paul M. Peterson Yolanda Herrera-Arrieta 
Department of Botany CHDIR Unidad Durango 
National Museum of Natural History COFAA-Instituto ies Nacional 
Smithsonian Institution Apartado Postal 57 
Washington, DC 20560-0166, U.S.A. Durango, Dgo,, ae MEXICO 
ABSTRACT 


Muhlenbergia includes 151 species of mostly New World origin; 133 species are indigenous to North 
America [although many of these range to Central America (33) and South America (14)k 38 species 
occur in sae einen a single species is endemic); 25 species occur in South America (10 are 

own i occur in n southern Asia. No modern subgeneric 


classification within the genus exists and speci é are not clear. An anatomical survey 
of the a Plage as viewed in Hate Y oie section n has pruned a waa set Bot 16 characters to test 


a J 
LOTITICQ 


on all but three species of Prem Based on this al Muhlenbergia appears to be divis- 
ae into oe aie anatomical ae colsespendine to two subgenera (M. subg. Muhlenbergia, 
oa)an ct | Podosemum) in M. subg. Trichochloa. Even 


ee ae presence et sclerosed ce isan » important apomorpby in the evolution of pon es in 
Muhlenbergia subg. Trichochloa, it ar ther species. 
Our study suggests that in Mahlenbemia subg. Muhlenbergia the C, photosynthesis, ia aoe. 


was a single evolutionary event since these species occur as a clade or an uninterrupted grade in our 


phylogenetic analysis. 


RESUMEN 


Muhlenbergia incluye 151 especies originarias principalmente del Nuevo Mundo; 133 especies son 
egan a distribuirse hasta Centro y Sudamérica 


pee) 
w 
ay 
— 


nativas de Norteamérica [aunque muchas 
33 y 14 respectivamente)|, 38 especies habitan en Centroamérica (sola una especie es endémica); 25 
especies se encuentran en Sudamérica (10 son endémicas); y solamente seis especies endémicas se 
distribuyen por el sur de Asia. No existe una clasificacion sungenence meseins del ae y las 


relaciones entre las especies no son claras 
versal ie ie a un See unico de - caracteres para probar las Pelnciones hipotéticas 


previas tos 16 caracteres de todas (excepto tres) las 
especies de Muhlen bergia es base a este andalisis Muhlenber ei 1 parece ser divisible en tres pens 
1 | (M.s Mu eee y Tr ichochloa) y dos 


secciones (M. sec. Epicampes y Podosemum) en nM. subg. Tic ociiae Aun cuando la presencia de 
floema esclerosado es una apomorfia importante en la evolucion de las especies de Muhlen bergia 
euDeen ero loa, parece aie ma ev VeCLOnadO reas veces ve que se » presenta en cuatro especies. 
CK en las especies 


del subgénero Mullenbertia fué un event luti , ya que estas dope es aparecen como 
un clado o un grado ininterrumpido en nuestro analisis sees 


SIDA 19(3): 469 — 506. 2001 


470 BRIT.ORG/SIDA 19(3) 


The subtribe Muhlenbergiinae (Poaceae: Chloridoideae: Eragrostideae) was first 
circumscribed by Pilger (1956) to include only species of Muhlenbergia Schreb. 
with narrow single-flowered spikelets, firm glumes often shorter than the 
awned lemmas, and cylindrical caryopses. In this same treatment Pilger recog- 
nized Epicampes J. Presl [>Muhlenbergia subg. Trichochloa A. Gray, M. sect. 
Epicampes (J). Presl) Soderstr] in subtribe Sporobolinae Ohwi. Pilger further di- 
vided Muhlenbergia into eight sections: Acroxis (Trin.) Bush, Bealia (Scribn.) 
Pilg., Cinnastrum CE. Fourn.) Pilg, Clomena (P. Beauv.) Pilg, Muhlenbergia, 
Podosemum (Desv.) Pilg., Pseudosporobolus Parodi, and Stenocladium (Trin.) 
Bush,. Subsequent authors have agreed that Pilger’s infrageneric treatment of 
Muhlenbergia was not particularly phylogenetically informative (Soderstrom 
1967: Pohl 1969; Morden 1985; Peterson and Annable 1991). More recently the 
following six genera have been shown to share common ancestry and have been 
placed in the Muhlenbergiinae: Bealia Scribn., Blepharoneuron Nash, 
Chaboissaea E. Fourn., Lycurus Kunth, Muhlenbergia, and Pereilema J. Presl 
(Duvall et al. 1994, Peterson 2000; Peterson et al. 1995, 1997). 

Many agrostologists have erected segregate genera to emphasize critical 
features of the large and diverse genus, Muhlenbergia. Desvaux (1810) recog- 
nized the genus Podosemum, based on the caespitose, open-panicled, and long- 
awned M. capillaris. Palisot de Beauvois (1812) described the genus Clomena 
based on the annual M. peruviana, and Presl (1830) described Epicampes based 
on M. robusta. Two relatives of the type species of the genus (M. schreberi), M. 
glomerata and M. andina, were given generic status by Link (1833) as 
Dactylogramma and by Thurber (1863) as Vaseya, respectively. Nuttall 1848) 
described the genus Calycodon based on the widespread and often important 
range grass, M. montana. The only other generic name given to a species pres- 
ently placed in Muhlenbergia is Crypsinna, described by Fournier (1886) and 
based on M. macroura. Hitchcock’s (1935) transfer of many of these segregate 
genera to Muhlenbergia has been followed by most American and European 
botanists. The morphological characters that delimit the genus are spikelets 
with single perfect florets and hyaline or membranous lemmas with three usu- 
ally prominent veins. These characters are not at all unique within the Eragro- 
stideae and seem to be possessed by about half of the genera in the tribe. 

The morphological diversity within Muhlenbergia is tremendous. Annuals 
lessthan 2 cm tall (M. depauperata, M. minutissima, M. peruviana, M. ramulosa) 
are not uncommon and there are numerous strongly caespitose perennials over 
2 m tall (M. gigantea, M. mutica, M. robusta). Rhizomes and/or stolons are found 
in 1/4 of the species and there is a single species (M. dumosa) that has a growth 
form similar to bamboos. Leaf blades can be flat, involute, or folded with a variety 
of pubescence types located on the abaxial and/or adaxial surface. All species of 
Muhlenbergia have open or contracted (spike-like) panicles with the branches 


=~ 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 471 


generally re-branched. At maturity or anthesis, the angle of the branches 
spreading from the culm axis and the total width of the inflorescence are diag- 
nostic characteristics used to separate the species. Pedicel orientation can vary 
from appressed or spreading, to nodding and reflexed from the branches, and 
the pedicels can be either round or flattened in cross section. Most species of 
Muhlenbergia have single-flowered spikelets although there are two species that 
are occasionally 2 or 3-flowered (M. asperifolia and M. uniflora). The lemma is 
perhaps the most critical structure, and its features such as length, presence or 
absence of an awn or mucro, pubescence type and location, shape, and color 
can all be used to differentiate among the species. The single lemma is 3-veined 
(1-veined in M. palmirensis) witha stout central vein and two lateral veins, although 
the lateral veins are sometimes very hard to discern witha good (20X) dissecting 
microscope. The caryopsis has a fused pericarp and is usually free from both 
the lemma and palea in most species of Muhlenbergia, however the length, shape 
and to lesser extent color are highly variable. 

At last tally, Muhlenbergia consisted of 151 species (Peterson 2000). The 
distribution of Muhlenbergia is almost entirely New World where 133 species 
are indigenous to North America [although many of these range to Central 
America (33) and South America (14)|, 38 species occur in Central America (a 
single species is endemic); 25 species occur in South America (10 are endemic): 
and only six endemic species are known to occur in southern Asia. One obvious 
hypothesis is that the genus arose where it is most diverse today, i.e., northern 
Mexico/southwestern U.S., and has since radiated. For a dispersal event, the 
longer the distance from the place of origin, would in theory, lessen the chance 
of a successful introduction. Therefore, there are many species of Muhlenbergia 
in North America, fewer in Central America, even fewer in South America, and 
finally very few in Asia. So far, all ten species in the subtribe Muhlenbergiinae 
that have been investigated genetically (Peterson and Herrera A. 1995; Peterson 
and Morrone 1998; Peterson and Ortiz-Diaz 1998; Peterson et al. 1993: Sykes et 
al. 1997) exhibit a north to south migration pattern, including Muhlenbergia 
torreyi (Peterson and Ortiz-Diaz 1998). 

All species of Muhlenbergia previously examined exhibit kranz (C4) leaf 
anatomy, particularly the parenchyma sheath subtype which is common in 
species pying the most arid regions (Brown 1977; Hattersley 1984; Hattersley 
and Watson 1992). Two main subtypes, NAD-ME (nicotinamide adenine dinucle- 
otide co-factor malic enzyme) and PCK (phosphoenolpyruvate carboxykinase) 
have been found, verified by biochemical assay, to occur within Muhlenbergia 
(Gutierrez et al. 1974; Hattersley and Browning 1981; Brown 1977; Hattersley 
and Watson 1976, 1992). These two biochemical subtypes differ in their predomi- 
nant C4 acid that is transported from primary carbon assimilation tissue (usually 
the mesophyll) to the photosynthetic carbon reduction (PCR) tissue (kranz 


472 BRIT.ORG/SIDA 19(3 


> 


sheath = parenchyma bundle sheath) [see Hattersley and Watson 1992], There 
usually is an associated anatomical structure in PCK-like species that is diag- 
nostic, such as, a looser arrangement of chlorenchyma tissue continuous between 
adjacent vascular bundles. In typical NAD-ME species the chlorenchyma is 
tightly radiate and usually separated from one vascular bundle to the next by a 
column of colorless cells. These differences have historically been used to separate 
these two subtypes; however, it has been shown in Enneapogon Desv. ex P. Beauv, 
Eragrostis Wolf, Eriachne R. Br, Panicum L., Pheidochloa S. T. Blake, Triodia R. 
Br, and Triraphis R. Br. that these anatomically PCK-like genera are actually 
biochemically NAD-ME (Ohsugi et al. 1982; Prendergast et al. 1986, 1987). 

The first major anatomical investigation of Muhlenbergia was done by Holm 
(1901) who looked at 10 species and was able discern three groups: woodland types 
(=M.subg. Muhlenbergia), dry, ey) mountain slopes [= M. subg. Trichochloa, sect. 
Epicampesand other species} and M, filipes (= M. subg. Trichochloa,sect. Podosemum). 
Holm (1901) pointed out that from an anatomical view-point these characteris- 
tics might prove useful in dividing Muhlenbergia into sections or subgenera. 

Schwabe (1948) later investigated 22 species of Muhlenbergia that occur 
in South America and found that they correspond to four major groups: hygro- 
phytes or mesic species (= M. subg. Muhlenbergia), xerophytic annuals one, 
xerophytic annuals two (= M. subg. Muhlenbergia), xerophytic perennials, and 
psammophytic perennials (= M. subg. Trichochloa, sect. Podosemum). Schwabe 
(1948) also suggested that Muhlenbergia should be separated from the genera 
of Agrostideae and incorporated into the Eragrosteae. 

On the basis of leaf blade transectional anatomy and morphology 
Soderstrom (1967) distinguished two subgenera in Muhlenbergia, Muhlenbergia 
and Podosemum (= Trichochloa, an older name), and divided M. subg. 
Trichlochloa into two sections, sect. Podosemumand sect. Epicampes. Soderstrom 
placed 46 species of Muhlenbergia, which have partially sclerosed phloem and 
caps of sclerenchyma associated with the primary vascular bundles, into M. 
subg. Trichochloa. Muhlenbergia sect. Epicampes was characterized as having 
a compound keel (midvein) composed of primary and secondary vascular 
bundles sunken in a confluent mass of thick-walled parenchyma, whereas M. 
sect. Podosemum had a simple midvein composed of a single primary vascular 
bundle with additional tertiary vascular bundles present (Soderstom 1967). 

Two years later Pohl (1969) completed a revision of 12 closely related species 
that he believed to represent the entire M.subg. Muhlenbergia in North America. 
Principal differences among the species in this study were size of the bulliform 
cells, size and degree of radial orientation of the chlorenchyma, and the extent 
to which the chlorenchyma was organized into discrete units surrounding each 
vascular bundle. Using morphological characteristics of the rhizome (posses- 
sion of very short internodes with imbricate scales) and leaf blade (thin, flat 


to 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 473 


blades with low length/width ratios), Pohl distinguished these species from 
others in the genus. However, these same characteristics are seen in M. 
californica, a species of the mountains and valleys of southern California, and 
many other species common to the southwestern United States/Mexico. 

Morden and Hatch (1987, 1996) investigated the anatomical and morpho- 
logical variation within the M. repens complex, which consists of six species in 
North and South America. Anatomical data supported the placement of M. 
squarvosa (Trin.) Rydb. as. a synonym of M. richardsonis, and supported the rec- 
ognition of two varieties of M. villiflora. 

Peterson et al. (1989) and Peterson and Annable (1991) investigated 29 annual 
species of Muhlenbergia and found 14 characters useful in distinguishing four 
major species groups. Species of group (1) had flat blades, two tertiary vascular 
bundles between primary vascular bundles, vascular bundles positioned on 
the median layer, distinctly radiate and compact chlorenchyma cells separating 
adjacent vascular bundles, and fan-shaped central bulliform cells. Species of 
groups (2, 3 & 4) shared three characteristics: 1) indistinctly or incompletely 
radiate and loosely arranged chlorenchyma cells, 2) chlorenchyma cells con- 
tinuous between vascular bundles, and 3) shield-shaped central bulliform cells 
(= M. subg. Muhlenbergia). 

More recently a biosystematic study investigating the anatomy of the M. 
montana complex (consisting of 15 species) has been completed (Herrera- 
Arrieta 1998; Herrera-Arrieta and Grant 1994). Herrera-Arrieta and Grant used 
18 characters to differentiate among these species and found four major species 
groups. Important characters appear to be the central midrib structure (similar 
in size to the other primary vascular bundles or presence of a prominent central 
midvein), the depth of the adaxial and abaxial furrows, sclerenchymatous girder 
development between the parenchyma bundle sheath and the epidermis, and 
epidermal vestiture (glabrous or papillose). 

Leaf anatomical characters within the Poaceae as viewed in transverse sec- 
tion have long been recognized as important diagnostic features used to deter- 
mine ic relationships, and have been critical in elucidating infrageneric 
relationships within Muhlenbergia (Herrera-Arrieta and Grant 1993; Holm 1901: 
Morden and Hatch 1987; Peterson 2000; Peterson and Annable 1991: Peterson et al. 
1989; Pohl 1969; Soderstrom 1967). A preliminary summary of our anatomical 
analysis is presented in Peterson (2000) where two subgenera, M. subg. 
Muhlenbergia and subg. Trichochloa, and a possible third group, ‘Clomena’ are 
recognized. In this current paper we will give a detailed summary of the ana- 
tomical features as viewed in cross section of 148 of the possible 151 species 
within Muhlenbergia and present a subgeneric hypothesis that most closely 
reflects a cladistic analysis of the data. This is the first anatomical survey of 
nearly all species within this large variable genus. 


474 BRIT.ORG/SIDA 19(3) 


MATERIALS AND METHODS 


Over the last 16 years fresh field-collected leaf blades were obtained from North, 
Central, and South America, as well as China for anatomical study (Appendix 
1). Five mm long leaf blade segments from the central third of the mid-culm 
region were fixed in FAA (10 parts EtOH; 1 part glacial acetic acid; 2 parts 37% 
formaldehyde; 7 parts distilled water). A few species (less than 5%) were studied 
from dried herbarium specimens because fresh field collected material was 
unavailable. Leaf blades were first desilicified in 100% hydrofluoric acid (HF) 
for 48 hours in order to ease microtomy, then dehydrated using 30, 50, 70, 90, 
95, and 100% (twice) ethanol, graded into xylene (twice) and transferred to 
xylene: paraffin oil (1:1, steps 1 hour minimum). Blades were then dehydrated 
by using 2-2 dimethoxypropane (DMP), acetone, and tertiary butyl alcohol 
(TBA) series while in a vacuum. Infiltration was accomplished using two, six 
hour minimum changes of liquid paraffin before being embedded. The tissue 
was softened using 95% EtOH: Glycerin: HF (8:1:1) to improve slicing (Foster 
and Gifford 1947). A standard rotary microtome set at 6-10 :m thickness was 
used and sections were stained with safranin/fast green or 0.05% toluidine blue 
(Berlyn and Miksche 1976). Samples were examined and photographed on an 
Olympus BH-2 photomicroscope using Kodak TMAX black and white or 
Ektachrome color slide film. 

Anatomical descriptions were completed following the procedure for stan- 
dardizing comparative leaf anatomy in grasses as outlined by Ellis (1976). For 
purposes of comparison and standardization, primary vascular bundles (1°) are 
defined as containing large metaxylem elements on either side of the protoxy- 
lem elements with additional lysigenous cavities, and are usually associated 
with sclerenchyma girders or strands (Ellis 1976; Peterson et al. 1989). Secondary 
vascular bundles (II°) resemble 1° vascular bundles by having distinguishable 
xylem and phloem but lack large metaxylem elements and lysigenous cavitities. 
Tertiary vascular bundles (111°) contain indistinguishable xylem and phloem 
areas and are usually smaller than the I° and II° vascular bundles. 

A list of the anatomical characters and their states used in all ensuing analyses 
appears in Table 1; a list of specimens used in this study is given in Appendix 1, 
and a complete data set is given in Appendix 2. The 153 taxon by 16 character 
data set was analyzed with WinClad2000. Parsimony heuristic analysis was 
performed with NONA (Goloboff 1998; Nixon 1999). We used 10 random taxon 
order replications in NONA, with TBR swapping holding 20 trees, followed by 
TBR swapping (to completion) holding up to 200 trees. A hard collapse of all 
unsupported nodes was selected to produce the cladogram in Figure 17. Since 
we were not testing for monophyly of the genus, the outgroup species were con- 
strained, i.e., there were no synapomorphies supporting the Muhlenbergia clade. 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 475 


Taste 1. List of anatomical characters used in the cladistic analyses, their states, and comments. 


1. 


Adaxial furrow depth: 1 = <1/5 blade thickness; 2 = 1/5-2/5 blade thickness; 3 = 1/2 or more than blade average 
thickness. 


2. Primary oes bundle shape; l= = rounded, = obovate /ellipti 4= rectangular The overall outline is used to 


w 


> 


LF] 


[oy) 


co 


‘oO 


_—= 


—_ 
—_ 


— 
Ww 


_ 
wn 


_= 
n 


: Central bulliform cells shape: ee or rregular- shaped, 2= almost fan- “shaped to shield- nape, een 


determ mine shape. Th VdSCUld? DUNIE. 


. Vascular bundle outline size: | = = primary vascular bundles about the same size lary and tertiary 


vascular bundles; 2 = subequal, secondary and tertiary vascular bundles 4/5 the size of primary vascular bundles; 3 = 
unequal, secondary and tertiary vascular bundles <2/3 the size of primary vascular bundles. 


. Median (keel) vascular bundle structure: | = simple keel, with only a single primary vascular bundle; 2 = com- 


pound keel, a single primary vascular bundle with only two additional tertiary vascular bundles; 3 = complex com- 
pound keel, with three or more additional primary, secondary and/or tertiary vascular bundles, In state 1 there are no 
associated parenchyma cells, whereas state 2 and 3 


. Vascular bundle position: 1 = one level, centered at same level from adaxial to abaxial ane 2 = two or three 


levels, usually closer to the abax'al surface 


. Vascular bundle composition: 1 = with primary and secondary vascular bundles; 2 = with primary and tertiary 


vascular bundles; 3 = with primary, secondary, and tertiary vascular eum A primary vascular bundle (I) olan: 
ws y more large metaxylem gual on ae side of the Pl toxy| 
habl secondary vascular bundle (II°) resembles a |° vasc ular bundle by having 
distinguishable xylem and phloem but He large metaxylem elements and lysigenous cavities. A tertiary vascular 
bundle (Ill?) contains indistinguishable xylem and phloem areas and is usually smaller than a |° and/or II° vascular 
ndle. 


. Chlorenchyma arrangement: | = radiate, compact; i loosely arranged. state | va with C, NAD-ME 


- tate 2 corresponds with C, 


i} 


p 
PCK species where the chlorenchyma i aces ah 


. Crown of inflated cells (adaxial) i in primary vascular bundles: | = absent; 2= eth inflated areas are 


usually composed of parenchyma or collenchyma cells.In addition there may be sclerenchyma cells. 


re is 
cal umn (contiguous between the adaxial and abaxial surface) of bulliform/colorless cells separating each ‘deen 
al bundle. 


1 / . I 1 i bons 4° i. 1 n i] 


0. 1h Number y / er lal y Vastulal t J 


= ]-3;2 = 4ormore. 


. Median vascular bundle structure: 1 = not differentiated from other primary vascular bundles; 2 = differentiated 


from other primary vascular bundles 


. Median vascular bundle (abaxial) projection: | = flattened, not enlarged; 2 = enlarged, bulbous with many 


strands of sclerenchyma. 


. Sclerosed phloem in primary vascular bundles: | = absent; 2 = present. State 2 is characterized by strands of 
sclerenchyma cells that divides the phloem 
4, C;l kL {ad any he I + 


Bnd ea ba 


vascular bundles: ] = absentt af W TIDETS, ne or two 


t 7 


layers; 3 = three or more layers. 


. Sclerenchyma (abaxial) development in the primary vascular bundles: | = one or more layers, continuous 


along the width . blade; 2 = one or more layers, discontinuous, only present below the vascular bundles; 3 = 
absent to a few 


. Inflated cells (abaxial) in primary vascular bundles: | = present; i= absent. Lh e le areas are usually 


composed of parenchyma or collenchyma cells. In addition there may his character is similar to 
number 8 but found on the abaxial surface. 


476 BRIT.ORG/SIDA 19(3) 


Fics. 1-6. Leaf blade anatomy of Muh/enbergia, adaxial surface upp t in all photographs, except in 3 where the 
lade is involute 1 ope “sh CLO ] vlig {-V/el £+h:-] \ chall n a | . L 
vascular bundles, | ly ged chl hyma, only primary and tertiary lar bundles t and shield-shaped 
bulliform cells. 2. M. pauciflora showing a flat lami di laxial f (1/5 — 1/3 leaf thickness), rounded pri- 

mary vascular bundle shape Jaxial rit pp i ll | I dles i | ial gi | f scl I bet 


n di + ale | ] | J nhl | kL £ I id L Oe | 


each vascular bundle, vascular | y y 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 477 


All autapomorphies were not included in the final cladogram because they add 
no additional information for inferring relationships among two or more taxa. 


RESULTS AND DISCUSSION 


The results and discussion are interpreted in two parts: 1) a general description 
combining all species of Muhlenbergia, and 2) results of the cladistic analysis. 

General Description of Leaf Structure.—Lamina (blades) are sometimes 
undulating, to more commonly flat, outwardly bowed, less commonly invo- 
lute (Fig. 3), or folded (Fig. 4.). The angle formed by the twoarms is broadly V or 
U shaped to expanded, occasionally loosely involute. Adaxial furrow depth in 
comparison to the leaf thickness can be slight, shallow (< 1/5 leaf thickness, 
Figs. 1, 11, 16), medium (1/5 to 1/3 leaf thickness, Figs. 2, 13), or deep (1/2 leaf 
thickness, Figs. 3, 10), and in the form of clefts located between all vascular 
bundles. Adaxial ribs are commonly present opposite all vascular bundles (Fig. 
2), the same size to generally smaller than abaxial ribs, to less frequently absent 
with shallow groves opposite the vascular bundles (Fig. l). Primary vascular 
bundle shape varies from rounded (Figs. 2, 4), to obovate /elliptic (Figs. 3, 6, 10) 
or rectangular (Figs. 5, 13, 14); secondary and tertiary vascular bundles also ex- 
hibit the same variation in shape. The secondary and tertiary vascular bundles 
are generally of the same size as the primary vascular bundles (Fig. 4), to about 
4/5 the size of the primary vascular bundles (Fig. 5), or very unequal, less that 
2/3 the size of the primary vascular bundles (Fig. 6). Abaxial projection of the 
median vascular bundle or midrib caused by sclerenchyma is sometimes large 
and with a protruding ridge (Fig. 7), to inconspicuous, often flat to round (Fig. 4). 


and adaxial fibers, and intereostal SCTE NGIyING: = M. dubia with an involute lamina, deep adaxial furrows (1/2 or 
more eMan blade thickness), ptic vascular bundle shape, ate crown of inflated cells abaxial inflated 
cells, le | i | i bundle, 2/3 the eof 
Bi y Fr J VYQOULUIdTI , ? WQORUIGE VULIMICS iy iit size Mi 
the primary bundles, with Jary and tertiary vascular bundles p 1 cl to the abaxial surface, 15 total 
1 L TT i kl rT ra 4h rT y J . y | L 4] L L p ig y L ij ,one or 
+ } £ ras L at | | dA 1 Fy 1 +1 hI J 14 4. M. brevivaginata 


VU ba TO Ul y y 


itt 1 ascul Iles al I i ne keel, rounded vascular bundles, an undiffer- 
entated median vascular bundle, centered vasculal bundle and : asa onde pet eG 5.M. pangiuma with 


7 
| L 4] hat L ° L Jle rectanqular I n 1] n 4 | L 1 anc 
U be J pcan” J 7 
L nl 


pe, 
| lart Jle, sclerosed phloem 
I I 

: : ; " 


kK 
yma, 
sheath extensions, one ort layers of adaxial scl k Jeveloy he p y 


| at . k 7 6.M.nig | . |} Jalliney lock rT 


—— and d secondary vascular bundles that are  <2/3 the length and width of the primary vascular bundles, tightly 
radia adaxial crown not intiated cals abarlal ioiated oy ane secondary and tertiary vencilet 


} bullif 
1UT 


| Il; 
renchyma, Ivb = pummel) vascular bundle; ! es = secondly vascular bundle; Ill es = feptiaey vascular binale mx = 


d yl NI, p ph i, p t y ATT nchyr Hd, op 


478 BRIT.ORG/SIDA 19(3) 


eS i 
6 


oe 


a8: - 


8:49 »* 


Figs. 7-12. fhilnd £ AAiphlantk . me | £. . IT nk L 7_M let ° ra 


be et ae” wee | hy teh | 4 1 J | } Tl + medium vas- 


> 
= 
5 
q 
. 
~ 


Pv imary vascular 
more primary and seontaly nesaulee bundle and abaniel sclerenchyma of two to four discantintious layers. 8. M. 
japennta with an arged proj ones median Uaealet bundle a complex compound keel 


with only pri- 


mary ane tertiary vascular bundles, sonsely altanged clorenchyma, and five tertiary vascular bundles between each 
primary. 9. M. gig more primal Y, See a oe 


| L 4] tal Hee er | } tiahel eee LI L 


1g! 


ing a primary, tertiary, and secondary vascular bundle (left to right), vascular bundles at three sed ade sahil 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 479 


The median vascular bundle or midrib is a simple keel (Fig. 4+) consisting 
of a single vascular bundle without associated parenchyma, a compound keel 
(Fig. 8) consisting of one primary and two secondary or tertiary vascular 
bundles with associated parenchyma, or a complex compound keel (Fig. 9) con- 
sisting of three or more primary, secondary, and/or tertiary vascular bundles 
with associated parenchyma. All vascular bundles are commonly situated with 
their position in the median layer of the blade, at the same distance from the 
adaxial and abaxial leaf surface (Figs. 2, 4,5, 11, 13), or are occasionally closer to 
the adaxial surface (Figs. 3, 6, 10) at two or three levels. Vascular bundle compo- 
sition consists of primary and secondary only in the same blade (Fig. 7), only 
primary and tertiary in the same blade (Figs. 1, 8, 11). The presence of primary, 
secondary, and tertiary vascular bundles combined in one blade is not as com- 
mon (Figs. 3, 6, 10). The total number of primary vascular bundles varies from 
5 to 15 (four in M. fastigiata, nine in M. pauciflora, 15 in M. gigantea). Secondary 
and/or tertiary vascular bundles are arranged in a regular fashion between 
consecutive primary vascular bundles, and the number varies between 1-3 (Figs. 
54.6), or 4-3 Fie.8): 

The chlorenchyma tissue consists of two major types (arrangements). It 
can be composed of a single radiate layer of tightly packed tabular cells that 
surround each vascular bundle [NAD-ME, centripetally positioned photosyn- 
thetic carbon reduction (PCR) cell chloroplasts, XyMS+ and PCR cell outlines 
that are even in transverse section; see Hattersley and Watson 1992] and is sepa- 
rated by uni-, bi- or tri-serial columns of colorless/bulliform cells (Figs. 3-6, 9, 
10, 12-15). Or it can be composed of tabular cells that are indistinctly radiate 
and continuous between the bundles [PCK type, defined as centrifugal/evenly 
distributed PCR cell chloroplasts (with grana), XyMS+ and presence of PCR 
cell wall suberized lamella, in Hattersley and Watson’s (1992) sense] (Figs. 1, 11, 
16). Colorless cells (Figs. 5, 12, 13) are smaller or similar in size and shape to 
bulliform cells and are often inflated. A crown of inflated cells is sometimes 
present over the primary vascular bundles on the adaxial surface and these 
inflated cells can be found over the secondary vascular bundles as well (Figs. 3, 
6, 7, 10, 14). Inflated cells sometimes can be found separating the primary vas- 
cular bundles from the epidermis on the abaxial surface (Figs 3, 6, 10). Strips of 


diate chl hyma, ok /ellipti lar bundle shape, deep adaxial furrows 1/2 or more than the blade ae 
ness | 4 LI - ag 1 1 A gad | Eiuflasad l kh sal tm flanenal Wi 
| f adaxial scl I in the primary lar bundles. 11.M sniene ee 11 arena 
bundles between oan femlaty vascular bundle hall axial f ular bundles, eel etranded 
chon ren chyma, bullif i gle lay of adaxial and abaxi | (di ) 
p | 12 Mc ifoli h radi hl | 1 pt ,anda dalam mn 
lorl II ing tl vascular bundles. rele pas — ay Pa symbols as ionews: a bllifrm cell d= 


vs | hl 
chlorenchyma, letaxy ;p=p ;ps=| y leren hyr Na; sp 


480 BRIT.ORG/SIDA 19(3) 


Fics. 13-16. Leaf blad f Muhlenbergia, adaxial surf. in all pt hs. 13.M. lindheimeri with 
adaxial ae ae es ane bade thickness, rectangular vascular bundle shape, centered vascular Pune non-scle- 
more 


layers si serene, ane one or more alae J discontinuous abaxial scprenenyiia: Las mt eeepansa auth 
oe rbun 


mary vas a dle, abaxial and adaxial i | | bundle sheath, abaxial girder of sclerenchyma 
fibers, circular shaped bulliform cells, three or more layers of adaxial ssotbieai and Haly ee of aes 
abaxial sclerenchyma. 15. M. curvula with non-sclerosed phloem, tightly y baxial and adaxial 
spe baenciyms aun ateat oe aay of Sdetenchym fibers, ee or more layers ee scleren- 
16.M with loosely 


nl 1. £. ere | Lahkauial asl L fil aL 


al 


chym 
nana corenchym, shields hay ; y 
primary bund] } bol foll b = bulliform cell; cl = chl h ,mx = metaxylem; p= 


Meee Hak } J Pr 


well-defined and regular bulliform cells are present in the epidermis and are 
distinct from normal epidermal cells. Bulliform cells can be closely associated 
with the colorless cells. Bulliform and colorless cells together form the uni-, bi-, 
or tri-seriate columns which extend from the adaxial furrow to the abaxial 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 481 


epidermis separating the vascular bundles (Figs. 5, 12, 13), or the columns do 
not extend to the abaxial surface (Fig. 2). The central bulliform cell can be cir- 
cular to fan-shaped (Fig. 14) or narrower than deep, shield-shaped (Figs. 1, 11, 
16). Outer tangential epidermal cell walls are unthickened to slightly thick- 
ened, with cells of similar size. Macrohairs have a sunken, nonconstricted base 
and are embedded between bulliform and/or colorless cells. 

The phloem of the primary vascular bundles can be homogeneous or 
unsclerosed (Figs. 2, 12, 14) or interrupted with sclerenchyma or sclerosed (Figs. 
5, 10, 14) where it adjoins the mestome sheath. Two enlarged metaxylem vessels 
are present adjacent to the phloem and one or two other enlarged protoxylem 
vessels are located adaxially to the phloem (Figs. 2,5, 10, 13). Metaxylem vessels 
are small, not wider than the parenchyma sheath cells, slightly thickened, and 
circular in outline. A mestome sheath surrounds the xylem and phloem. The 
mestome cells are small with uniformly thickened walls in all bundles (Figs. 2, 
5, 10, 13, 14). 

Bundle sheaths in the primary vascular bundles sometimes include ex- 
tensions (Fig. 5) and are entire (form a complete circle) to adaxially interrupted, 
or adaxially and abaxially interrupted (Figs. 14, 15) by a broad girder of a few to 
many sclerenchyma fibers (Figs. 14, 15), or colorless inflated cells (Fig. 14). Sec- 
ondary and tertiary vascular bundle parenchyma sheaths are mostly entire, 
not interrupted, to abaxially interrupted in some species. The median vascular 
bundle parenchyma sheath is mostly abaxially interrupted, to interrupted on 
both sides, or less frequently not interrupted. Commonly 6-21 cells form the 
parenchyma sheath of primary vascular bundles (Figs. 2, 5, 10-12, 15, 16), with 
up to 24 cells found in some species (Fig. 14), while 3-14 cells commonly com- 
prise parenchyma bundle sheaths of secondary (Figs. 5, 10) and tertiary (Figs. 
1, 11, 16) vascular bundles. 

Adaxial and abaxial sclerenchyma development is extremely variable, from 
a few fibers (Fig. 1) to 1-3 layers or strands (Figs. 5, 10, 14). In a few species a 
continuous layer of sclerenchyma can form beneath the epidermis on the 
adaxial or more commonly abaxial surface (Figs. 3, 4, 6, 10). Sclerenchyma is 
usually present along the margins of the blades forming a “cap” that may be 
rounded or pointed (Figs. 3, 4). This sclerenchyma cap adjoins normal meso- 
phyll cells. Sclerenchyma is usually absent between each vascular bundle where 
there are no continuous sclerenchyma layers. However, a few species, M. 
pauciflora (Fig. 2) and M. seatonii, form a conic abaxial girder of intercostal 
sclerenchyma. An abaxial projection of midrib caused by sclerenchyma is some- 
times enlarged and bulbous (Figs. 7, 8). 

Cladistics.—For the overall analysis of 148 species (plus two infraspecific 
taxa) of Muhlenbergia and four outgroup species representing four genera 
(Eragrostis acutiflora, Erioneuron avenaceum, Leptochloa virgata, and 


BRIT.ORG/SIDA 19(3) 


Eragrostis acutiflora 
Erioneuron avenaceum 
Leptochloa virgata 
Spor Spa oe 


M. dumosa 
+— M. glabriflora 
i-— omerata 
M. appres 
M. mone 
M. ciliata 
M. diversiglumis 


a 


-microsperma 
ec 


— M. pe ee 
-——M.a 
1— M. ae 
60 | M. fragilis 
7-1, 9-1, 10-1 -— M. depauperata 


M. crispiseta 
Bi a9 M. eludens 
12-2 15-2 M. sinuosa 
M. texana 


Fic. 17. One of 20 equally parsimonious trees (length = 97 steps, Cl = 0.30; RI = 0.89) analyzing 148 species of 
Muhlenbergia with Eragrostis acutiflora, Erioneuron avenaceum, Leptochloa virgata, and Sporobolus airoides used as 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 483 


M. arizonica 
Zz -— M. fastigiata 


12-1 —- MI. ramulos 


folia 
ee | M. caxamarcensis 
6-1 |__ M. cleefii 
-——~ MI. jonesii 
-— M. palmirensis 


4 
15-2 


jab 
M. villiflo 

M. vill.var. vite 
wrigh 


tii 
M. orophil 
84 
= M. strictior | E 
6-2 M. tenuissima 
M. cuspidata 
M. arenace 
M. flexuosa 
a9 M. arenicola 
14-2 = M. setifolia 
fe M. torreyi 
* -— M. argentea 
15-2 — M. flaviseta 
M. uniflora 
watsonia 
pee MI previvacinats 
-+— M. eriophylla 
82 | -— M. michisensis 
3-2 torreyana 
M. asperifolia 
filiculmis 
plumbea 
31] M. seatonii 
a utilis 
TWh -. M. aguascalientensis 
i__ M. curvula 
+— M. hinton 
+— M. quadridentata 
89 straminea F 
3-3 M. virescens 
ng M. cualensis 
id M. ria ana 


84 ‘ 14-2 sy flabe 
4-3 M.jaime-intoni 
n 


an 
» | 3Lom. voicaloe 
2-4, 13-2 


cE n | 1 L hal ae L n fall 


followed by the 


outgroups N 


state. Groups A-F are discussed in the text. 


BRIT.ORG/SIDA 19(3) 


—— M. mutica 
+—— M. pubigluma 
-—— M. expansa 
}+— M. ag 


igida 
| lm. subaristata 


TM. distchophyla 
}+—_— M. emersleyi 
}+— M. is 
/-— M. eects A 
-— M. iridifolia 


veal 
a) 


M. xanthodas 


Fig. 17 Continued 


CwanyvAnA lit iV | 
P E 


;) were used simultaneously and in all possible binations 
All possible combinations were obtained by changing the order of each 
outgroup listed in the data set and sequentially eliminating one, two or three of 
the outgroups. These 200 trees from the single overall analysis are 97 to 99 steps 
long, with a consistency index (CD of 0.30 and a retention index (RD) of 0.89. 
Twenty of these 200 trees were only 97 steps long and therefore one of these 
was randomly selected for illustration (Fig. 17). As indicated in the methods 
there were no synapomorphies supporting monophyly of the Muhlenbergia 
clade, therefore the outgroup species were constrained. These 16 anatomica 
characters are not, by themselves robust enough to test for monophyly within 
the genera of Muhlenbergiinae, Eragrostideae, or the entire Chloridoideae. 
There is little resolution in the strict consensus tree for the overall analysis 


— 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 485 


(using the 200 trees). However, the strict consensus tree separates a clade (Fig. 
17A) containing M. aurea, M. breviligula, M. distans, M. distichophylla, M. 
emersleyi, M. gigantea, M. grandis, M. inaequalis, M. x involuta, M. iridifolia, M. 
lehmanniana, M. lindheimeri, M. longiglumis, M. longiligula, M. pilosa, M. 
pubescens, M. reederorum, M. robusta, M. scoparia, M. speciosa, and M. xanthodas 
(bootstrap value of 100%); a clade (when rooted with Erioneuron avenaceumin 
a separate analysis at 0.28 Cl and 0.88 RI) or a grade (B), when rooted with 
Eragrostis acutiflora in a separate analysis) containing the following 37 species: 
M. alamosae, M. andina, M. appressa, M. arsenei, M. brandegei, M. bushii, M. 
californica, M. ciliata, M. curtifolia, M. x curtisetosa, M. curviaristata, M. 
diversiglumis, M. dumosa, M. glabriflora, M. glauca, M. glomerata, M. hakoensis, 
M. himalayensis, M. huegelii, M. japonica, M. mexicana var. mexicana, M. 
mexicana var. filiformis, M. microsperma, M. pauciflora, M. pectinata, M. 
polycaulis, M. porteri, M. racemosa, M. ramosa, M. schreberi, M. setarioides, M. 
sobolifera, M. spiciformis, M. sylvatica, M. tenella, M. tenuiflora, M. tenuifolia, 
and M. thurberi (bootstrap value of 100%); and a grade of all other fale in 
the genus. The former oe of a species all appear to be members of M. subg. 
Trichochloa, sect. Epi 1967; Peterson 2000). Two apomorphies 
support a clade (Fig 17, clade A): complex compound — ma anee ¢ or more 
additional primary, secondary and/or tertiary vascular | 4(3) 
and median vascular bundles that are differentiated from other primary vascular 

bundles [character 11(2)]. However, complex compound keels [character 4(3)] 
appear in five additional species in two clades. One of these clades, containing 
M. curviaristata, M. hakoensis, M. himalayensis, and M. japonica, is composed 
only of species endemic to southeast Asia. 

We must point out that M. mexicana var. mexicana and M. mexicana var. 
filiformis occur in two separate clades (Fig. 17, grade A). Both species have identical 
scores for the data set. However, abaxial sclerenchyma development (character 
15) isambiguously scored as having one or more layers (state 2) or three or more 
layers (state 3) for each of these taxa. I am not completely familiar with the al- 
gorithm used in WinClad2000 but it appears that it selects either state 2 or 3 
when reading ambiguous character scores. That would account for the inclu- 
sion of M. mexicana var. filiformis after the node supported by character 15(3), 
whereas M. mexicana var. mexicana was selected as 15(2). This is just one of the 
20 shortest trees and most of the other trees include both varieties of M. 
mexicana in the same clade. 

Three apomorphies of deep adaxial furrows greater than 1/2 the blade 
thickness [character 1(3)], vascular bundles positioned in two or three levels 
[character 5(2)|, and inflated cells located below (abaxial to) the primary vascu- 
lar bundles [character 16(1)] support a core group of 12 species (Fig. 17, clade C) 
that correspond to members of M. subg. Trichochloa, sect. Podosemum (M. 


— 


486 BRIT.ORG/SIDA 19(3) 


angustata, M. dubia, M. gypsophila, M. jaliscana, M. lucida, M. macroura, M. 
mucronata, M. nigra, M. palmeri, M. rigens, M. rigida,and M. subaristata). These 
12 species plus M. articulata, M. capillaris, and M. stricta form a clade charac- 
terized by having vascular bundles composed of primary, secondary, and ter- 
tiary types [character 6(3)]. 

Muhlenbergia capillaris, M. expansa, and M. filipes are problematic since 
in other trees these species comprise a single clade or in separate clades. In 70 of 
the 200 trees these three species form a clade with the 14 previously discussed 
species tentatively placed in M. subg. Trichlochloa, sect. Podosemum. In all other 
trees they are aligned with species of M. subg. Trichochloa in a grade containing 
clades of each section (Epicampes and Podosemum). Therefore, placement in 
either section of M. subg. Trichochloa is premature. Interestingly, these three 
species were treated by Morden and Hatch (1989) as a single species with three 
varieties. Although they can be somewhat difficult to distinguish using gross 
morphological features, there appears to be sufficient differences in habitat, 
flowering time, and anatomical structure to warrant recognition at the species 
level. We believe these three species clearly belong in M. subg. Trichochlod since 
they form a clade with other members of sect. Epicampes and sect. Podosemum by 
possessing a crown of inflated cells just below (abaxial to) the primary vascular 
bundles [character 8(2)]. 

Soderstrom (1967) delineated M. subg. Trichochlod (as M. subg. Podosemum) 
based on possession of sclerosed phloem, caespitose perennial habit with erect, 
usually stout and robust culms, and glumes veinless or l-veined. The single 
apomorphy (symplesiomorphy?) of sclerosed phloem [character 13(2)] appears to 
be an important character aligning at least Ll additional species in our analysis: 
M. elongata, M. jaime-hintonii, M. montana, M. mutica, M. pubigluma, M. 
pungens, M. purpusii, M. reverchonii, M. setifolia, M. versicolor, M. virlettii. All 
these species except M. pungens have the morphological characteristics that 
Soderstrom described for M. subg. Trichochloa. Muhlenbergia pungens is rhi- 
zomatous, decumbent near the base, and short (culms 20-70 cm tall). Based on 
our morphological observations M. pungens appears related to M. arenacea, M. 
arenicola, and M. torreyi. The last two species are the only other taxa in our 
study that have sclerosed phloem [character 13(2)|, however, these two species 
with M. setifolia always form a separate clade. There are no obvious morpho- 
logical characteristics that align M. setifolia with either M. arenicola or M. 
torreyi. Therefore, the evolution of sclerosed phloem within Muhlenbergia ap- 
pears to have occurred twice. Even though M. montana exhibits some individuals 
with sclerosed phloem, others lack this character state. Muhlenbergia montana 
was aligned with M. straminea and M. virescens in about half of the 200 trees 
and therefore should not be included in M. subg. Irichochloa at this time. These 
three densely caespitose species all have 3-veined upper glumes that are usually 


~ 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 487 


3-toothed as well. It seems best to tentatively place these eight species (excluding 
M. montana and M. pungens) in M. subg. Trichochloa without further affinities. 

These 200 trees in the overall analysis appear to support a group (Fig. 17, 
grade B) of 37 species with apomorphies of loosely arranged chlorenchyma[C4 
PCK type; character 7(2)] and fan- to shield-shaped bulliform cells without for- 
mation of a sclerenchyma girder from the adaxial to the abaxial surface [char- 
acter 9(2)]. These 37 species correspond to M. subg. Muhlenbergia. All of these 
species except M. arsenei and M. polycaulis have an additional apomorphy of 
four or more secondary and/or tertiary vascular bundles between consecutive 
primary vascular bundle [Character 10(2)]. However, four species (M. curtifolia, 
M. glauca, M. pauciflora,and M. thurberi) exhibit both states for character 10. 
A homoplasious state in these 37 species is the occurrence of only primary and 
tertiary vascular bundles[character 6(2)] also shared with annual or short-lived 
perennial* (Fig. 17, clades D & E) species (M. annua, M. brevis, M. breviseta*, M. 
capillipes, M. crispiseta, M. depauperata, M. eludens, M. filiformis, M. flavida, 
M. fragilis, M. implicata, M. ligularis*, M. majalcensis, M. minutissima, M. 
peruviana, M. ramulosa, M. schmitzii, M. sinuosa, M. strictior, M. tenuissima, M. 
texana, M. vaginata*). However, 120 of the 200 trees suggested direct descent, 
ie., derived froma single ancestor, for the derivation of this state [char- 
acter 6(2)]. 

The evolution of C4 photosynthesis in grasses is a complicated subject, 
however, it seems clear that the pathway has originated at least four times (Sinha 
and Kellogg 1996) or more (seven or more times in Brown 1977). One of those 
origins appears to be the subfamily Chloridoideae lineage (Renvoize and Clayton 
1992). Our study suggests that in Muhlenbergia the PCK subtype of photosyn- 
thesis was a single evolutionary event [character 7(2)]. Since the occurrence of 
the PCK subtype is found in three of the four outgroup species (Eragrostis acuti- 
flora, Leptochloa virgata, and Sporobolus airoides), it is not surprising that in 
all 200 trees this state appears plesiomorphic when rooted with these species. 
Hattersley and Watson (1992) hypothesized that the PCK subtype evolved from 
NAD-ME since in the C4 acid cycle PCK subtype isan enhancement of the NAD- 
ME subtype, and PCK is only known in grasses and may therefore have evolved 
subsequent to the NAD-ME type which is known in other monocotyledons and 
dicotyledons. Jacobs (1987) earlier suggested that the PCK subtype is perhaps 
primitive since it is found in other groups, ie., Panicoideae, whereas the NAD- 
ME subtype is restricted to Chloridoideae. We agree with Hattersley and 
Watson's t and prefer to view the development of the PCK subtype in 
Muhlenbergia as the derived state. An alternative hypothesis, although this 
would require additional morphological or molecular evidence, might be that 
the PCK species or M. subg. Muhlenbergia actually represent a separate lineage 
and deserves generic status. 


488 BRIT.ORG/SIDA 19(3) 


The remaining 64 taxa (M. aguds dlientensis, M. annua, M. arenacea, M. 
arenicola, M. argentea, M. arizonica, M. asperifolia, M. brevifolia, M. brevis, M. 
breviseta, M. capillipes, M.caxamarcensis, M. cleefii, M. crispiseta, M. cualensis, 
M. curvuld, M. cuspidata, M. d 


lepauperata, M. durangensis, M. eludens, M. 
eriophylla, M. fastigiata, M. filiculmis, M. filiformis, M. flabellata, M. flavida, M. 
flaviseta, M. flexuosa, M. fragilis, M. hintonii, M.implicata, M. jonesii, M. ligularis, 
M. majalcensis, M. michisensis, M. minutissima, M. montana, M. orophila, M. 
palmirensis, M. peruviana, M. plumbea, M. pungens, M. purpusii, M. quadri- 
dentata, M. ramulosa, M. repens, M. richardsonis, M. schmitzii, M. seatonii, M. 
sinuosad, M. sinuosa, M. straminea, M. strictior, M. tenuissima, M. texana, M. 
torreyana, M. torreyi, M. utilis, M. vaginata, M. villiflora var. villiflora, M. villi- 
flora var. villosa, M. virescens, M. watsoniana, and M. wrightii) seem to contain 
sympleisiomorphies, ie., they lack anatomical synapomorphies. These species 
all exhibit radiate, compact chlorenchyma or the classical NAD-ME subtype 
characteristic of many chloridoid grasses [character 7(1)|, contain primary vas- 
cular bundles without sclerosed phloem [character 13(2)], although present in 
M. arenicola, M. pungens, and M. torreyi; have rounded vascular bundles [char- 
acter 2(1)|, although M. torreyana and M. pungens have obovate/elliptic or rect- 
angular bundles; have simple keels [character 4()], although M. torreyana has 
a complex compound keel like species in M. subg. Trichochloa sect. Epicampes; 
and have circular or irregular to fan-shaped bulliform cells [character 9()]. Even 
though the cladistic analysis using these 16 anatomical characters does not sug- 
gest a monophyletic lineage for these 63 species, we prefer to recognize them 
informally as the ‘Clomena’ complex (Peterson 2000). 

Within ‘Clomena’ there exists some resolution, for instance, a clade con- 
taining annual or short-lived perennial species (Fig. 17, clade D) is based on the 
occurrence of primary and secondary vascular bundles [character 6(2)]. Other 
annuals (M. annua, M. brevis, M. crispiseta, M. depauperata, M. eludens, M. 
fragilis, M. minutissima, M. sinuosa, and M. texana) occur asa grade. A grade 
(Fig. 17, grade F) within ‘Clomena’ containing M. aguascalientensis, M. cualensis, 
M. curvula, M. durangensis, M. flabellata, M. hintonii, M. quadridentata, M. 
straminea, and M. virescens, along with the remaining species in the analysis is 
also depicted. These species along with M. argentea, M. crispiseta, M. eriophylla, 
M. filiculmis, M. flaviseta, M. jonesii, M. michisensis, M. montana, M. peruviana, 
and M. watsoniana have been referred to as the Muhlenbergia montana complex 
(Herrera-Arrieta 1998; Herrera-Arrieta and De la Cerda-Lemus 1995; Herrera- 
Arrieta and Grant 1993,1994). This complex consists of highly caespitose species 
usually with a three-veined upper glumes, and in our analysis, leaf blades 
mostly with three or more layers of adaxial sclerenchyma in the primary 
bundles [character 14(3)|. The Muhlenbergia repens complex (Morden 1985, 
1995; Morden and Hatch 1987, 1996) which includes M. fastigiata, M. plumbea, 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 489 


M. repens, M. richardsonis, M. utilis and M. villiflora) is not monophyletic, ie. 
these species are found in two or more clades or as a grade with many addi- 
tional species. 

One of the two least homoplasious characters in the analysis is chloren- 
chyma arrangement (character 7, consistency index = 0.50). All species in M. 
subg. Muhlenbergia appear to be PCK [character 7(2)] whereas all other members 
of the genus are NAD-ME [7(1)]. Only two species are ambiguously scored for 
character seven, M. glauca, more than likely amember of M. subg. Muhlenbergia, 
and M. capillaris, clearly a member of M. subg. Trichochloa. Median vascular 
bundle structure (character 11, CI = 0.50) is the other least homoplasious char- 
acter. Allmembers of M. subg. Trichochloa sect. Epicampes have median vascular 
bundles that are differentiated from other primary vascular bundles [11(2)]. If 
you choose to disregard the outgroup species in the cladistic analysis, then 
within Muhlenbergia there is no homoplasy (CI = 1.00) for these two characters 
(7 & 1D. 

In conclusion, our data support the division of Muhlenbergia into two sub- 
genera (M. subg. Muhlenbergia and Trichochloa) and two sections (M. sect. 
Epicampes and Podosemum) in M. subg. Trichochloa. Preliminary investigations 
of Muhlenbergia and relatives based on internal transcribed spacer region se- 
quences of nuclear ribosomal DNA provide support for a clade containing only 
PCK species (= M. subg. Muhlenbergia) and another clade containing only M. 
subg. Trichochloa (Peterson, Columbus, Cerro Tlatilpa, and Kinney 2001). We 
prefer to view this partial classification based on anatomical characters as a 
work in progress and realize that with additional morphological and molecular 
data our understanding of the evolution of this genus will improve. We feel it is 
important to present this anatomical information since it is the first time the 
entire genus has been surveyed in this manner, therefore this serves as a foun- 
dation for further taxonomic research. 


APPENDIX 1 


Specimens used in this study, all housed at the United States National Herbarium 
(US) unless otherwise indicated. Those marked with an asterisk * appear in the 
figures. Collectors are abbreviated as follows: A=C.R. Annable; AC = S. Acevado:; 
B = S.M. Braxton; C = A. Cortes O.; CA = M.A. Carranza; CV = A. Cam pos- 
Villanueva; D = C.H. Dietrich; DC = M. De la Cerda-Lemus; DU = WC. Dunn: G = 
M.S. Gonzalez-Elizondo; H = Y. Herrera-Arrieta; J = EJ. Judziewicz; K = MB. 
Knowles; KI = R.M. King; L=J. Linkins; LB = R.J. LeBlond:; M =O. Morrone; P = PM. 
Peterson; PO = M_E. Poston; R=N. Refulio-Rodriguez; S$ = RJ. Soreng; V=J. Valdes- 
Reyna; VI=J.A. Villarreal; W = A.S. Weakley. 


APPENDIX 1. 
TAXON COLLECTORS LOCALITY 
M. eae Y. Herrera & H&DC 1185 MEXICO. Aguascalientes: San Jose de Gracia, 12 km NW of La Congoja 
de la Cerda-Lem 
M. alamosae Vasey P&A 8293 MEXICO. Chihuahua: 76 mi W La Junta and 35.2 mi W Tomochic in Parque Nal. 
Cascada de Basaseachic 
PA&V 10807 MEXICO. Chihuahua: Parque Natural Barranca del Cobre, 12.6 mi NE La Bufa and 2 
mi S Basigochi 
M. andina (Nutt.) Hitche. P&A 4982 USA. California: San Benito Co. 9.8 mi SW New Idria along Clear Creek 
M. angustata (J. Presl) Kunth 
PA&PO 8817 ECUADOR. Provincia de Chimborazo: 8.9 km N Palmira on the Panamerican Hwy 


M. annua (Vasey) Swallen 


M. appressa C.O.Goodd. 


M. arenacea (Buckley) Hitchc. 


M. arenicola Buckley 


M. argentea Vasey 
M. arizonica Scribn. 
M. arsenei Hitchc. 
M. articulata Scribn. 


M. asperifolia (Nees & 
eyen ex Trin.) Parodi 


M. aurea Swallen 


P&A 4036, 4053 

P&A 4102 

P&A 4189 

Holmgren & Holmgren 7051 


P&A 5703 
PA&V 10033 
P&A 5521 


PA&V 10032 
PA&H 8044 
P&A 5329 
P&A 5142 


P&K 13386 
P&K 13365 


P&A 4851 
PA&M 10177 
M de Koninck 1954 


MEXICO. Chihuahua: NW of Hernandez Javales 

MEXICO. Durango: Navios 

USA. Arizona: Graham Co. 6.6 mi SW Hwy 366 above jtn. with Hwy 666 
(NY)USA. Arizona: Graham Co., Pinaleno Mts 


USA. Arizona: Cochise Co., Triangle T Road, 1.2 mi E Dragoon 

MEXICO. Coahuila: 29.2 mi S Saltillo on Mex 54 to Concepcion del Oro 

one Arizona: Cochise Co., 10 mi S Rucker Canyon on Tex Canyon Road & 6 mi NE 
80. 


wy 
res Coahuila: 29.2 mi S Saltillo on Mex 54 to Concepcion del Oro 
MEXICO. Chihuahua: 15.3 mi S Mex 127 and 6.9 mi NE La Bufa 
USA. Arizona: Santa Cruz Co. 3.3. mi W Hwy 289 & Pena Blanca 
MEXICO. Baja California: Sierra San Pedro Martir, 1.8 mi S Vallecitos 
MEXICO. San Luis Potosi: 2.5 mi E Hwy 57 on road towards Guadalcazar 
MEXICO. Nuevo Leon: ca 36 mi N Dr. arroyo on Hwy 61 towards Linares 


USA. Oregon: Klamath Co. 1.5 mi S Worden on Hwy 97 
ARGENTINA. Provincia Salta: at km 1137, 26.7 km SE Molinos on Hwy 40 
GUATEMALA. Quetzaltenango: Retalhuleu 


06 


(€)6L VOIS/DYO'LINE 


TAXON COLLECTORS LOCALITY 
M. brandegei C. Reeder Moran 9361 MEXICO. Baja California Sur: Isla Catalina 
P&A 4760 MEXICO. Baja California Sur: |sla La Partida 
PA&V 10811 MEXICO. Chihuahua: 12.6 mi NE of La Bufa and 2 mi S of Basigochi 


M. brevifolia Scribn. ex Beal 
M. breviligula Hitche 
M. brevis C.O.Goodd 


M. breviseta Griseb. ex E. Fourn. 


M. brevivaginata Swallen 


M. bushii R.W. Pohl 

M. californica Vasey 

M. capillaris (Lam.) Trin. 

M. capillipes (M.E. Jones) 
P.M. Peterson & Annable 


M. caxamarcensis Laegaard & 
M. ciliata (Kunth) Trin. 


M. cleefii Laegaard 
M. crispiseta Hitch. 


M. cualensis Y. Herrer & PM. Peterson 


M. curtifolia Scripn. 


M.xcurtisetosa (Scribn.) Bush 
M. curviaristata (Ohwi) Ohwi 
M.curvula Swallen 


A.S. Hitchcock 9063 

P&A 4005 

P&A 4030 

A.S. Muller 1853 

PD,B&K 13396 

PG&K 13660* 

D.M. ec 30573 

P&A 5 

MeL 23 
P&A 

Pp i ie 


P&R 14013 


P&A 4679 
Cleef & Florschutz 5578 


P&A 4063 
P&A 4067 

&A 4103 
Guzman 6090 
P&A 5631 
G.P. Clinton 1892 


T. Koyama 6390 
PG&K 13636 
P 9686* 


GUATEMALA. Guatemala city 
SA.New Mexico: Grant Co., NE of San Lorenzo 
MEXICO. Chihuahua: NW of Hernandez Javales 
MEXICO. Veracruz: Orizaba 
MEXICO, Sinaloa: 3 mi SW Estado de Durango and 2.2 mi S El Palmito on Hwy 40 
MEXICO. Durango: 7.3 mi S of Charcos on road towards San Juan de Michis 
USA. Arkansas: Benton Co., Monte Ne 
USA. California: San Bernardino Co., Mtn. Home Village, along Mtn. Home Creek 
USA. North Carolina: Brunswick Co., Sunset Beach 
MEXICO. Chihuahua: 24 mi SW La Junta and approx. 44 mi N Creel, at P Arroyo 
Ancho crossing 
Chihuahua: 23 mi SW La Junta on road to Creel at the Puente Arroyo Ancho 
PERU. Depto. Cajamarca: Prov. Cajamarca, 18 km W of Central Plaza of Cajamarca 
up road to Cumbemayo Sanchez Vega 
MEXICO. Chiapas: 8.2 mi SE of San Cristobal de las Casas 
COLOMBIA. Boyaca: Sierra Nevada del Cocuy, Alto he Lagunillas 
MEXICO. Chihuahua: 12.1 mi NE of El Vergel on Hw 
MEXICO. Chihuahua: 10.9 mi NE of El Vergel on Hwy - 
MEXICO. Durango: 5.4 mi W of Navios, 42 mi W of Durango 
MEXICO. Jalisco: E of Zimapa 
USA yaaa Cocoino Co,, mae eae. 22.5 mi SE of Fredonia on Forest Service 


Road 422 

eee 

JAPAN. Honshu: Prov. Shinano, Togakushi, 2 km NW of Chusha 
MEXICO. Durango: 30 mi SE Mezquital on road to Charcos 
MEXICO. Guanajuato: 18.5 mi SE San Felipe on Mex 37 to Leon 


VIDUIENITHNW 40 AWOLYNY J0V19 4V37 ‘VISINYV-VHINYIH ONY NOSYILId 


L6p 


TAXON COLLECTORS LOCALITY 
PA&V 10056 MEXICO. Coahuila: 32 mi SE Saltillo and 8 mi SE JAme on road to Sierra La Viga 
PA&V 10057 MEXICO. Coahuila: 32 mi SE Saltillo and 8 mi SE JAme on road to Sierra La Viga 
M. cuspidata (Torr.) Rydb. P&A 5544 USA.New Mexico: Grant Co.,8 mi E Central on Hwy 90 
M. depau Scribn. P&A 4082 MEXICO. Chihuahua: Just N of Villa Matamoros on Hwy 45 
P&A 4088 MEXICO. Durango: 64 km N of Durango on Hwy 45 
P&A 4091 MEXICO. Durango: 20 km S of Durango on road towards Aserradero La Flor 
M. distans Swallen P&A 5886 MEXICO. Chihuahua: 5.3 mi S of Cusarare on road to Guachochi 
P&A 6010 MEXICO. Durango: 40 km W of Durango on Hwy 40,6 mi W of Rio Chico 
M. distichophylla (J.Presl) Kunth PD,B&K 13583 MEXICO. Chihuahua: Parque Natural Barranca del Cobre, 1 mi E of La Bufa 
M. diversiglumis Trin. P&A 4132 MEXICO. Durango: 18.6 mi W of El Salto, 81.2 mi W Dgo. 
P&A 4137 MEXICO. Durango: 22.7 mi W of La Ciudad on Hwy 40 
P&A 4147 MEXICO. Sinaloa: 2 mi E of Sta. Rita 
P&A 4163 MEXICO. Sinaloa: 1.1 mi NW of Mocorito 
M. dubia E. Fourn. P&A 5550 USA.New Mexico: Grant Co., 12 mi E Central on Hwy 90 
P&A 5558 USA.New Mexico: Grant Co.,0.7 mi NW junction Hwy 61 & 35,0on Hwy 35 
P&A 5809 MEXICO. Chihuahua: Colonia Cumbres de Majalca, approx. 20 mi W Hwy 45,N of 
Chihuahua 
PA&H 8028 MEXICO. Chihuahua: 25.6 mi S Creel on road to Batopilas 
PV,VI 8391 MEXICO. Coahuila: SE San Antonio de las Alazanas & SE of Saltillo, at end of road 
near summit of Coah 
P&A 10593 MEXICO. Coahuila:87 mi NW Muzquiz on Hwy 53 towards Boquilla del Carmen 
P&A 10594* MEXICO. Coahuila: 87 mi NW Muzquiz on Hwy 53 towards Boquilla del Carmen 
P&K 13328, 13330 MEXICO. Nuevo Leon: 6.7 mi W 18 de Marzo, up road towards Cerro del Potosi 
M.dumosa Scribn.ex Vasey P&A 5942 MEXICO. Chihuahua: 12 mi SE Balleza towards Parral 
M. durangensis Y. Herrera PG MEXICO. Durango: 6 mi S Charcos on road towards Juan de Michis 


M. elongata Scribn. ex Beal 


M. eludens C. Reeder 


P&A 4014 


MEXICO. Durango: Ca. 10 km W of San Juan de M 

USA. Arizona: Pima Co., Santa Rita Mts. Box ack. - mi W Hwy 83 on Forest 
Service Road 62 

USA. Arizona: Cochise Co., along Rucker Creek 


t6b 


(€)6L VaIS/O¥O"LINd 


TAXON COLLECTORS LOCALITY 
P&A 4096 MEXICO. Durango: 2.1 mi W of Rio Chico crosing, 21 mi W Dgo. 
P&A 4106 MEXICO. Durango: 5.4 mi W of Navios, 42 mi W of Durango 
P&A 4516 MEXICO. Chihuahua: 9.1 mi E of Cumbres de Majalca 
M.emersleyi Vasey P&A 5068 MEXICO. Baja California: Sierra San Pedro Martir, 0.6 mi E the W Park entrance 
P&A 5326 USA. Arizona: Santa Cruz Co. 3.3 mi W Hwy 289 & Pena Blanca 
P&A 7917 USA. New Mexico: Grant Co., Line along Hwy 78,6 mi W of Mule Creek 
P&A 7918 USA. New Mexico: Grant Co., Line along Hwy 78,6 mi W of Mule C 
PA&H 8018 MEXICO. Chihuahua: 24.3 mi S Creel on road to mee at the ne : Cobre 
PA&V 10765 MEXICO. Chihuahua: 35 mi W Balleza and 24 mi E Guachochi 
PA&V 10805 MEXICO. Chihuahua: Parque Natural Barranca de Cobre, 12.6 mi NE La Bufa and 2 


M. eriophylla Swallen 
M. expansa (Poir.) Trin. 
M. fastigiata (J. Pres!) Henrard 


M. filiculmis Vasey 


M. filiformis (Thurb. ex S. Watson) 


M. filipes M.A. Curtis 
M. flabellata Mez 


G 1626 (CIIDIR) 
H&C 930 (CIIDIR) 


Boyce & Godfrey 1581 


PRA 5627 
P&A 7860 


P&A 3994 
P&A 2648 


E.P Killip 42315 
Pittier 3372 
Pohl & Davidse 


1162 


mi S Basigochi 

MEXICO. Durango: A El Temazcal, 4 km al SW de pee Herrada 
MEXICO. Michoacan: Ge Ucarero, Zinapecuaro, 2 km al S de Querendaro 
USA. North Carolina: Cumberland Co., 13 mi N of a. on route 15 
USA. North Carolina: Columbus Co., Old Dock Savanna. 

ARGENTINA. Provincia Jujuy: 4 km E of Tres Cruces on road to Humahuaca 
ARGENTINA. Jujuy: 24 km W La Quiaca on Hwy 5 towards Sta Cabalina 
USA. ieee ae Co,, Kaibab Plateau, 4 mi N Kaibab Lodge, Pleasant 
Valley, along H 

USA. Colorado: an Co., NW of Saguache, 14 mi up Cochetopa pass road 
(F5750) from Hwy 114 

USA. Arizona: Apache Co., E of McNary 

USA. California: Tulare Co., Lion Meadow 

USA. Washington: Klickitat Co., Washington State Fish Hatchery 

USA. Arizona: Apache Co., 7.4 mi E of McNary off Hwy 260 

USA. Colorado: Pitkin Co., 13 mi S of Leadville on Hwy 24 

USA. Florida: Monroe Co., Big Pine Key 

COSTA RICA. San Jose: Cerro Buena Vista 

COSTA RICA. San Jose: Cerro Buena Vista 


VISUIGNITHNW JO AWOLYNY 3019 4V37 ‘VLIINNV-VuaYNaH ONY NOSHILad 


€6r 


b6r 


TAXON COLLECTORS LOCALITY 
M. flavida Vasey P&A 4138 MEXICO. Durango: 22.7 mi W of La Ciudad on Hwy 40 
P&A 4153 MEXICO. Sinaloa: 48.6 mi NE of Mococrito, 8.5 mi S of Surutato 
P&A 4162 MEXICO. Sinaloa: 1.1 mi NW of Surutato, 61.1 mi NE of Mocorito 
M. flaviseta Scribn. P&A 5911 MEXICO. Chihuahua: side of Barranca El Cobre, approx. 20 mi S Cusarare on road 
to Guachochi 
H 993 (CIIDIR) MEXICO. moa Parque El Tecuan, 58 km ESE of Durango on Hwy 40 to Mazatlan 


M. flexuosa Hitche. 
M. fragilis Swallen 


M. gigantea (E. Fourn.) Hitchc. 


M.glabriflora Scribn. 
M. glauca (Nees) B.D. Jacks. 


M.glomerata (Willd) Trin. 


M. grandis Vasey 
M. gypsophila Reeder & C. Reeder 


ee 982 (CIIDIR) 
J.F. Macbride s.n. 


PD,B&K 13414* 
R.M. Kriebel 5802 
P&KI 8237 


PA&V 10072 
P&A 5511 


P&A 5482 
P&A 5562 
PD,B&K 13413 
P&K 13289 


P&K 13299 


MEXICO. Durango: 4.5 km SW of San Juan de Michis on road to Piedra Herrada 
PERU. Huacachi: Estacion near Muna 
USA. Arizona: Santa Cruz Co., SW of Camelo 
USA. Arizona: Santa Cruz Co., Sycamore Canyon 
MEXICO. Chihuahua: 13.5 mi W of Parral on Hwy 24 
MEXICO. Sinaloa: 34.5 mi NE of Mocorito, 22.6 mi S of Surutato 
MEXICO. Nayarit:8 mi E of Compostela on roads towards Chapalilla and 
Guadalajara 
MEXICO. Sinaloa: 3 mi SW Estado de Durango and 2.2 mi S El Palmito on Hwy 40 
USA. Indiana: Daviess Co.,6 mi N of Washington 
MEXICO. Chihuahua: 76 mi W La Junta and 35.2 mi W Tomochic in Parque Nacional 
Cascad Basaseachic 
MEXICO. Coahuila: 17 mi SE Saltillo and 7.4 mi NW Jame at Bosque de Montana 
USA. Arizona: Cochise Co., Chiricahua Mts., Red Rock Canyon, aprox. 2 mi Rucker 
Canyon Road 
USA. Arizona: Cochise Co., 2.4 mi above Upper Picnic Area, Fort Huachuca 
Military Reservation 
SA.New oie Catron Co., Canyon leading to Cliff Dwellings and upper end of 


Hwy 15 

MEXICO. Sinaloa: 3 mi SW Estado de Durango and 2.2 mi S El Palmito on Hwy 40 
MEXICO. Nuevo Leon: 5.6 mi E junction Hwy 57 on Hwy 58 towards Linares 
MEXICO. Nuevo Leon: 13.4 mi E Hwy 57 on Hwy 58 at crossin Rio Potosi 


(€)6L YaIS/D¥O"LINd 


TAXON 


COLLECTORS 


LOCALITY 


M. hakonensis (Hack. ex Matsum.) 
Makino 


M. himalayensis Hack. ex Hook. 


M. huegelii Trin. 


M. hintonii Swallen 
M.implicata (Kunth 


) 


Trin. 


M. inaequalis Soderstr. 


M.xinvoluta Swallen 


M. iridifolia Soderstr. 


M.jaime-hintonii P.M. Peterson & 


Valdes-Reyna 


M.jaliscana Swallen 


M. japonica Steud. 


M.jonesii (Vasey) Htichc. 


M.lehmaniana Henrard 


M. ligularis (Hack.)Hitche. 


M. lindheimeri Hitchc. 
M. longiglumis Vasey 
M. longiligula Hitche. 


T. Tateoka s.n. 
S&P 5666 
S&P 5324 


S&P 5344 

G.B. Hinton 3059 
P&A 4514 

P&A 4090 

P&A 4095 
A.S.Muller 953 
P&A 6267 

P&A 6281 

P&A 6133,6135 
V&C 2560 


P&A 6137 
P&A 6149 


PA&V 10068* 
P 13710 
P&A 5408 


JAPAN. Hakone: Kanagwa-ken 


CHINA. Xizhang (Tibet) Prov.: Markam Co. Ningjing Shan, Markham Range 
between Mekong & Yantze 

CHINA. Sichuan Prov.: Wenchuan Co.W side of front range NW of Sichuan Basin 
CHINA. Sichuan: Qiunglai Slan,ca 40 km W Wezlou and ca. 120 km NW Clongdu 
MEXICO. Mexico: crucero Temazcaltepec 

MEXICO. Chihuahua: 0.7 mi W of Nuevo Majalca, 8.5 mi W of Hwy 45 

MEXICO. Durango: 20 km S$ of Durango on road towards La Flor 

MEXICO. Durango: 2.1 mi W of Rio Chico crossing, 21 mi W of Durango on Hwy 40 
VENEZUELA. Trujillo: Quebrada de Duri 

MEXICO. Nuevo Leon: 10 mi E of Los Lirios and 12 mi W of Laguna de Sanchez 
USA. Texas: Blanco Co., 0.3 mi E Hwy 280 on Tex 473 

MEXICO, Jalisco: 50 mi W of Ameca on road to Mascota 

MEXICO. Nuevo Leon:La Joya, Cuesta Blanca, 15 km S of Aramberri 


MEXICO. Jalisco: 50 mi W of Ameca on road to Mascota 
MEXICO. Jalisco: Pass above Talpa de Allende, 3.6 mi W of Rio Mascota 
CHINA. Yunnan Prov.: Fugong (N 1/2 Bijiang) Co.W slopes of Bilou Mts. 
CHINA. Kiangwang Shan:ca 15 km E of Dongchuan, ca 120 km NNE of Kunming 
USA. California: Siskiyou Co. Shasta-Trinity National Forest, 9.5 mi SE Hwy 97, 
on military pass road 
PANAMA. Province of Chiriqui: between Rio Quebrado 

CUADOR. Provincia de Azuay: 5.6 km S LA Paz on the Panamerican Hwy, and 36.3 
km N Ona 
MEXICO. Coahuila: 26.7 mi Se Saltillo and 2.7 mi SE Jame on road to Sierra La Viga 
MEXICO. Jalisco: 8.2 mi NW Cuautla on road towards Los Volcanes 
USA. Arizona: Santa Cruz Co. Patagonia Mts., along road to Red Mt., 7.6 mi SE of 
Patagonia 


VIDUIANITHNW 4O AWOLWNV 3019 4¥31 ‘VLalNuy-VududIH ONY NOSYILId 


S6b 


96P 


TAXON COLLECTORS LOCALITY 
P&A 7919 USA. New Mexico: Grant Co., Line along Hwy 78,6 mi W of Mule Creek 
P&KI 8207 MEXICO. Chihuahua: 33.7 mi W La Junta on road to Parque Nacional Cascada de 
Basaseachic 
P 9605 MEXICO. Chihuahua: 20.8 mi SW La Junta on road to Creel 
M. lucida Swallen P&A 5882 MEXICO. Chihuahua: 5.3 mi S Cusarare on road to Guachochi 
PA&H 8039 MEXICO. Chihuahua: 10.7 mi S Mex 127 on road to Batopilas 
M. macroura (Kunth) Hitche. P&CA 9769 MEXICO. Oaxaca: 11.4 mi W San Juan Mixtepec and 1.5 mi E San Isidro 
Chicahuaxtla 
PA&V 10986 MEXICO. Michoacan: 8.4 mi SE Zacapu on Mex 15 towards Quiroga 
M. majalcensis PM. Peterson P&A 4573 MEXICO. Chihuahua: E of Cumbres de Majalca 
M. mexicana (L.) Trin. P&A 4765 USA. Idaho: Lemhi Co., 2.5 mi E of Salmon River on Warm Springs Creek 
M. mexicana var. filiformis (Torr.) Morton 11689 CANADA. Ontario: Lake Timiskaming (Dawson Point) 


cribn 
M. michisensis Y. Herrera & 
M. Peterson 


M.microsperama (DC.) Trin. 


M. minutissima (Steud.) Swallen 


M. montana (Nutt.)Hitchc. 


H&A 986 (CIIDIR) 


tei 8913 


PA&H 8033 
P&KI 8214 


P&CV 9733 


MEXICO. Durango: San Juan de Michis, Potrero Las Escobas 


ECUADOR. Provincia de Azuay: 10.2 km N Ona on the Panamerican Hwy 
ECUADOR. Provincia de Pichincha: 13 km N Calderon on the Panamerican Hwy 
MEXICO. Baja California Sur: 25 km S of La Paz, W side of Isla La Partida 

MEXICO. Sonora: 18.2 mi E of Los Tanques on road to Milpillas 

USA. Arizona: Santa Cruz Co., Sycamore Canyon 

USA. Nevada: Clark Co., Lake Mead 

USA. Arizona: Coconino Co., W of Flagstaff 

MEXICO. Chihuahua: 12 mi SW of Madera off Hwy 16 towards Cuauhtemoc 
MEXICO. Durango: 3.2 mi W of Rio Chico crossing, 22.1 mi W of Durango on Hwy 40 
MEXICO. Chuhuahua: 9.1 mi E of Cumbres de Majalca 

MEXICO. Chihuahua: 25.6 mi S Creel on road to Batopilas 

MEXICO. Chihuahua: 33.7 mi W La Junta on road to Parque Nacional Cascada de 
Basaseachic 

MEXICO. Oaxaca: 4.8 mi NW Tlaxiaco on road to San Juan Mixtepec 


(€)6L VOIS/D¥O LIS 


TAXON COLLECTORS LOCALITY 
P&A 9971 MEXICO. Tlaxcala: 5.2 mi N Tlaxco on Mex 119 to Zacatlan 
P&KI 8171 USA. Arizona: Cochise Co., 10 mi W Portal on F542, E of Onion Pass 
P&A 5438 USA. Arizona: Santa Cruz Co, Patagonia Mts., 12.3 mi S Patagonia on Forest Service 
Road 49 
M. mucronata (Kunth) Trin. P&A 10778 MEXICO. Chuhuahua: 40 mi W of Balleza and 19 mi E of Guachochi 


M. mutica (Rupr. ex E. Fourn.) Hitche. 


M. nigra Hitchc. 
M. orophila Swallen 
M. palmeri Vasey 


M. palmirensis Grigi 
M. pauciflora Buckley 


on & Laegaard 


M. peruviana (P. Beauv.) Steud. 


M. pilosa PM. Peterson, Wipff & 
S.D. Jones 


M. plumbea (Trin.)Hitche. 


M. polycaulis Scribn. 


M. porteri Scribn. ex Beal 


M. pubescens (Kunth) 
M. pubigluma Swallen 


Hitchc. 


A.S. Hitchcock 6348 
A 11081 

P&A 11105 

P&KI 8322 

P&A 11134 

P&A 5478 


P&A 5681 


PA&PO 8810 
P&A 5715* 
P&J 9308 
P&A 4071 
P&A 4125 
P&A 11061 


PA&V 10765 
P&A 5406, 5407 


PA&V 10764 

P&KI 8144 
PD.B&K 13440 
P&A 10593, 10594 
P&K 13329 


MEXICO. Veracruz: Orizaba 

MEXICO. Mexico: 28.3 mi NE Temascaltepec on Mex 134 towards Toluca 
MEXICO. Mexico: 15.6 mi E Amecameca and 2 mi N Paso de Cortes 

MEXICO. Tamaulipas: 63 mi SW Cd. Victoria on Mex 101 towards San Luis Potosi 
MEXICO. San Luis Potosi:45 mi NE San Luis Potosi on road towards Guadalcazar 
USA. Arizona: Cochise Co., 3.1 mi above Upper Picnic Area, Fort Huachuca Military 
Reservation 

USA. Arizona: ae Co., Santa Rita Mts. Box Canyon, 7 mi W Hwy 83 on Forest 
Service Road 6 

ECUADOR. ie: de Chimb 8.9 km N Palmira on the Panamerican Hwy 
USA. Texas: Culberson Co,, Giacsiipe Mts, Pine a on Hwy 62 (180) 
ECUADOR. Provincia Cotopaxi: Lago Limpiopun 

MEXICO. Durango: 14 mi SW of El Vergel on Hwy 7 

MEXICO. Durango: 7.0 mi W of El Salto on Hwy 40 

MEXICO. Mexico: 5 km NE of Tejupilco on Mex 134 towards Temascaltepec 


MEXICO. Chihuahua: 35 mi W Balleza and 24 mi E Guachochi 

USA. Arizona: Santa Cruz Co. Patagonia Mts., along road to Red Mt., 7.6 mi SE of 
Patagon 

(coc mi W Balleza and 24 mi E Guachochi 

MEXICO. Chihuahua: 17 mi S of Nuevo Casas grandes on Mex 2 

MEXICO. Durango: 4.5 mi N of Borbollones, N of Hwy 40 

MEXICO. Coahuila: 85.5 mi NW of Muzquiz on Hwy 53 towards Boquilla del Carmen 
MEXICO. Nuevo Leon: 6.7 mi W 18 de Marzo up road towards Cerro del Potosi 


VISUTSNITHNW JO AWOLVNY 30V19 4V31 ‘VIdINdY-VuIdyIH ONY NOSYIL3d 


L6¥ 


TAXON COLLECTORS LOCALITY 
M. folent Thurb. ex A. Gray P&A 5614 USA. Arizona: Apache Co., 20 mi N St. Johnson Hwy 666 (61) 
M. purpusti Me P&K! 8325 MEXICO. Tamaulipas: 63 mi SW Cd. Victoria on Mex 101 towards San Luis Potosi 
M. eee (Kunth) Trin. P&A 6009 MEXICO. Durango: 40 km W Durango on Hwy 40, and 6 mi W Rio 
P&A 11082 MEXICO. Mexico: 28.3 mi NE er ieecniens on ee 134 towards ie 


M. racemosa (Michx.) Britton, 
Sterns & Poggenb 
M. ramosa (Hack. ex Matsum.) 


Makino 
M. ramulosa (Kunth) Swallen 
M. reederorum Soderstr. 


M. repens (J. Presl) Hitche. 

M. reverchonii Vasey & Scribn. 
M. richardsonis (Trin.) Rydb. 
M. rigens (Benth.) Hook. 


M. rigida (Kunth) Trin. 


H&C 906 (CIIDIR) 
H&C 899 (CIIDIR) 
H&C 917 (CIIDIR) 
M.W. Talbot 814 


S&P 5302 
P&A 4109, 4113 
P&A 4121 

K 13643 


PRA 5422 


PA&PO 8820 
P&KI 8187 
P&K 13300 
P&K 13301* 
P&KI 8316 


MEXICO. Mexico: Parque Nacional Nevado d 

MEXICO. Oaxaca Ladera SW del Cerro Pelon, oo m antes del mirador 
MEXICO. Puebla: Ladera E del Popocatepetl, 10 km W de Santiago Salicintla 
USA. New Mexico: Santa Fe Co., Santa Fe Canyon 


CHINA. Kiangwang Shan:ca 15 km E of Dongchuan, ca 120 km NNE of Kunming 


MEXICO. Durango: W of Navios 

MEXICO. Durango: W of El Salto 

MEXICO. Michoacan: E of Opopeo 

MEXICO. Durango: 56 km W Durango,on Hwy 40 

MEXICO. Durango:6 mi S Charcos on road towards San Juan de Michis 
MEXICO. Sinaloa: 3 mi SW Estado de Durango and 2.2 mi S El Palmito on Hwy 40 
USA. Arizona: Santa Cruz Co. Patagonia Mts., junction of Forest Service Road 135 & 49 
USA. Texas: Bexar Co.5 mi N of San Antonio, city limits on Hw 

MEXICO. Chihuahua: 10 mi W of Cuahutemoc on Hwy 16 

MEXICO. Chihuahua: 5.6 mi N Cuahutemoc on Mex 23 

MEXICO. Chihuahua: 76 mi W La Junta in Parque Nacional Cascada de Basaseachic 
MEXICO. Chihuahua: 52.5 mi SE Villa Matamoros and 1 mi N Ejido Revolucion 
ECUADOR. Provincia de Azuay: 17.7 km N Ona on the Panamerican Hwy 
ECUADOR. Provincia de Chimborazo: 8.7 km W Alausi on road to Sibambe 
MEXICO. Chihuahua: 44.5 mi SE Madera on Mex 16 and 1 mi S Temosachic 
MEXICO. Nuevo Leon: 13.4 mi E Hwy 57 on Hwy 58 at crossin Rio Potosi 
MEXICO. Nuevo Leon: 13.4 mi E Hwy 57 on Hwy 58 at crassin Rio Potosi 
MEXICO. Tamaulipas: 63 mi SW Cd. Victoria on Mex 101 towards San Luis Potosi 


86r 


(€)6L VdIS/9¥O'LINE 


TAXON COLLECTORS LOCALITY 
P&CV 9707 MEXICO. Oaxaca: 6.5 mi NE Villa de Tamazulapan and 2 mi NE Teotongo 
P&CV 9728 MEXICO. Oaxaca: 5 mi SW Teposcolula and 2.5 mi NE Yolomecatl on Mex 125 
P&A 5455 USA. Arizona: Cochise Co., 2 mi SW Sunnyside along Forest Service Road 228 
P&A 5434 USA. Arizona: Santa Cruz Co. Patagonia Mts., 12.3 mi S Patagonia on Forest Service 
Road 49 
M. robusta (E. Fourn.) Hitche. PG&K 13594 MEXICO. Mexico: Durango: 5 mi E of Mezquital on road to Charcos 
M. schmitzii Hack. P&A 4631 MEXICO. Michoacan: 6.1 mi W of Ciudad Hidalgo on Hwy 15 
M. scoparia Vasey P&A 6079 MEXICO. Nayarit: 29 mi SW of Tepic on Hwy 15 to Guadalajara 
M. schreberi J.F.Gmel. P 14231 USA. Maryland: Montgomery Co. Bethesda, 4520 Cheltenham Dr. 
M. seatoni Scribn. P&A 9946 MEXICO. Puebla: 3.5 mi SE Cd. Serdan on Mex 140 
M. setarioides E. Fourn. P&CV 9897 MEXICO. Oaxaca: 1.4 mi E Ayutla on Mex 179 towards Mitla 
M. setifolia Vasey P&K 13376 MEXICO. San Luis Potosi: 2.5 mi E Hwy 57 on road towards Guadalcazar 
P&A 5716 USA. Texas: Culberson Co., Guadalupe Mts., Pine Springs on Hwy 62 (180) 
M. sinuosa Swallen P&A 12590 MEXICO. Chihuahua: Sierra El Nido, 16.7 mi W of Hwy 45 on road up Los Prietos Canyon 
M. sobolifera (Muhl.ex Willd.) Trin. — C.H. Bissell s.n. USA. Connecticut: Rocky woods near Savage St. 
M. speciosa Vasey PG&K 13626 MEXICO. Durango: 7 mi SE of Mezquital on road to Charcos 
PD,B&K 13409 MEXICO. Sinaloa: 3 mi SW Estado de Durango and 2.2 mi S El Palmito on Hwy 40 
M. spiciformis Trin. P&A 6244 MEXICO. Coahuila: approx. 20 mi SE Saltillo on road to Los Lirios 
P&KI 8334 MEXICO. Nuevo Leon: 3.8 mi S Allende on Mex 85 towards Montemorelos 
P&A 9896 MEXICO. Oaxaca: 1.4 mi E Ayutla on Mex 179 towards Mitla 
P&A 9945 MEXICO. Puebla: 3.5 mi SE Cd.Serdan on Mex 140 
P&A 10590 MEXICO. Coahuila: 85.5 mi NW Muzquiz on Hwy 53 towards Boquilla del Carmen 
P&KI 8332 MEXICO. Tamaulipas: 55 mi SW Cd. Victoria on Mex 101 towards SanLuis Potosi 


M. straminea Hitche. 
M. stricta (J. Presl) Kunth 


M. strictior Scribn.ex Beal 


R.Endlich 1210 
P 13709 


P&KI 8324 
P&A 4520 
P&A 4039 
P&A 4054 


MEXICO. Chihuahua. 

MEXICO. Jalisco: 8.2 mi NW Cuautla on road towards Los Volcanes 

MEXICO. Tamaulipas: 63 mi SW of Ciudad Victoria on Mex 101 to San Luis Potosi 
MEXICO. Chihuahua: 21.1 mi W of Hwy 45, 0.4 mi E of Cumbres de Majalca 
MEXICO. Chihuahua: 3.1 mi S of Hernandez Javales, 32 mi SW of Colonia Juarez 
MEXICO. Chihuahua: 15.5 mi W of La Junta on road to Tomochic 


VIDUISNITHNW 40 AWOLWNY 3018 4¥31 ‘VLSINuY-VudduaH ONY NOSYIL3d 


66¢ 


TAXON 


COLLECTORS 


LOCALITY 


M. subaristata Swallen 

M. sylvatica (Torr.) Torr. ex A. Gray 
M. tenella (Kunth) Trin. 

M. tenuiflora (Willd.) Britton, 


Sterns & Poggen. 
M. anes (Kunth) Trin. 


M. tenuissima (J. Presl) Kunth 
M. texana Buckley 


M. thurberi (Scribn. 


Rydb. 


M. torreyana (Schult.) Hitch. 
M. torreyi (Kunth) Hitche. ex Bush 


P&A 4098 

FW Pennel 18572 
C.A.Weatherby 5139 
P&A 4618 

P&A 4755 

Ch.C. Deam 46,172 


P&A 4513 
P&A 8104 


P 9613 
P&A 4545 


P&A 4045 


P&A 11418 


MEXICO. Durango: 2.1 mi W of Rio Chico crossing, 21 miW of Durango on Hwy 40 
MEXICO. Durango: El Salto (aserradero 
USA. Connecticut: Stafford 

MEXICO. Michoacan: S of Uruapan on Mex 37 

MEXICO. Nayarit: 10.2 mi W of Tepic on road to Miramar 
USA. Indiana: Noble Co.,5 mi SE of Albion 


<7 


MEXICO. Chihuahua: 0.7 mi W of Nuevo Majalca, 8.5 mi W of Hwy 45 

MEXICO. Chihuahua: 54.4 mi N Parral on Mex 24 to Chihuahua 

MEXICO. Chihuahua: 3 mi NE of Parral on Hwy 45 towards Chihuahua 

MEXICO. Chihuahua: 15.6 mi NE of El Vergel on Hwy 24 

MEXICO. Durango: 21 mi of Rio Chico, 21 mi W of Durango on Hwy 40 

USA. Texas: Jeff Davis Co., 7.5 mi SW Hwy 118 on Hwy 166, NE Valentine 

Hahee Jalisco: 2 mi NW of Magdalena on Mex 15 and 15 mi from Guadalajara 
MEXI ihuahua: 10 mi SW La Junta on road to Creel 

nC Chihuahua: Cascada de Basaseachic, 37 mi W of Tomochic, 0.6 mi from 
overloo 

ee eee 5.0 mi S of Hernandez Javales 

USA. Arizona: Santa Cruz Co., 7 mi SW of Canelo on Hwy 83 

USA. Arizona: Apache Co., Antelope House Overlook, N rim above Canyon del Muerto 
USA. Arizona: Apache Co, Antelope House Overlook, N rim above Canyon del Muerto 
USA. New Mexico: Rio Arriba Co., on Hwy 84, at Echo Amphitheate 

USA. New Jersey: Burlington Co.,0.1 mi N of Sale Burlington Co. lines on Hwy 206 
ele Provincia Salta: 48 km E of Cachi on Hwy 40 to Salta 

ARGENTIN Mendoza; San Carlos: near ae ne Hondo on junctn road 


Hw 
ARGENTINA. Mendoza; Depto. Lujan de Cuyo: approx. 21 km SW Potrerillos on road 
toward San Jose 


00S 


(€)6L VaIs/O¥O'LINa 


TAXON COLLECTORS LOCALITY 
P&A 11621 eed declan: Depto. Tafi del Valle: 30 km SE Amaicha de Valle & 25 km 
NW Tafi del V 
P&A 11701 ARGENTINA. - Depto. Chicoana: just E Piedra del Molino on Hwy 33 between E| 
Carril and Cachi 
P&A 11726 ARGENTINA. Salta; Depto. San Carlos: 3 km S Isonza and 23 km N Amblayo 
P&A 5549 A.New Mexico: Grant Co., 12 mi E Central on Hwy 90 
M. uniflora (Muhl.) Fernald JV. Haberer 3266 USA. New York: Oneida Co., Forestforth 
M. utilis (Torr.) Hitche. P&A 6259 MEXICO. Coahuila: 8 mi E of Los Lirios on road to Laguna de Sanchez 
M. vaginata Swallen P&A 4070 MEXICO. Chihuahua: 0.5 mi NE of El Vergel 
P&A 4111 MEXICO. Durango: 5.4 mi W of Navios, 42 mi W of Durango on Hwy 40 
P&A 4124, 459] MEXICO. vale 7.0 mi W of El Salto on Hwy 40 
M. versicolor Swallen P&A 11053 MEXICO. Mexico: 1.1 mi N Tejupilco on Mex 134 towards Temascaltepec 
M. villiflora Hitchc. var. villiflora P&A 6228 MEXICO. San Luis Potosi: 10.3 mi NW Matehuala on road to Cedral, near Km 


M. villiflora var. villosa 
(Swallen) Mor 
M. virescens (Kunth) Kunth 


M. virletii (E. Fourn.) Soderstr. 


M. watsoniana Hitchc. 


M. wrightii Vasey ex J.M.Coult. 


M.xanthodas Soderstr. 


JR.& C.G. Reeder 4536 


P&A 5876 
P&A 5589 


PD,B&K 13429 

P&CA 9709 

P&CA 9723 

P&CA 9724 

P&CA 9729 
Hernandez s.n. (HUAA) 


P 9586 
Hernandez & Sharp 311 


marker 12 
USA. Texas: Glasscock Co., 15 mi S of Stanton 


MEXICO. Chihuahua: 15 mi S San Juanito & 3 mi N Creel 

USA. New oe Co., 2.1 mi E Hwy 180 on Forest Service Road 35, San 
Francisco Mts. 

MEXICO. cee 2.4 mi N Borbollones, N Hwy 40 

MEXICO. Oaxaca: 10 mi NE Villa de Tamazulapan and 5.5 mi NE Teotongo 
MEXICO. Oaxaca: 2.6 mi E Teposcolula on Mex 125 

MEXICO. Oaxaca: 2.6 mi E Tepascolula on Mex 12 

MEXICO. Oaxaca: 5 mi SW Teposcolula and 2.5 mi NE Yolomecatl on Mex 125 
MEXICO. Aguascalientes: San Jose de Gracia, Sierra Fria 

USA.New Mexico: San Francisco Mts., Potato Patch, 3 mi E Hwy 180 on Forest 
Service Road 35 

MEXICO. Chlntanua: 13 mi W Cuahutemoc on Mex 16 

MEXICO. C B Escuipulas and Canada Honda 


VIDUJENITHNW 40 AWOLVNV 30V19 4V37 ‘VI9INYY-VUINYIH ONY NOSHIL3d 


LOS 


502 


BRIT.ORG/SIDA 19(3) 


APPENDIX 2. 
Data set used in the cladistic analysis. 

Eragrostis acutiflora 2[12]111221221112[23]2.  M.emersleyi (12]413111211212322 
Frioneuron avenaceum 1211111111121232 ) M.eriophylla 2121111111111[23]22 
Leptochloa virgata (12]123122122212322  M.expansa (23]431111211112322 
porobolus airoides 3[24]33232211212322 ~~ M.fastigiata 2111111111111132 
M. aguascalientensis 2121111111111322 — M. filiculmis (23)(14)11111111111[23)[23]2 
M. alamosae Speen ines M. filiformis 2111121111111132 
M. andina 21211221 . 1232 M.filipes (23]431111211112322 
M. angustata crea os 2)1— M. flabellata 4121111111111222 
M. annua 11121 121132. M.flavida (12)1111211111111[23]2 
M. appressa namie 11132 M< flaviseta 2121111111121[23]22 
M. arenacea 2111111111121[12]22 ~~ M. flexuosa 2111111111121222 
M. arenicola St eet M. fragilis (12]111121111121132 
M. argentea 21211111111212(23]2  =M.gigantea 2433111211212222 
M. arizonica 2111111111121132 = M.glauca 111112[12)12[12]111132 
M. arsenei 11211221211112[23]2 — M.glabriflora 1121122122111132 
M. asperifolia 21111111111112[23]2) M.glomerata 1121122122111132 
M.articulata (23]231131211112322 9 M.grandis 2(24]23111211212322 
M. aurea 2(24)23111211212322 = M.gypsophila 3231231211112311 
M. brandegei (12]111122122111132 9M. hakoensis 1123122122121222 
M. brevifolia 2111111111111122) MM. des aa 1123122122121222 
M. breviligula 2(24]33111211212322 9 M.hintonil 2121111111111322 
oe 111{12]1211111211[23]2 ee 1122122122121[12]22 
M. brevi 21111211111111[23]2) M.implicata 2111121111111112 
M. ie inata = 4)21111111111212 ) M_inaequalis 2423111211212322 
M. bushii 121122122111232 ) M.xinvoluta 2433111211212322 
M. californica oo 122122111132 ~~ M.iridifolia 2(24]23111211212322 
M. capillaris 2(24]31 ere Lease M. jaime-hintonii 2411131111112(23]22 
M. capillipes 1121111111132.) M.jaliscana 3231231211112311 
M. caxamarcensis eens M. japonica 1123122122121222 
M. ciliata 11111221221111[23]2 M. jonesti 2(14]111111111111[12]2 
M. cleefii 2111111111111122)) M.lehmanniana 2[24]23111211222322 
M. crispiseta (12]111121111121122 M. ligularis 2111121111111132 
M. cualensis 1121111111111322 ) M.lindheimeri 2423111211212222 
M. xcurtisetosa 1121122122111232 = M. longiglumis 2(24]33111211212[23]22 
M. curtifolia 212112212({12]121322 ~~ M.longiligula 2(24]2311121[12]212222 
M. curviaristata 11231221221212([23]2 = M. lucida 3231231211112321 
M.curvula {23][14J21111111111322 M.macroura 3231231211112311 
M. cuspidata 2111111111121232 = M.majalcensis a pees 
M. depauperata 2111121111121132 > M.mexicana 11211221221112(23]2 

M. distans 2(24]2[23]111211222322 M. mex. var-filiformis 11211221221112[23]2 

M. distichophylla 2(24]23111211212322 = M.michisens 2121111111111222 
M. diversiglumis 1111122122111132) M.microsperma 1111122122111132 
M. dubia 3231231211112311 = M.minutissima 2111121111121132 
M.dumosa 1121122122111132 M.montana (23)[14)3111111111[12)[23]22 
M. durangensis 1121231111111222.)) M.mucronata 3231231211112311 
M. elongata 2(24]31111111112322 M. mutica 2[24]21111211112322 
M. eludens (12}1111211111211[23] M. nigra 323123121111231] 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 


M. orophila 1711111111111[12]22) MM. setifolia 
M. palmeri 3231231211112321 ) M.sinuosa 
M. palmirensis 2111111111111122 ) M. sobolifera 
M. pauciflora 212112212[12]111222 — M.speciosa 
M. pectinata 11111221221111[23]2  M. spiciformis 
M. peruviana 2111121111111132 ) M.straminea 
M. pilosa 241 - ; 1211212222 ~~) M stricta 
M. plumbea a 111112[23]2) M.strictior 
M. polycaulis en 1111[23]2. M.subaristata 
M. porteri oer 22122111[12]22— M. sylvatica 
M. pubescens See | 212222 M.tenella 
M. pubigluma 2421 112(23]22. M. tenuiflora 
M. pungens screener M. tenuifolia 
M. purpusii 2221111111112[23]22) M. tenuissima 
M. quadridentata (23]121111111111322 MM. texana 
M. racemosa 11211221221112[23]2 MM. thurberi 
M.ramosa 1121122122121[12][23]2  M. torreyana 
M. ramulosa (12]111121111111132 — M. torreyi 
M. reederorum 2433111211212322 ~~) M.uniflora 
pen 21111111111111[23]2 utilis 
M. reverchonil 2(14]21111111112[23]22 M. vaginat 
M. richardsonis 2111111111111132  M. versicolor 
M. rigens 3231231211112321 — M.villiflora 
M. rigida 3231231211112311 M. villvar.villosa 
M. robusta 2433111211212322 ~~ M. virescens 
M. scoparia 2(24)23111211212322 — M. virleti 
M. schmitzii 2111121111111132 ) M.watsoniana 
M. schreberi 1121122122111222 ) M.wrightii 
M. seatonii 2111111111111222)) M_.xanthodas 
M. setarioides 1121122122111222 
ACKNOWLEDGMENTS 


2(24]231 Peta 2 
211112212211 ] 
at ee 
caren 12(23]2 
11211 111 
eon 111 
11211221 cae 
1111122122111) 
: 


112212211 
ohne 
2111121111121122 
211112212[12]111[12]22 


2(24)23111111111[12)[23]2 
2111111111122[23]22 
2(14]721111111121[12][12]2 
211111111111112]22 
21111211111111[23]2 
2(24]31111111112[23]22 
2111111111111122 
2111111111111122 
3(12]3121111111[12][23}12 
oe 
21211 ae 
ane eet 
2(24)23112121121 — 


We thank Stanley Yankowski for advice in preparing the anatomical prepara- 
tions; Carol R. Annable and Marjorie B. Knowles for preparing a portion of the 
anatomical slides; Robert J. Soreng for assistance with CLADOS: J. Travis Co- 
lumbus and Robert D. Webster for reviewing the manuscript; the Smithsonian 
Institution’s Fellowships and Grants for providing a Short-term Visitor Grant 
to YH, the Scholarly Studies Grants Program; and the Natural Museum of Natu- 
ral History’s, Research Opportunities Fund. 


REFERENCES 


Beauvols, PB.1812.Essai d’une | 


avec figures représentant les eee de fouls les genres, 16, 26. Paris. 
Bertyn, G.P.and J.P. Miksce. 1976. Botanical microtechnique and cytochemistry. Ames: lowa 


State University Press. 


Brown, W.V. 1977.The kranz syndrome and its subtypes in grass systematics. Mem. Torrey 


Bot. Club 23:1-97. 


504 BRIT.ORG/SIDA 19(3) 


Desvaux, A.N. 1810. Extrait d’un mémoire sur quelques nouveau genres de la famille des 
Graminées. Nouv. Bull. Sci. Soc. Philom. Paris. 2:187-190,. 

Duvatt, MLR., PM. Peterson, and A.H. CHRISTENSEN. 1994, Alliances of Muhlenbergia (Poaceae) 
within New World Eragrostideae are identified by phylogenetic analysis of mapped 
restriction sites from plastid DNAS. Amer. J. Bot. 81:622-629. 

E.uis, R.P. 1976. A procedure for standardizing comparative leaf anatomy in the Poaceae. I. 
The leaf-blade as viewed in transverse section. Bothalia 12:65-109. 

Foster, A.S. and E.M. Girrorp. 1947. Improvements in paraffin method. Stain Technology 
22(4):129-131. 

Fournier, E. 1886. Mexicanas Plantas. Pars secunda: Gramineae 2:1-160. 

Gotosorr, PA. 1998. NONA, version 1.8. Program and Documentation 

Gutierrez, M., V.E. Gracen, and G.E. Eowarps. 1974. Biochemical and cytological relationships 
in C4 plants. Planta (Berlin) 119:279-300. 

Harterstey, PW. 1984. Characterization of C4 type leaf anatomy in grasses (Poaceae). Meso- 
phyll: bundle sheath area ratios. Ann. Bot. 53:163-179. 

Harrersiey, PW. and AJ. Browninc. 1981. Occurrence of the suberized lamella in leaves of 
grasses pf different photsynthetic types. |. In parenchymatous bundle sheaths and 
PCR (‘Kranz’) sheaths. Protoplasma 109:371-401 

Harterstey, PW.and L.WATSON. 1976.C4 grasses:an anatomical criterion for distinguishing 
between NADP-malic enzyme species and PCK or NAD-malic enzyme species. Aus- 
tral. J. Bot. 24:297-308. 

Harterstey, PW. and L. WATSON, 1992. Diversification of photosynthesis. In: C.P. Chapman, 
ed. Grass evolution and domestication. Cambridge University Press: Cambridge. Pp. 
38-116. 

Herrera Arrieta, Y. 1998. A revision of the Muhlenbergia montana (Nutt.) Hitchc. complex 
(Poaceae: Chloridoideae). Brittonia 50:23-50. 

Herrera Arrieta, Y. and be tA Cerda-Lemus, M. 1985. Muhlenbergia aquascalientensis, a new 
species from Mexico. Novon 5:278-280. 

HerReRA Arrieta, Y. and W.F. Grant. 1993. Correlation between generated morphological 
character data and flavonoid content in the Muhlenbergia montana complex.Canad. J. 
Bot./1:816-826. 

Herrera Arrieta, Y.and W.F. Grant. 1994. Anatomy of the Muhlenbergia montana (Poaceae) 
complex. Amer. J. Bot. 81:1038-1044. 

HitcHcock, A.S. 1935. Muhlenbergia. (Poales) Poaceae (pars). North Amer. Fl. 17:431-476. 

Howm, H.T. 1901. Some anatomical characters for certain Gramineae. Botanisches 
Centralblatt 11:101-103. 

Jacoss, S.W.L. 1987. Systematics of the chloridoid grasses. In: Soderstron, T.R., K.W. Hilu, CS. 
Campbell, and M.E. Barkworth, eds. Grass systematics and evolution. Smithsonian |n- 
stitution Press, Washington, D.C. Pp. 277—286. 

Link, G.M. 1933. Hortus Regius Botanicus Berolinensis 2:248.G. Reimer: Berlin. 

Morben, C.W.1985.A biosystematic study of the Muhlenbergia repens complex (Poaceae). 
Ph.D. dissertation. Texas A&M University: College Station. 


— 


PETERSON AND HERRERA-ARRIETA, LEAF BLADE ANATOMY OF MUHLENBERGIA 505 


Morpen, C.W.1995.A new combination in Muhlenbergia (Poaceae). Phytologia 79:28-30. 

Moroen, C.W. and S.L. HatcH. 1987. Anatomical study of the Muhlenbergia repens 
complex(Poaceae: Chloridoideae: Eragrostideae). Sida 12:347-359. 

Moroen, C.W. and S.L. Hatch. 1989. An analysis of morphological variation in Muhlenbergia 
capillaris (Poaceae) and its allies in the southeastern United States. Sida 13:303-314. 

Morben, C.W.and S.L. Harcu. 1996. Morphological variation and synopsis of the Muhlenbergia 
repens complex (Poaceae). Sida 17: 349-365. 

Nixon, K.C. 1999. The parsimony ratchet,a new method for rapid parsimony analysis. Cla- 
distics 15:407-414. 

Nurtatt,T. 1848. Descriptions of plants collected by Mr.William Gambel in the Rocky Moun- 
tains and Upper California. Proc. Natl. Acad. Sci. Philadelphia 4:23. 

Onsuai, R., T. Murata, and N. CHonan. 1982.C,4 syndrome of the species in the Dichotomiflora 
group of the genus Panicm (Gramineae). Bot. Mag. (Tokyo) 95:339-347. 

Peterson, PM. 2000. Systematics of the Muhlenbergiinae (Chloridoideae: Eragrostideae). 
Pp. 195-211 in Grasses: Systematics and Evolution,eds.S.W.L.Jacobs and J.Everett. CSIRO, 
Melbourne. 

Peterson, PM. and C.R. Annable. 1991. Systematics of the annual species of Muhlenbergia 
(Poaceae-Eragrostideae). Syst. Bot. Monogr. 31:1-109. 

Peterson, PM. and C.R. Annaste, and V.R. FrancescHi. 1989. Comparative leaf anatomy of the 
annual Muhlenbergia (Poaceae). Nordic J. Bot. 8:575-583. 

Peterson, P.M., J.T. Cotumsus, R. Cerros Taticpa, and M.S. Kinney. 2001. Phylogenetics of 
Muhlenbergia and relatives (Poaceae: Chloridoideae) based on internal transcribed 
spacer region sequences (nrDNA). Amer. J. Bot. abstract, http://www.botany2001.org/. 

Peterson, PM.,M.R. Duvatt, and A.H. CHrisTeNseNn. 1993. Allozyme differentiation among Bealia 
mexicana, Muhlenbergia argentea, and M. lucida (Poaceae: Eragrostideae). Madrono 
40:148-160. 

Peterson, PRM. and Y. Herrera-Arrieta. 1995. Allozyme variation in the amphitropical disjunct, 
Chaboissaea (Poaceae: Eragrostideae). Madrono 42:427-449, 

Peterson, PM. and O. Morrone. 1998. Allelic variation in the amphitropical disjunct Lycurus 
setosus (Poaceae: Muhlenbergiinae). Madrono 44:334-346. 

Peterson, RM. and J.J. Ortiz-Diaz. 1998. Allelic variation in the amphitropical dis 
Muhlenbergia torreyi (Poac Muhlenbergii Brittonia 50:381-391. 

Peterson, PM., R.W. Wesster, and J. Vatpes-Reyna. 1995. Subtribal classification of the New 
World Eragrostideae (Poaceae: Chloridoideae). Sida 16:529-544. 

Peterson, P.M., R.W. Weester, and J. VaLpes-Reyna. 1997. Genera of New World Eragrostideae 
(Poaceae: Chloridoideae). Smithsonian Contr. Bot. 87:1-50. 

Piicer, R. 1956. Gramineae ||. Unterfamilien: Micraioideae, Eragrostideae, Oryzoideae, 
Olyroideae. In: H.Melchoir and E.Werd ,eds. Die naturalichen Pflanzenfamilien 
2d ed. 14:1-168. Berlin: Duncker & Humblot. 

Pout, R.W. 1969. Muhlenbergia, subgenus Muhlenbergia (Gramineae) in North America. 
Amer. Midland Nat. 82:512-542. 


unct 


se 


506 BRIT.ORG/SIDA 19(3) 


PRENDERGAST, H.D.V., PW. Harrerstey, N.E. Stone, and M. Lazarives. 1986.C,4 acid decarboxylation 
type in Eragrostis (Poaceae): patterns of variation in chloroplast position, ultrastructure 
and geographical distribution. Pl. Cell Environm. 9:333-344. 

PRENDERGAST, H.D.V., RW. Harterscey, and N.E. Stone. 1987. New structural/biochemical asso- 
ciations in leaf blades of C4 grasses (Poaceae). Austral. J. Pl. Physiol. 14:403-420. 

Prest, K.B. 1830. Reliquiae Haenkeanae 1:207-356. 

Renvoize, S.A. and W.D. CLayton. 1992. Classification and evolution of the grasses. In: C.P. 
Chapman, ed. Grass evolution and domestication. Cambridge University Press: Cam- 
bridge. Pp. 3-37. 

ScHwase, H. 1948. Contribucidn a la anatomia foliar de algunas Agrostideas. Las especies 
Argentinas de los géneros Muhlenbergia y Lycurus, sus relaciones con especies 
Americanas y las relaciones intraenéricas de Muhlenbergia, Lycurus, Sporobolus, y 
Epicampes. Lilloa 16:141-160. 

SINHA, N.R. and E.A. Kettoaa. 1996. Parallelism and diversity in multiple origins of C4 photo- 
synthesis in the grass family. Amer. J. Bot. 83:1458-1470. 

SODERSTROM, T.R. 1967. Taxonomic study of the subgenus Podosemum and section Epicampes 
of Muhlenbergia (Gramineae). Contr. U.S. Natl. Herb. 34:75-189. 

Sykes, G.R., A.H. CHRISTENSEN, and P.M. Peterson. 1997.A chloroplast DNA analysis of Chaboissaea 
(Poac Erag id ). Syst. Bot. 22:291-302. 

THURBER, G. 1863. Gramineae. In: Enumeration of the species of plants collected by Dr.C.C. 
Parry,and Messrs. Elihu Hall and J.P. Harbour, during the summer and autumn of 1862, 
on and near the Rocky Mountains, in Colorado Territory, lat.39 -41 , by Asa Gray. Proc. 


PSEUDOGNAPHALIUM AUSTROTEXANUM (ASTERACEAE: 
GNAPHALIEAE), A NEW SPECIES FROM SOUTHEASTERN 
TEXAS AND ADJACENT MEXICO 


Guy L.Nesom 
Botanical Research Institute of Texas 
09 Pecan Street 
Fort Worth, TX 76102-4060, U.S.A. 


ABSTRACT 
Pseudognaphalium austrotexanum rs sp. nov., is described from collections in eleven counties of 
southern and southeastern Texas and from one locality in Nuevo Leén, Mexico. The new species is 


similar to Pseudoyraphaliam vi viscosum, his occurs in ae aucune n Texas and widely in Mexico 


and Central A lar stems, non-decurrent leaves, smaller, 
fewer-flowered heads and keeled, apically rounded, ane phyllaries. 


RESUMEN 
Se describe Pseudognaphalium austrotexanum Nesom, sp. nov, a partir de recolecciones en once 
condados del sur y sureste de Texas y de una localidad en Nuevo Leon, México. La nueva especie es 
similar a Pseudognaphalium viscosum, del suroeste de ae y muy comun en México y América 


entral; P austrotexanum se diferencia por sus tallos lulares, hojas no decurrentes, capitulos 
gl 


mas pequenios y con menos flores, y filarios a ee apiculados y apicalmente redondeados. 
REQ y y q Pp y 


In study of Gnaphalieae of Mexico and adjacent United States, various undescribed 
taxa and range extensions have come to light. The present paper reports a new 
species whose geographic range is centered in near-coastal and adjacent inland 
localities in southeastern Texas, some localities more than 100 airline miles from 
the coast (Fig. 2). One collection of the new species was made from the out- 
skirts of the city of Monterrey, Nuevo Leon, Mexico. 


Pseud hali t Nesom, sp. nov. (Fig. 1). TyPE: UNITED STATES. 
TEXAS. Jim Wells Co. 10.1 mi S of Alice, railroad right-of-way, Santa Gertrudis Divi- 
sion of King Ranch, open brush on light brownish-gray, clayey, sand loam, 24 Nov 
1954, see 542108 (HOLOTYPE: TEX!; ISOTYPE: SMU}). 


Dp q L | ] s ey as ‘ Ve . } ] 
I A mngere | 
“1 | een (eee | ] Joolayil 4 1] ] ne (eco ines ae 4 Soe ene 
UCCULICL ntipu 
oO t fo | 
1: : “1 Piutt caer, Le Sgt Cee | : a pee Pores Weare tees dy 
r 


Annual herbs from a woody taproot. Stems 3-7 dm tall, densely and closely 
white-tomentose-floccose, glabrescent, eglandular, strictly erect, unbranched 
until near the inflorescence or rarely more highly branched and “a large bushy 
herb” (fide comments on Runyon 1954). Leaves congested on very short intern- 
odes, continuing congested to immediately beneath the heads, the upper loosely 
to strictly ascending, the lower (moribund) loosely spreading to deflexed, the 


SIDA 19(3): 507 — 511. 2001 


" 
AND 


£5 
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BRIT.ORG/SIDA 19(3) 


NESOM, A NEW SPECIES OF PSEUDOGNAPHALIUM 509 


blades epetiolate, linear to linear-lanceolate, 2-5 cm long, 1-3 mm wide, rela- 
tively uniform in size, shorter immediately below the heads, subclasping but 
not basally ampliate, auriculate, or decurrent, apically acute, strongly bicol- 
ored, the upper surface green, densely stipitate-glandular, otherwise glabrate, 
the lower surface densely and closely white-tomentose, the margins revolute, 
sometimes closely sinuate. Capitula 4.5-5 mm high, in tight glomerules, the 
glomerules usually borne in a flat-topped inflorescence 10-30 cm wide, the 
ultimate branches 1.5-4 cm long; phyllaries narrowly ovate to oblong or ellip- 
tic, persistently wooly at the base, the inner with an elongate stereome, gland- 
otted near the apex, the outer ca. 2/3 as long as the inner. Outer, pistillate flo- 
rets (in Mexico, 46-)76-102; inner, bisexual florets (6-)8-Ll. Cypselae oblong, 
brownish-yellow, 0.5-0.7 mm long, 4-6 striate-ridged longitudinally, minutely 
papillate; pappus of fragile, separate, basally caducous barbellate bristles. 


Additional collections examined. MEXICO. Nuevo Leon: 7 mi SE of the Santa Catarina bridge in 
onterrey on the Montemorelos highway, gravel pit of the Cia. Fundidora de Fierro y Acero, 2000 ft, 
reddish limestone and caliche gravel, 11 Nov 1959, Johnston 4618 (TEX). UNITED STATES. Texas. 


Brazoria Co.: Brazoria National Wildlife Refuge, Bastrop Bayou, across bayou from N end of island, 
27 Oct 1967, one zi EX), 3.5 mi E of Angleton, 9 Oct 1934, Cory 11495 (GH). Brazos Co.: 
College Station, 3 Oct 1946, Parks s.n. (TEX). Brooks Co.: 5 mi N of Falfurrias, 28 Oct 1973, Everitt s.n. 


(SMU). Frio Co.: 13.6 mi ao of Dilly on Hwy 117, scrub pasture, reddish soil, 31 Oct 1981, Mahler 
9225 (BRIT). Harris Co.: Seabrook, 18 Oct 1939, Fisher 39038 (TEX). Jim Hogg Co.: 9 mi SW of 
Hebbronville, sandy loam, not plentiful, 10 Dec 1972, Bone s.n. (TEX). Kenedy Co.: near Rudolph, S of 
Norias, sandy knoll along highway, 3 Jan 1963, Correll 26926 (NCU, TEX); 4.4 mi S of Armstrong, 
sandy sacahuiste prairie at edge of caliche flat, 29 Nov 1954, Johnston aa (TEX); S of Armstrong 
on side of Hwy 96, sandy loam soil, scarce, only a few ape seen, 17 Oct 1938, Runyon 1954 (TEX-2 
sheets, US); Norias, highway right-of-way, dune sand, 4 Dec 1948, Tharp, Johnson, and Webster o 108 
(TEX). Matagorda Co.: Matagorda, 14 Oct 1936, Fisher ce (ARIZ, TEX, US). San Patricio Co.: 

W of Taft, near shore of Nueces Bay but above high tide level, soil not saline, 10 Nov 1956, Jones oe 
(SMU); ca. 2 mi SE of Odem, in sandy open pasture, locally abundant, 31 Oct 1959, Jones 3610 (TEX). 
Uvalde Co.: Uvalde, 1880, Palmer 550 (GH 


These plants are similar to eee viscosum (Kunth) A. Anderb. in 
general appearance and previously have been identified as that species. Plants 
of both species are taprooted annuals with white-tomentose, strictly erect stems 
mostly unbranched until the inflorescence, leaves linear to linear-lanceolate, 
strongly bicolored (green and glandular above, white-tomentose beneath), 
loosely to strictly ascending, crowded on short internodes and continuing to 
immediately below the heads, and basally subclasping but not strongly auricu- 
late, phyllaries silvery, thin-hyaline, and achenes minutely papillate. They are 
distinguished by the following contrasts: 
1. Capitula ca. 250-flowered, bisexual florets (13-)16-29, pistillate florets ca. 200- 

250; inner phyllaries narrowly acute at apex, not apiculate, not keeled or with a 

barely perceptible thickening along the midrib; stems stipitate-glandular; leaves 

not basally ampliate or subclasping, the lower usually decurrent 3-6(-10) mm 
Pseudognaphalium viscosum 


BRIT.ORG/SIDA 19(3) 


510 


fa 


}—4—_j 


| | if we quan 
a on {~~ 
jf ee Cae ly i 

4 1 a 


1 


|S — 
i s eas 


y 


P. viscosum A 
P. austrotexanum © 


Fic.2. Distrib 


NESOM, A NEW SPECIES OF PSEUDOGNAPHALIUM 511 


. Capitula ca. 100-flowered, bisexual florets (6-)8-11, pistillate florets (46-)76-102; 
inner ee ee) kounged or Lac uminels: distinct apiculate nom a thick- 


} claspin 
not deeiene pecudeananhalium austrotexanum 


Pseudognaphalium viscosum is distributed through Mexico and Central America 
and also apparently is common on the Caribbean island of Hispaniola. In the 
United States, it is known only from southwestern Texas, widely separated by 
distance, habitat, and climate from P. austrotexanum (Fig. 2). 

Plants of Pseud halium austrotexanum were included by Correll and 
Johnston (1970) as “rare in s.e. Tex.” with plants identified as Gnaphalium 
macounii Greene, a name that has sometimes been misapplied to Pseudognapha- 
lium viscosum. The other Texas plants of “G. macounii” “local in Rio Grande Plains 
and Trans-Pecos ... and the Llano region of the Edward Plateau”) are Pseudo- 
gnaphalium viscosum rather than Pseudognaphalium(Gnaphalium) macounii 
(Greene) Kartesz, which does not occur in Texas. The only known Mexican local- 
ity for P austrotexanum is separated by about 200 kilometers from the closest 
Texas site, although the geology and topology are generally similar. The Nuevo 
Leon site is alongside a major highway, and a fruit may have been accidentally 
transported (vehicle-dispersal) from Texas. 

The distinctiveness of Pseudognaphalium austrotexanum also has been 
recognized by several other botanists: Marshall Johnston (by notation); Billie 
Turner (pers. comm.); and Harvey Ballard (by annotation and pers. comm.). 


ACKNOWLEDGMENTS 
Iam grateful to the staffs at BRIT, GH, TEX, and US for help during visits, to 
Billie Turner for arranging for the fine illustration by Linny Heagy, and to an 
anonymous reviewer for helpful comments. 
REFERENCE 


Corrett, D.S. and M.C. JoHNsTON. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner, Texas. 


512 BRIT.ORG/SIDA 19(3) 


Book REVIEW 


Kay YATSKIEVYCH. 2002. Field Guide to Indiana Wild Flowers. (ISBN 0-253-21420- 
3, pbk.). Indiana University Press, 601 North Morton Street, Bloomington, 


IN 47404-3797, U.S.A. (Orders: i li edu, 800-842-6796, 812-855- 
4203, 812-855-8507 fax). $17.95, 372 pp, 640 color photos, line drawings, 5 
1 Ole eo Bae 


How should a book review be done about a subject (Wild Flower Field Guides) that has seen a great 
revival during the past few decades? It seems to me that the importance of field guides goes far be- 
yond their immediate practical usefulness a identifying a species of plant that oa attract ones 

‘icance of wild flowers and 


interest for a short moment. Her { my feelings about the signi 


Kay paces s boo 
seen spare ai come and BO) for neany nly years in Indiana, and having contem- 
are ee | ever-changing complex phe- 
nomena for at least 10, 000 years in the glaciated parts of indiana, | personal know that the places 
on earth where these flashes of springtime brilliance that I have witnessed in the virgin beech-maple 
and oak- aa forests in some of the State parks and State poe In diana are quickly vanishing. 
at if an unexpected, terrible disaster such as a tornado, earthquake, fire, explosion, or even 


a terrorist act destroyed one of the fine buildings on the campus of Indiana University or Purdue 
University? By using the blueprints that were used to build the original structure, highly sophisti- 
cated, well-trained Indiana architects, engineers, and other scientists ha ey up a ae 


ee pani buil een: exactly asit was, even to the { the 
I pr ope rties sa ps material oe eee rpction Soil scientists and pele 
tot ldd 1344 f } 
5 5 } t ) the ground W here the build 
ing stood. 
lar q ] ] ] } ] Jiyet ld ] 


But what if human ¢ 
a 


L vt I [ 
] 


ment | a 1 or 
p eae o 


L t I 
pristine eco-habitats of beech-maple and oak-hickory forests? Where could the dynamic structural 
details of these rapidly vanishing, primeval treasures be found? 
Where are the brilliant, highly sophisticated, well-trained architects, engineers, and other sci- 


entists, with great library and internet resources, who could draw up the plans for a virgin beech- 
maple forest that would be exactly like it was, even to the minutest detail of the myriads of unseen 
structural features represented by the millions of microscopic living creatures, before it was care- 
lessly dest 

Kay eae has produced a field guide to 1,564 pieces (wild flower species) that might fit 


into the dynamic ESUEUCI UTE of these Best master DISCS Just as the architects and engineers have their 
technical pieces that go into their structures, Kay Yatskievych 
has eraercdae us in 374 pages an easy to carry, detailed, and easy to use guide for naming and talking 


about many traits of these 1,564 species. Because of the excellent‘ flower finder” and other line draw- 
f 


ings, color photos of examples of all genera, and clear word descript important traits of each 
species, her guide will help ae lovers of all kinds know these “ ee of the puzzel of the forest” in 
much greater detail. Hopefully, some of the younger users of this book will be inspired to dig much 
deeper into the forests and uncover more of ee webs that help tie the dynamic struc- 
ture together—Joe F Hennen, Resident Research Associate, Botanical Research Institute of Texas, 509 
Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 


SIDA 19(3): 512.2001 


TAXONOMIC NOTES ON KEYSSERIA AND PYTINICARPA 
(ASTERACEAE: ASTEREAE, LAGENIFERINAE) 


Guy L.Nesom 


Botanical Research Institute of Texas 
Pecan Street 
Fort Worth, TX 76102-4060, U.S.A. 


ABSTRACT 


ia (9 


New Guinea, Celebes, and Borneo) and the Ha- 
waiian islands (3 species). The Hawaiian species are formally segregated here as Keysseria sect. 
Sandwicactis Nesom, sect. nov., on the basis of their bisexual and fertile disc florets (vs. functionally 


Keysseria is divided between I 


staminate in Indonesia) and other ec differences in ray and dise corollas. Keysseria 
penn A. rey ages to Vanua Levu Island, Fiji, is excluded from Keysseria and formally 


(A 


Gray) Nesom, comb. nov. A nomenclatural update and sum- 

aes are provided for /Pytinicanoe. whose other two species are New Caledonian. The genera of 

Lageniferinae, including nce ia and Pytinicarpa, are divided into seven “core genera” and five 
vid 


“peripheral genera”, a key g is provi 


RESUMEN 


Keysseria esta dividido entre Indonesia (9 especies en Nueva Guinea, Celebes, y Borneo) y las islas 
Hawai (3 especies). las especies Hawaianas se segregan formalmente aqui como Keysseria sect. 
Sandwicactis Nesom, sect. nov, en base a sus flésculos bisexuales y fértiles (vs. funcionalmente 
estaminadas en Indonesia) y otras diferencias morfologicas en las corolas centrales y periféricas. 
Roser pickerigy A. my, encore ce 2 oe Vanua Levu, Fiji, se excluye de Keysseria y se 

i(A. Gray) Nesom, comb. nov. Se of rece una puesta 


al dia resumen nomenclatural para Pytinicarga a Nueva Caledonia. 
Los de Lageniferinae, que incluyen Keysseria y Pytinicarpa, se araden en siete ete y cinco 
“géneros periféricos”; se ofrece una clave para los géneros centrales. 


Keysseria Lauterbach (Lauterbach 1914) is a genus of 12 species divided be- 
tween Indonesia (9 species plus infraspecific taxa; Koster 1966) and Hawaii (3 
species; Mill 1990, 1999 - see below). All of the Indonesian species occur in New 
Guinea; K. radicans (F Muell.) Mattf.also is known from Celebes and K. gibbsiae 
(Merrill) Cabrera ex Steenis also occurs on Mt. Kinabalu in Borneo. The genus 
was treated as a member of Astereae subtribe Lageniferinae by Nesom (1994a, 
2000) anda recent morphological and taxonomic overview is available (Nesom 
2000). 

The three Hawaiian species were transferred to Keysseria from Lagenifera 
Cass. by Cabrera (1966). Mill (1990, 1999) maintained them in a broadly con- 
ceived Lagenifera, indicating that Keysseria was to be regarded as a synonym. 
Swenson and Bremer (1994) also explicitly regarded Keysseria as a synonym of 
Lagenifera. Perspective on morphological distinctions among these genera is 
provided below in the key to the “core genera” of Lageniferinae. 


SIDA 19(3): 513 — 518. 2001 


a 


514 BRIT.ORG/SIDA 19(3) 


The Hawaiian plants of Keysseria are similar to the Indonesian ones in 
habitat, habit, and morphological details but differ as follows: disc florets with 
fertile ovaries (vs. functionally staminate), ray corollas usually deeply and asym- 
metrically 2-4-lobed at the apex (vs. apically entire), and disc corollas some- 
times 5-lobed (vs. consistently 4-lobed). Mill (1990) noted that the Hawaiian 
species apparently are “from one founder,” in this summary of supraspecific 
taxonomy of Keysseria, they are treated as a separate section. 


Keysseria Lauterbach, Repert. Spec. Nov. Regni Veg. 13:241. 1914. Myriactis Less. 

subg. Hecatactis F Muell., Trans. Royal Soc. Victoria 1, 2:13. 1889. Hecatactis (F- 

Muell) F Muell. ex Mattf., Bot. Jahrb. Syst. 62:407. 1929. Keysseria sect. Hecatactis 

(FE Muell.) Mattf., Bot. Jahrb. 68:250. 1937. TyPE: Keysseria papuana Lauterbach (= 
Keysseria radicans (E Muell.) Mattf.). 


The nine Indonesian species constitute sect. Keysseria. 
Keysseria Lauterbach sect. Sand wicactis Nesom, sect. nov. TYPE: Keysserid erici 


(C.N. Forbes) Cabrera. 


H {] 1 a hs es s f, 144, 


Syne a Keysseria sect. K 


corollis radii plerumque ad apicem 
2-4-lobatis, et c -orollis disci il aliqu ando 5-lobatis. 


Species included: K. erici, K. maviensis (H. Mann) Cabrera, and K. helenae (CN. 
Forbes & Lydgate) Cabrera. 


The “core genera” of Lageniferinae 
Lagenifera Cass. and genera similar to it have been grouped together as the 
subtribe Lageniferinae (Table 1) (Nesom 1994a). Three genera have been added 
to the Lageniferinae since the recent classification and overview: Lagenocypsela 
U. Swenson and K. Bremer (Swenson & Bremer 1994); Pytinicarpa Nesom 
(Nesom 1994b); and Pappochroma Raf. (Forbes & Morris 1996; Nesom 1994, 
1994d, 1998; synonyms = Lagenopappus Nesom and Lagenithrix Nesom). The 
12 genera and approximately 71 species of Lageniferinae are distributed from 
India to southeast Asia, Australia, and various Pacific islands, except for the 
nine species of Lagenifera and Myriactis native to South America and Central 
America. Plants of the seven “core genera” of Lageniferinae (Table 1) are prima- 
rily characterized (with exceptions) by a herbaceous habit, leaves mostly in a 
basal rosette, heads borne singly on scapose or scapiform stems or few ina loose 
panicle, pistillate florets with reduced lamina and in several series, a tendency 
to produce functionally staminate disc florets, and flat, 2-nerved, epappose 
cypselae, commonly with a glandular apex or beak. The remaining five “pe- 
ripheral genera,” while similar in geography, may not be closely related to the 
others; they differ in combinations of various features, particularly in subterete, 
multinerved cypselae. 

A key to the “core genera” provides perspective for the positioning of 
Keysseria and its distinction within the group. 


NESOM. TAXONOMIC AND DYVTINICARDA 


TIHINILARFA 515 


Taste 1. Composition of subtribe Lageniferinae. 


“core genera” 


Keysseria Lauterbach (12 species; Hawaiian Islands and Indonesia-New Guinea, Celebes, and 
Borneo) 

Lagenifera Cass. (15 species; Australasia and South America) 

Lagenocypsela U. Swenson & K. Bremer (2 species; New Guinea) 

Myriactis Less. (ca. 19 species; Central America, South America, southeast Asia, Indonesia) 

Pappochroma Raf. (9 species; Australia and Tasmania). 
Synonyms = Lagenopappus Nesom and Lagenithrix Nesom 

Piora Koster (1 species; New Guinea) 

Solenogyne Cass. (3 species; Australia, New Zealand) 


“peripheral genera” 


Pytinicarpa Nesom (3 species; New Caledonia and Fiji 

Rhamphogyne S. Moore (1 species; Rodrigues Island) 
Rhynchospermum Reinw. (1 species; Japan, Korea, Ryukyus, Formosa, Malaysia, and India) 

Sheareria S. Moore (2 species; southern and southeastern China) 

Thespis DC. (3 species; southeast Asia) 


a 


ee of barbellate bristles. 


Pappochroma 
te 


. Disc corollas infundibular, the tube aunty Opening into a broad limb, 4-lobed 
or 5-lobed; flowering stems leafy or perennial and arising from 

a thick rhizome or a procumbent stem (Keysseria Bioin and Myriactis p. p.) or 
annual from a thin base (Myriactis s. str.). 

2. Disc corollas tubular-funnelform, gradual ning into the limb, 5-lobed; flow- 
ering stems scapiform; plants on lived, ine arising from a barely evi- 
aes caudex clos 

Phylla lanceolate to linear, acute to acuminate; capitula radiate— 
lamina of ray florets strongly developed; cypselar beak well-developed, 
glandular. 

. Phyllaries elliptic-obovate to oblong, bluntly rounded to obtuse; capitula disci- 

form—lamina of ray florets rudimentary or absent; eal beak pronounced 

to rudimentary or absent, the fruits mostly eglandul 

4, wea beak conspicuous; ovaries of disc florets pen absent;leaves 
tire. 


Lagenifera 


Ww 


BgEnOeypseie 


+ 
q 


: ° pselar or ies of disc florets present, sterile 
leaves toothed. Solenogyne 
5. Annual or perennial herbs, not aromatic; leaves thin-herbaceous, flat- 
margined, sometimes SUeer sells a not ees dilated or sheathing, 
the basal often persistent but th tinuing unreduced in size 
halfway to nearly completely up thes stem. Myriactis 


Sa) 


. Perennial herbs to subshrubs or shrubs, at least some species aromatic; 
leaves thickened to coriaceous, usually with revolute or deflexed mar- 
gins, basally dilated and sheathing, evenly arranged along the stems or 
in rosulate clusters 
6. Leaves of current year in a rosulate or subrosulate cluster, the heads 


516 BRIT.ORG/SIDA 19(3) 


on long, scapiform peduncles; anthers with a short apical appendage. 


Keysseria 
6. Leaves densely and more or less evenly arranged on the stems, the 
heads without an evident peduncle; anthers without an apical 
appendage. Piora 


Transfer of Keysseria pickeringii to Pytinicarpa 

Keysseria pickeringii, originally described by Gray (1861), is endemic to Vanua 
Levu Island, Fiji. Smith and Carr (1991, p. 302) noted “it is not possible to sug- 
gest a New Guinean relative of Keysseria pickeringii, which is remarkable for 
its very small heads and its costate achenes.” This species, which has a basal 
rosette of narrow leaves loosely villous-strigose on both surfaces, 
monocephalous and minutely bracteate, scapiform stems, small, radiate heads, 
convex receptacles, functionally staminate disc florets, and subcylindric, 
eglandular cypselae 4 mm long with 8-10 longitudinal, strongly raised nerves 
and a truncate, epappose apex, is a member of the recently described genus 
Pytinicarpa Nesom (Nesom 1994b), which originally included two species from 
New Caledonia. Some features of Pytinicarpa (geography, solitary capitula on 
scapose stems, functionally staminate disc florets, epappose cypselae) suggest 
that it shares close ancestry with genera of Lageniferinae, but the conical-con- 
vex receptacles, ray florets in a single series, and multinerved, subcylindric 
cypselae are unusual in that subtribe. 


Pytinicarpa pickeringii (A. Gray) Nesom, comb. nov. Lagenophora Paes A. 
Gray, Proc. Amer. Acad. Arts 5:121. 1861. Keysseria pickeringii (A. Gray) Cabrera, 
Blumea 14:307. 1966. Type: FUJI: Vanua Levu Island, Mathuata Mts., ica. Jul 1840], 
Wilkes Expl. Exped. s.n. (HOLOTYPE: US). 


Pytinicarpa pickeringii differs from both New Caledonian species in its cypselar 
surface, which is minutely papillate, the center of each cell abruptly raised into 
a sharp point. In the New Caledonian species, these epidermal cells are simi- 
larly quadrate but the whole surface of each cell is convex and the cypselar 
surface has a “frothy” appearance. The relatively broad leaves of P. pickeringii 
are more like those of P. sarasinii (see below) rather than the linear leaves of P. 
neocaledonia, but those of P. pickeringii are obovate-spatulate, abruptly nar- 
rowed toa petiolar base, and the margins are shallowly crenulate-mucronulate 
(vs. oblanceolate, without a distinct petiolar portion, with margins coarsely 
toothed only near the apex). 

There is some indication that the narrow “coronal rim” of the cypselae in 
Pytinicarpa pickeringii might have been sticky when fresh, which is a general 
feature of the Lageniferinae. This rim is similar to that illustrated for the cypsela 
of P sarasinii (Nesom 1994b), where stickiness was not evident. 


N lature of New Caledonian Pytinicarpa 
Lagenifera neocaledonica S. Moore predates Brachyscome sarasinii Daniker 


NESOM, TAXONOMIC AND PYTINICARPA 517 


which was the basionym for one of the two original species in Pytinicarpa (Nesom 
1994b). Cabrera (1966), in his list of species excluded from Lagenifera, recog- 
nized that these two names refer to the same species. But because the epithet 
“neocaledonica” already exists in Pytinicarpa for the second species, P sarasinii 
is the correct designation of the broad-leaved New Caledonian species named 
by Moore and Daniker. The two New Caledonian species are as follows. 


1, Pytinicarpa sarasinii (Daniker) Nesom, Phytologia 76:138. 1994. Brachyscome 
sarasinii Daniker, Mitt. Bot. Mus. Univ. Ztrich 142:479. 1933. TypE: NEW 
CALEDONIA: Am obern Abhang des Mut. Koniambo bei der Mine Boume I, 
zerstreut im lichten Gebusch an felsigen stellen, 14 Jan 1925, Daniker 880 (HOLO- 
TYPE: Z). 


ge ets neocaledonica S. Moore, J. Linn. Soc. Bot. 45:345. 1921 (non Pytincarpa 
(Brachyscome) neocaledonica (Guill.) Nesom 1994). Type: NEW CALEDONIA: 
oo pau red serpentine earth, 2500 ft, rare, 2 Dec 1914, R.H. Compton 2305 (HO- 
LOTYPE: BM, photocopy!). 


2. Pytinicarpa neocaledonica Sia occ 76:138. 1994. Brachyscome 
neocaledonica Guill., Bull. . Bot. France 84:61. 1937. SYNTYPES: NEW 
CALEDONIA: Gatope, ee 2823 ae GH!); Néhoué, Pancher 425 and 
Deplanche 425. 


ACKNOWLEDGMENTS 


I am grateful for help from the staffs at GH and US during recent visits, the 
staff of the Couch Botanical Library (UNC-Chapel Hill) in obtaining literature, 
John Strother and Mark Wetter for comments on the manuscript, and Roy Vick- 
ery of the BM staff in sending a photocopy of the type of Lagenifera 


eee a 
MTLEOCGALEGOFLCA 


REFERENCES 

Caprera, A.L. 1966.The genus Lagenophora (Compositae). Blumea 14:285-307. 

Forses, S.J. and D.. Morris. 1996. A review of the Erigeron pappocromus Labill. complex. 
Muelleria 9:175-189. 

Gray, A. 1861.Characters of some Compositae in the collection of the United States South 
Pacific Exploring Expedition under Captain Wilkes, with observations, &c. Proc. Amer. 
Acad. Arts 5:114-146. 

Koster, J.T. 1966. Keysseria. The Compositae of New Guinea |. Nova Guinea, Bot. 24:597- 
608. 

LAUTERBACH, C. 1914. Neue Bergpflanzen aus Kaiser-Wilhelms-Land. Repert. Spec. Nov. Regni 
Veg. 13:239-250 

Mit, S.W. 1990 (1999, rev. ed). Lagenifera. In: Wagner, W.L., D.R. Herbst, and S.H. Sohmer. 
Manual of the flowering plants of Hawai'i. Vol. 1, pp. 329-331. Univ. Hawaii Press, Honolulu. 

Nesom, G.L. 1994a. Subtribal classification of the Astereae (Asteraceae). Phytologia 76:193- 
274. 


518 BRIT.ORG/SIDA 19(3 


ua 


Nesom, G.L. 1994b. Pytinicarpa (Asteraceae: Astereae),a new genus from New Caledonia. 
Phytologia 76:136-142. 

Nesom, G.L. 1994c. Taxonomic dispersal of Australian Erigeron (Asteraceae: Astereae). 
Phytologia 76:143-159. 

Nesom, G.L. 1994d.Pappochroma Rafin.is the correct generic name for Erigeron pappocroma 
Labill. Phytologia 76:426. 

Nesom, G.L. 1998 [2000]. Full constitution of the Australian genus Pappochroma Raf. 
(Asteraceae: Lageniferinae). Phytologia 85:276-279. 

Nesom, G.L. 2000.Generic conspectus of the tribe Astereae (Asteraceae) in North America, 
Central America, the Antilles, and Hawaii. Sida, Bot. Misc. 20:i-vili, 7-100. 

Situ, A.C. and G.D. Carr. 1991. Asteraceae. In A.C. Smith. Fl. Vitiensis Nova 5:254-320. 

Swenson, U. and K. Bremer. 1994. The genus Lagenocypsela (Asteraceae, Asteraceae) in New 
Guinea. Austr. Syst. Bot. 7:265-273. 


A NEW SPECIES OF SACCOGLOSSUM (ORCHIDACEAE) 
FROM THE HANS MEYER RANGE, NEW IRELAND, 
PAPUA NEW GUINEA 


N.H.S. Howcroft 


ENB Balsa |r try Str he one Project (ITTO) 
PNG POB 406 


Rabaul, East New Bri tain 614 
PAP W GUINEA 
howcroft@daltron.com.pg 


ABSTRACT 


A new species, Saccoglossum takeuchii, is described from the Hans Meyer Range in southern New 
land, Papua New Guinea. The Poveley is most nna related to S. lanceolatum L.O. Williams, but 
i the petals esser number of cilia-like stelids, 


the sommes enen on around the cli lrium, and by the differentl y shaped leaves. 
ABSTRACT (MELANESIAN TOK PISIN) 


Nupela kain orchid Saccoglossum takeuchii, ol bin kisim long Hans Meyer Range insait long Nu Ailan, 
Papua New Guinea, em ol bin deskraibim. Em I wankain long Saccoglossum lanceolatum L.O. Will- 


an 


iams. Dispela em minim olsem long sais na saip long petals, na namba long stelids na dentations 
bilong clinandrium, na saip long lip. 


INTRODUCTION 


The genus Saccoglossum is presently comprised of four taxa: S. lanceolatum L.O. 
Williams, S. maculata Schltr, S. papuanum Schltr, and S. verrucosum L.O. Will- 
iams. The latter species is recorded only from Irian Jaya whereas the others are 
known primarily from Papua New Guinea (PNG). O’Byrne (1994) suggested 
there are about 5 species and that some new ones could be expected from the 
PNG Highlands. In the orchid treatments for Vanuatu, the Solomon Islands, and 
Bougainville (Lewis & Cribb 1989, 1991) no records are provided for 
Saccoglossum. The new species extends the generic range to New Ireland and 
apparently marks the easternmost limit for the genus. The Hans Meyer species 
is most closely related to S. lanceolatum L.O. Williams but differs in the size 
and shape of the petals, the structure of the clinandrium, and toa lesser extent 
the shape of the leaves. 


Saccoglossum takeuchii Howcroft, sp. nov. (Fig. 1). Typus: PAPUA NEW GUINEA. 
RELAND PROVINCE: Hans Meyer Range, 27 Jan 1994, W. Takeuchi & J. Wiakabu 
9601, 27 Jan 1994 (HOLOTYPUs: LAE). 


S 1 Sac ‘coglossum 


isad bases 
] : are 


cubirimeatiis clinandrio stelidiis pauci | iversis ] isp ile disting 


SIDA 19(3): 519 — 521. 2001 


520 BRIT.ORG/SIDA 19(3) 


Fic. 1, Saccogl keuchi(d from tyy terial). A. Plant habit. B. Flower, f iew. CD. Flower lateral views. 
E-F. Dorsal and lateral sepals.G—H. Petals. |!.Labellum, lateral view.J. Col itl her | 
lateral view. L. Anther. M-N. Pollen, lateral and oblique ventral (scale as for L). 


K.Column stelids 


Epiphyte, erect to suberect, up toca. 7 cm tall. Rhizome creeping, concealed by 
sheaths. Pseudobulbs obovoid, 1.25 to 1.5 cm long, up to 0.75 cm in diameter, 
unifoliate, surface smooth, yellow green. Leaf blade elliptic to lanceolate, 4.5- 


HOWCROFT, AWEW SFELILS Ue 521 


5.5 x L.0-1L.7 cm, apically obtuse, base duplicative, sessile, surfaces glossy green, 
texture firm, costa impressed above. Inflorescences more or less fasciculate, lateral 
from base of the pseudobulb, erect, single flowered, peduncle short, 3.1-3.8 cm 
long, 0.2-0.3mm in diameter, provided witha single sheath, glabrous. Perianth 
purple, glabrous; dorsal sepal ovate-lanceolate, 1.05 x 0.33-0.5 cm, slightly 
hooded, subacute; lateral sepal obliquely ovate, 1.18 x 0.66-0.70 cm, apiculate, 
glabrous; petals broadly obovate to obliquely quadrate, 0.28 x 0.24 cm, apically 
truncate, attenuate at base, glabrous, median nerve slightly raised and thick- 
ened; labellum saccate, 0.55-0.66 cm high, 0.55 cm broad, 0.78 cm long, front 
part incised; column arched, glabrous, thickened to the base and with a short, 
thick foot, ca. 0.24-0.25 cm long: clinandrium in two segments on either side, 
the lower with one cirrhose stelid on each side, the upper subpalmate with a 
subulate margin; anther cordate in outline witha raised, thickened keel, 0.07 x 
0.08 cm; ovary with glabrous pedicel, ca. 1.3 cm long. 

Distribution —Known only from the Hans Meyer Range in New Ireland, 
Papua New Guinea. 

Habitat—Mossy montane forest on trees at ca. 1175 m elevation. 

Etymology.—The new species has been named for Wayne Takeuchi, the prin- 
cipal collector of this orchid who submitted the type specimen to me for iden- 
tification, together with many other orchids from New Ireland. 

Notes.—The description and illustration of Saccoglossum takeuchii was 
made from spirit material. The new species differs in flower color from its con- 
geners (based on Takeuchi’s fieldnotes) and the floral parts are also smaller than 


those from previously described species. 
ACKNOWLEDGMENTS 


| wish to thank Wayne Takeuchi and Joe Wiakabu for providing the specimen 
of Saccoglossum takeuchii and John Pipoly for his help with the Latin diagno- 
sis. To John Ohana a very special thanks for his assistance with the Melanesian 
Tok Pisin abstract. 


REFERENCES 


Lewis, B.and P. Criss. 1989. Orchids of Vanuatu. Royal Botanic Gardens, Kew. 

Lewis, B.and P.Crigs. 1991.Orchids of the Solomon Islands and Bougainville. Royal Botanic 
Gardens, Kew. 

O'Bryne, P. 1994. Lowland orchids of Papua New Guinea. SNP Publishers, Singapore. 

SCHLECHTER, R. 1911-14. Die Orchidaceen von Deutsch-Neu-Guinea. Fedde Repert. Spec. 
Nov. Regni Veg., Beih. 1:1-1079. English translation 1982, D.F. Blaxell, ed. The Australian 
Orchid Foundation, Melbourne 

SCHLECHTER, R. 1928. Figuren atlas zu den Orchidaceen von Deutsch-Neu-Guinea. 


522 BRIT.ORG/SIDA 19(3) 


Book REVIEW 


DENI Bown. 2000. Aroids: Plants of the Arum family (ed. 2). ISBN 0-88192-485-7, 
hbk.). Timber Press, 133 SW Second Ave., Suite +50, Portland, OR 97204-3527, 
U.S.A. (Orders: www.timberpress.com, 800-327-5680, 503-227-2878, 503- 
227-3070 fax). $34.95, 468 pp, 108 color photos, 50 line drawings, 6" x 9". 

Contents.—Foreword to the Second Edition, Peter C. Boyce; Foreword to the First Edition, Simon Mayo; 

Preface; sella: seat Introduction. 

1. Variations on a theme: What are aroids and where do they as 
2. Of tails and traps and the underworld: Mechanisms of reproduc 
3. Woodlanders: Species of temperate woodland and higher Sanus of the tropics and 


oO. 


subtropics. 
4. Aquatics and Amphibians: Species of wetlands and water. 
5. A place in the sun: Species of arid and seasonally a0 —e 
6. In the shadows: Species of the monies saueueos 
lam ber 4 


7. Towards the light: mopic 
8. The titans: Giant s species of the inopiee 
9. An acquired taste: Aroids as food plants. 
10. Acids and aes = eet and ee of aroids. 
ailed notes on how to do it, arranged by genus and species). 


Aroids in 


= Checklist of aroid genera hands overview of classification, with number of species, distribu- 
tion, habit, and ecology for each genus). 

* Glossary 

* References 

* Index 
Its an amazing family—so much structural diversity and so much biological diversity—with about 
109 genera and 3200 species. Philodendron (Philodendron), anthurium (Anthurium), green dragon 
and jack-in-the-pulpit (Arisaema), dumb cane (Dieffenbachia), duckweeds Pera ane pReauiyes), 
monstera (Monstera), pothos (Pothos and relatives), caladium (Caladium), golden 
skunk cabbage (Symplocarpus), calla (Calla), taro (Colocasia), water ace (Pistia), crypocoryne 
and re esa (Cryptocoryne and Lagendndra), titan arum } n lily 
(Zantedeschi ven those with only a passing interest in horticulture ay west of nee names 


And even the ae oriented see these plants in aquaria, malls, and shady, highly tamed eaten 
“Sweet flag” is out of the family (Acorus calamus, “The Aroid that never was,” pp. 124-126) but still 
given discussion; the duckweeds are in (Lemna et al., “Deceptively delicate drifters,” pp. 100-102). 
The new edition of Bown’s “Aroids” conveys so much interesting information with as much 
clarity and wit as anything botanical I’ve ever read. The beautiful photos are integral. The author 
intends the book as a oe introduction to the family, “readable rather than aly referable” t dut 
] 


| ” 


“sulficie ntly i [ enthusiasts who already grow these )] and as 
an offering “lor all those who find structure and design in naure an endless fascination.” You may 
want one copy for yourself, one fora friend, and one for the local public acne! npr) The Genera of 


Ardceae (SJ. Mayo et al. 1997, Kew Gardens) is more technical and more tecl inically organized | buta 
natural companion to the Bown volume.—Guy L. Nesom, Botanical Research Institute of Texas, 509 
Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 


SIDA 19(3): 522, 2001 


NOMENCLATURAL CHANGES IN PENNISETUM 
(POACEAE: PANICEAE) 


Joseph K.Wipft 


Pure Seed Testing, Inc. 
Ae) 449 
Hubbard, OR 97032 U.S.A. 


ABSTRACT 


During preparation of the account of Pennisetum Rich. for the Manual of Grasses for North America, 
it was determined that three taxa formerly treated unaer Cenc ucts L. required new combinations in 


Pennisetum: PB. setigerum (Vahl) Wipff, comb. nov, P. if (Hochst. & Steud. ex Steud.) Wipff; 


BP 


mb. nov, and P. somalensis (Clayton) Wipff, comb. nov. 


RESUMEN 
l | ion del inf de P Rich. para el Ma lof Grasses for North America, 
ee taxa tratados previamente como Ce nents L Hecesteabans nuevas combinaciones en 
ee P. setigerum (Vahl) Wipff, comb. nov, P. (I & Steud. ex Steud.) Wipff; 


comb. nov, y P. somalensis (Clayton) WipIf, éoiien nov. 


There has been considerable debate concerning the generic limits of Cenchrus 
L. and Pennisetum Rich. The predominant character traditionally used to dis- 
tinguish the two genera is fusion, or lack of fusion, of the bristles (e.g., Henrard 
1935; DeLisle 1963; Clayton & Renvoize 1982; Filguerias 1984, Clayton & 
Renvoize 1986; Watson & Dallwitz 1992), but its variation across the two genera 
is continuous, making the placement of numerous species arbitrary (Webster 
1987). DeLisle (1963), though basing his treatment on the traditional criteria of 
bristle fusion, recognized the difficulty in the interpretation of this character, 
and refined his generic criteria with the addition of the follow characters. 
Pennisetum has bristles that are seldom more than 0.2-0.4 mm wide, and the 
base of the fascicle rarely exceeds 0.5 mm in width; whereas in Cenchrus, “the 
spines usually 0.5 mm or (more) wider, and are generally united for a consider 
able distance above the base of the bur, with the base itself usually at least 1.5 
min in diameter. These characteristics, although admittedly arbitrary, are used 
in the present treatment of the genus Cenchrus” (DeLile 1963, p. 269). The in- 
crease in base diameter is probably a structural response to the fusion and thick- 
ening of the bristles and is closely correlated with bristle fusion. The more fu- 
sion and thickening of the bristles that occurs the wider the base of the fascicle 
must be to support them. 

Filgueiras (1984), using criteria similar to that of DeLisle (1963), separated 
the two genera as follows. Cenchrus has fused bristles, at least basally, forming 
a basal disc at least 1 mm in diameter, whereas Pennisetum has bristles to the 


SIDA 19(3 


— 


523 — 530, 2001 


524 BRIT.ORG/SIDA 19(3) 


base, not forming a disc. Webster (1987) used only the presence of this disc or 
callus to separate the genera; in Cenchrus the callus is pronounced, with the 
apex flared to form a discoid receptacle, whereas in Pennisetum the pronounced 
callus is absent or, when present not differentiated as in Cenchrus species. Web- 
ster went on to say that this character allows for the separation of the species 
along traditional grounds, which is based on bristle fusion. In addition to fu- 
sion, Clayton and Renvoize (1986) and Watson and Dallwitz (1992) also men- 
tioned that Cenchrus usually had ‘spiny’ bristles. However, Chase (1920) sepa- 
rated the two genera by bristle type, in addition to fusion: Pennisetum has 
bristles that are usually very slender, not rigid, and are free or rarely united at 
the very base; whereas Cenchrus has rigid bristles that are united below. 
Webster (1988) stated that even within a number of species it is open to 
interpretation as to whether the bristles are fused or the callus flared. In regards 
to bristle fusion, Pennisetum ciliare (L.) Link is extremely variable and has been 
treated in both Cenchrus and Pennisetum. Hignight et al. (1991) evaluated 800 
accessions of P.ciliare collected in South Africa and selected accessions based 
on extreme differences in morphology, including differences in bristle fusion. 
Thirteen of the most diverse morphological types were studied for morphology, 


cytology, and fertility. Five of these diverse morphologically types were used in 
hybridization studies with a sexual genotype (Bashaw 1969) of P.ciliare. Though 
they found most accessions to have at least some fusion, two of the accessions 
studied had a complete lack of bristle fusion. These plants were verified at Royal 
Botanical Gardens, Kew (K) to be P.ciliare. Hybridization studies with the sexual 
genotype showed a close relationship between the plants. Some of the F) prog- 
eny from the hybridization studies segregated for union of bristles similar to 
the bristle fusion found in Cenchrus setigerus Vahl. Hignight et al. (1991) con- 
cluded, “that bristle union is an arbitrary character that varies with genotype 
and is unreliable for the taxonomic classification of buffelgrass [P. ciliare].” 

Read and Bashaw (1969) hybridized the same sexual accession of P ciliare 
with an apomictic accession of C. setigerus. The resulting progeny represented 
a complete intergradation in morphology between the parents. Read an 
Bashaw concluded that the chromosome homology and cross-compatability 
of P-ciliare and C. setigerus, plus the high fertility and morphological intergra- 
dation, observed in the F) progeny provided overwhelming evidence of a very 
close relationship between the species and concluded that they belonged in the 
same genus. 

Sohns’(1955) examination of fascicle organization in eight species of Cenchrus 


and six of Pennisetum suggests an additional differentiating character: whether 
the axis of the fascicle is prolonged asa, usually prominent, bristle (Pennisetum) 
or terminates into a spikelet and is not prolonged (Cenchrus). Unfortunately, 
the prolonged bristle in Cenchrus setigerus and P. clandestinum, although 
present, is less prominent than in the other species of Pennisetum studied and 


— 


WIPFF, NOMENCLATURAL CHANGES IN PENNISETUM 525 


was overlooked by Sohns. This may be why most subsequent taxonomists con- 
sidered the presence or absence of the prolonging bristle not to be of generic 
significance. Also, this character has historically been evaluated as a secondary 
character in conjunction with bristle fusion (e.g., DeLisle 1963), which is known 
to be arbitrary in its separation of the genera, and would explain why Sohns’ 
character has appeared to be of little taxonomic value. 

Avdulov (1931) and Nunez (1952) reported that the genus Cenchrus has a 
base number of x = 17. Pohl (1980) used chromosome base number as part of his 
generic criteria. He distinguished the two genera on the following characters. 
Cenchrus has inner br istles that are spine-like or pungent, are — retrorsely 
scabrous, and usually have a base chromosome number of x = 17: whereas 
Pennisetum has bristles that are not spine-like or pungent and are antrorsely 
scabrous; and have base chromosome numbers of 5, 7, 8, or 9. However, despite 
these observations, Pohl later (Pohl & Davidse 1994), without explanation, fol- 
lowed Delisle (1963), Filgueiras 1984), and Clayton and Renvoize (1986) in his 
generic concept of Cenchrus and Penniseum. 

From the examination of specimens of the following species of Cenchrus and 
Pennisetum: Cenchrus agrimonioides Trin., C. biflorus Roxb., C. caliculatus Cav. 
C. distichopyllus Griseb., C. brownii Roem. & Schult., C. echinatus L., C. 
gracillimus Nash, C. longispinus (Hack.) Fern., C. pilosus Kunth, C. palmeri Vasey, 
C. platycanthus Anderss., C. spinifex Cav. C. tribuloides L., Pennisetum advena 
Wipff & Veldkamp, P. alopecuroides (L.) Sprengel, P. annum Mez, P. 
bambusiforme (Fournier) Hemsley, P basedowii Summeth,, P. chilense (Desv.) 
Jackson, P. ciliare (L.) Link, P clandestinum Hoch. ex Chiov. P complanatum 
(Nees) Hemsley, P crinitum (Kunth) Sprengel, P. distachyum Rupr, P. divisum 
(Gmel.) Henr, P. domingense (Sprengel) Sprengel, P durum Beal, P. elymoides (F 
Muell.) Gardn., P. flaccidum Munro ex Griseb., P frutescens Leeke, P glaucum 
(L.) R. Br, P hohenackeri Steud., P hordeoides (Lam.) Steud., P intectum Chase, P 
karwinskyi Chase, P. lanatum Klotzsch, P latifolium Sprengel, P macrostachys 
(Brong.) Trin. P. macrourum Trin., P. massaicum Stapf, P mezianum Leeke, P 
montanum (Griseb.) Hack., P nervosum (Nees) Trin., P occidentale Chase, P 
orientale Rich., PR. pauperum Nees ex Steud., P. pedicellatum Trin., P. 
pennisetiformis (Hochst. & Steud. ex Steud.) Wipff, P peruvianum (Dol) Trin., 
P. petiolare (Hochst.) Chiov, P. polystachion (L.) Schultes, P. prieurii Kunth, P. 
prolificum Chase, PB purpureum Schumach., P. ramosum (Hochst.) Schweinf., P. 
rigidum (Griseb.) Hack., P rupestre Chase, P sagittatum Henr, P setaceum (Forsk.) 
Chiov, P. setigerum(Vahl) Wipff, P somalenisis (Clayton) Wipff, P sphacelatum 
(Nees) Dur. & Schinz, P. ia Fresen., P. tempisquense Pohl, P thunbergii 
Kunth, P. tristachyon (Kunth) Sprengel, P. unisetum (Nees) Benth., P. villosum 
R.Br. ex Fresn., P vulcanicum Chase, and P. weberbauri Mez; as well as cytologi- 
cal examinations of 9 species of Cenchrus and 26 species of Pennisetum, and in 
addition to the cytological work already published (for a review see Jauhar 1981; 


526 BRIT.ORG/SIDA 19(3) 


Wipff 1995; Schmelzer 1998), it is concluded that the generic ue aee that 
Pohl adopted in 1980 is correct phylogenetically. The degree of fusion of the 
bristles is generally liable at the generic level and should not be used as the 
primary character in separating the two genera. 

The following characters are considered the most important in delineat- 
ing the two genera: 

Pennisetum: |) bristles are not spine-like or pungent and are antrorsely scabrous 
(one South American species is both antrorse/retrorse); 2) the axis of the fas- 
cicle is prolonged as a, usually, prominent bristle; 3) inner bristles free or fused; 
and 4) have base chromosome numbers of 5,7,8, or 9. 

Cenchrus: |) inner bristles are spine-like or pungent, and usually retrorsely sca- 
brous (when antrorsely scabrous, the inner bristles are fused and not grooved); 
2) the axis of the fascicle terminates in a spikelet; 3) inner bristles are fused, at 
least at the base: and 4) have a base chromosome number of x = 17. 

Though, there are still species in Cenchrus and Pennisetum whose generic 
placement still needs clarification. For example, C. myosuroides Kunth, which 
has a base number of x = 9 or 10 and a fascicle structure very different from 
Cenchrus s.s.,as well as some South Pacific taxa. The process of obtaining the 
materials needed to resolve these problems has begun. 


New ComBINATIONS IN THE PENNISETUM CILIARE COMPLEX 


Pennisetum setigerum (Vahl) Wipff, comb. nov. BAsionyM: Cenchrus setigerus Vahl, 
Enum. PL. 2:395. 1805. Pennisetum vahlit Kunth, nom. illeg., Rév. Gram. 1:49. 1829. 
Pennisetum ciliare (L.) Link var. setigerum (Vahl) Leeke, Z. Naturwiss. 79:22. 1907. 
Cenchrus ciliaris L. var. setigerus (Vahl) Maire & Weiler, Fl. Afr. Nord. 1342. 1952. 
Type: Arabia. Forsskdl (HOLOTYPE: C)). 


— 


Fisher et al. (1954) reported that the type of reproduction was identical between 
Pennisetum ciliare and Cenchrus setigerus and that there was variation 
in morphological characteristics between the two species. He concluded that the 
two species are members of a single agamic complex. Snyder et al. 1955) also 
reported that these two species had similar reproductive behavior. Bashaw (1953), 
after studying the morphology, cytology, and mode of reproduction of Cenchrus 
setigerus, concluded that C. setigerus and Pennisetum ciliare were “much more 
closely related than our present classification indicates, perhaps even varieties of 
the same species.” DeLisle (1963), after examining specimens of each taxon from 
throughout their range, only observed a few specimens that could be considered 
intermediates and recognized the two taxa as distinct species of Cenchrus. 
Read and Bashaw (1969) hybridized a sexual genotype of P.ciliare with an 
apomictic genotype of C. setigerus. The resulting F) population consisted of both 
sexual and apomictic plants that represented a complete intergradation in 
morphology between the parents. They also stated that some of the hybrids were 
so different from either parent that populations from them might be mistaken 


WIPFF, NOMENCLATURAL CHANGES IN PENNISETUM 527 


for new species. The hybrids were highly fertile and had fewer quadrivalents 
and more bivalents than either parent. They concluded that the two species 
were certainly congeneric and possibly conspecific. They noted, however, that 
“They have been effectively isolated in nature by obligate apomixis and their 
morphological distinctness was sufficient to permit valid taxonomic treatment 
at the species level. It is also apparent that with sexuality in buffelgrass 
[Pennisetum ciliare], we are able to produce an unlimited number of distinctly 
different hybrids. At present it would be convenient to retain specific rank al- 
though we feel it would be justifiable to merge the species.” (Read & Bashaw 
1969, p. 806). Although they recognized both taxa as species of Cenchrus, Read 
and Bashaw stated that it might become necessary in the future to reconsider 
the generic rank of this entire agamic complex. 


Pennisetum pennisetiforme (Hochs. & Steud. ex Steud.) Wipff, comb. nov. (Figs. 1, 
2). BASIONYM: Cenchrus pennisetiformis Hochs. & Steud. ex Steud., Syn. Pl. Glumac. 
1:109. 1854. TyPE: Saudi Arabia: Jedda, “In deserto pr. oppid. Deschedda.,” 28 Jan 
1836, Schimper 973 [LECTOTYPE, here designated: P!; ISOLECTOTYPES: K! (3 sheets). 
Steudel (1854) cited two collections (Schimper 973 and 974) in the protologue. Sheets 
of these collections were examined from P and found to be similar. Schimper 973 was 
chosen as the lectotype because duplicate sheets are known to exist at K, whereas 
presently there is only one sheet of Schimper 974 known to be in existence. 


Delisle (1963) considered Cenchrus pennisetiformis as part of C. ciliaris. Clayton 
(1982) reported that the boundary between C. ciliaris and C. pennisetiformis 
was indistinct, but that the species could be separated as follows: Cenchrus 
pennisetiformis has inner bristles basally connate for 1-2.5 mm of their length, 
is usually annual (short-lived perennial), smaller in stature and found mostly in 
sub-desert grasslands; whereas, C. ciliaris has the inner bristles basally connate 
for (O-)0.5-L5 mm of their length, isa stout perennial, with or without rhizomes, 
usually forming a hard, knotty, sometimes almost woody base, and is found in 
deciduous bushland and wooded grasslands. 


Pennisetum somalensis (Clayton) Wipff, comb. nov. BASIONYM: Cenchrus somalensis 
Clayton, Kew Bull. 32:3. 1977. TyPE: SOMALI REPUBLIC: Erigavo, 5000-7000 ft 
[1524-2133 m], under shade of bush and trees, Nov 1938, A.S. McKinnon S221 (HO- 
LOTYPE: K!). 

Clayton (1977) reported that C. somalensis and C. pennisetiformis were closely 

related, but that C. somalensis isa densely tufted perennial with inrolled leaves 

about 1 mm wide; whereas, C. pennisetiformis is an annual, or short-lived pe- 
rennial, with flat leaf blades, 2-5 mm wide. 


ACKNOWLEDGMENTS 


lam grateful to the following herbaria for loaning specimens: C, ISC, MO, NY, P, 
US; and to K for providing photographs of type specimens. | would like tothank 
Mary E. Barkworth (UTC) and Kathleen M. Capels (UTC) for assistance while 


528 BRIT.ORG/SIDA 19(3) 


1 


i 


P hull SihtdlutQhnbubliubibn 


= 


emchrus pemmichiferned 

Cenchrus /é ii he 

| yi ciple ht | 
' : Ane on 7 | 


Bieséne Sheet 


Hebimper ¥V Aral fel BA TL Hedemachor. 1549, 
anther fis 


wala, 


Dhat kh af€ +k | (D\ n¥€ D, 5 H if; 
Fic. 1 g pe (P) 


WIPFF, NOMENCLATURAL CHANGES IN PENNISETUM 


Lie ey : 
Contents PE ak 

hn se MA oft 
: : (ec Neet we bt, By 
tn. : 


e 


sat cane bes mason 


W. Schimper. PI. Arab. fel. 


fee 


» FZ. 22 ees Ch ec Pe LEBEL LSLS UY “> £ + J Te, LA 


Ed. Il. Hohenacker. 1848. 
2 CF 4 
Ve a Seco. 


Z 


In deserto pr. oppid. Dschedda. D. /% Jan. 1836. 


Dhat kK fel lab | Lal 1 bat (p)\ £p M = A 
Fig. 2. | g ype (P) 


preparing the manuscript, and for their editorial reviews of the manuscript, 
Alison Kelly (UTC) for her assistance with the digital images of the types, and 
UTCss assistance in obtaining loans of type specimens; and to Stanley Jones 
(BRCH) and two anonymous reviewers for their insightful comments. 


REFERENCES 


Avoutov, N.P. 1931.Karyo-systematische untersuchung der familie Grarnineen. Trudy Prikl. 
Bot. 44, Suppl.:119-123 

BasHaw, E.C. 1953. An investigation of the morphology, cytology, and mode of reproduc- 
tion of Cenchrus setigerus. Ph.D. dissertation. Texas A&M University, College Station. 

BasHaw, E.C. 1969. Registration of buffelgrass germplasm. Crop Sci. 9:396. 

CHASE, M.A. 1920.The North American species of Cenchrus.Contr. U.S. Natl. Herb. 22:45—-77. 


Ciayton, W.D. 1977. New grasses from eastern Africa. Studies in the Gramineae: XLII. Kew 
Bull. 32:1-4. 


530 BRIT.ORG/SIDA 19(3) 


Crayton, W.D. and S.A. Renvoize. 1982. Gramineae, Pt. 3:451—898. In: R.M. Polhill, ed. Flora of 
Tropical East Africa. A.A. Balkema, Rotterdam 

Crayton, W.D. and S.A. Renvoize. 1986. Genera graminum: grasses of the world. Kew Bull. 
Addit. Ser. 13. Her Majesty's Stationery Office, London. 

Deuste, D.G. 1963. Taxonomy and distribution of the genus Cenchrus. lowa State J. of Sci. 
37:259-351. 

FILGuEIRAS, T.S. 1984. O Género Cenchrus L. No Brasil (Gramineae: Panicoideae). Acta 
Amazonica 14:95-127. 

Fisher, W.D.,E.C. BasHaw, and E.C. Hout. 1954. Evidence for apomixis in Pennisetum ciliare and 
Cenchrus setigerus. Agron. J.46:401-404. 

Henrard, J.1. 1935. Identification of some Malaysian grasses. Blumea 1:305-311. 

HIGNIGHT, K.W., E.C. BasHaw, and M.A. Hussey. 1991. Cytological and morphological diversity 
of native apomictic buffelgrass, Pennisetum ciliare (L.) Link. Bot. Gaz. 152:214-218, 

JAUHAR, P.1981.Cytogenetics and breeding of pear! millet and related species. Vol. 1.Progress 
and topics in Cytogenetics In: A.A. Sandberg, ed. Alan R. Liss, Inc., New York, NY. 

Nunez,O.1952.Investigaciones cariosistematicas en las Gramineaes Argentinas de la tribus 
“Paniceae.” Revista Fac. Agron. Univ. Nac. La Plata 28:229-256. 

POHL, R.W. 1980. Gramineae, Family # 15.In:William Burger, ed. Flora Costaricensis. Fieldiana 
Botany, New Series No. 4. Field Museum of Natural History, Chicago, IL. 

Pout, R.W.,and G. Davibse. 1994. Cenchrus L.Pp 374-375. |In:G. Davidse, M. Sousa S.,and A.O. 
Chater, eds. Flora Mesoamericana, Vol. 6, Alismataceae a Cyperaceae. Universidad 
Nacional Autonoma de México, México City, México. 

Reap, J.C. and E.C. BasHaw. 1969. Cytotaxonomic relationships and the role of apomixis in 
speciation in buffelgrass and birdwoodgrass. Crop Sci. 9:805-806. 

ScHmelzer, G.H. 1989. Pennisetum Section Brevivalvula in West Africa: Morphological and 
genetic variation in an agamic species complex. Ph.D. dissertation. Landbouwuniversiteit 
Wageningen. 

SoHNs, E.R. 1955. Cenchrus and Pennisetum: Fascicle morphology. J. Wash. Acad. Sci. 45: 
135-143. 

Snyper, L.A., A.R. HerNanbez,and H.E.Warm«e. 1955.The mechanism of apomixis in Pennisetum 
ciliare. Bot. Gaz. 116:209-221. 

Steupet, E.G. 1854. Synopsis plantarum glumacearum. Part 1.Gramineae.J.B. Metzler; Stuttgart. 

Watson, L.and MJ. Datiwitz.(1992 +).’Grass genera of the world: Descriptions, illustrations, 
identification, and information retrieval; including synonyms, morphology, anatomy, 
physiology, phytochemistry, cytology, classification pathogens, world and local distri- 
bution, and references.’ http://biodiversity.uno.edu/delta/. Version: 18th August 1999. 

Weester, R.D. 1987. The Australian Paniceae (Poaceae). J. Cramer: Berlin. 

Weester, R.D. 1988. Genera of the North American Paniceae (Poaceae: Panicoideae). Syst 
Bot. 13:576-609. 

Wierr, J.K.1995.A biosystematic study of selected facultative apomictic species of Pennise- 
tum (Poaceae: Paniceae) and their hybrids. Ph.D. dissertation, Texas A&M University, 
College Station. 


REEVALUATION OF AYLACOPHORA AND PALEAEPAPPUS 
(ASTERACEAE: ASTEREAE) 


José M. Bonifacino 


Departamento Cientifico de Plantas Vasculares 
acy, ae de Ciencias Naturales y Museo 
Pa Ge eos La Plata, ARGENTINA 


“iboniaemines fenym.unip.edu.ar 


ane cca 


fs d de Agronomia 


Casilla de Correos 1738 Montevideo, l URt UGUAY 


Gisela Sancho 


Department of Botan 
Naweng! Mpeulp eee att SOU 
nian Inctit. 
Waser DC 20560-0166, U.S.A. 
sancho.gisel 


JH.SLEDU 
Permanent addre 
Departamento Aries de ie Vasculares 
ey tad de Ciencias Naturales y Muse 
P. CP 1900 La Plata, pe NA 


ABSTRACT 


=~ 


Nardophyllum deserticola (Cabrera) Nesom and Nardophyllum patagonicum (Cabrera) Nesom are 
two endemic species from Argentinean Patagonia that were originally — under the mono- 
typic genera Aylacophora and Paleaepappus in 1953 and 1969, respectively. In 1993, Nesom included 
Aylacophora and Paleaepappus within the genus Nardophyllum, arguing oe the discontinuities 
between the paleaceous pappus of Aylacophora and Paleaepappus and the pappus bristles of 
Sea do not justify ae them as distinct genera. Careful observation of the her- 
arium material revealed that: 1) the us of N. deserticola and N. patagonicu nsists of ca. 10 
linear- ie paleae, while in dete de hs m ana the pee is composed of ca. 30 
bristles, and 2) the receptacular paleae of Nard 
are wide and enclose the florets, ane in Nardopia ilu species, the paleae, if ee are narrow 
and do not enclose the florets. These obec? Levene DEED N. desert icola and N. 
ce iii with respect to the rest of ideration of both species 
o two distinct monotypic genera, Pera raand a eis respectively. 


RESUMEN 
Nardophyllum deserticola (Cabrera) Nesom y Nardophyllum patagonicum (Cabrera) Nesom son dos 
especies endémicas de la Patagonia argentina que fueron originalmente descritas bajo los generos 
monotipicos Aylacophora y age oe: en ae y oe eee aio Posteriormente en — 3; 


Nesom naluye Aylacophora y P. 
discontinuidades entre el papo palaceo de Aylacophora Paleaepapps y fa papo lee de 


| observacion detallada 


Nardophyllum no justifican consi 


oO 


SIDA 19(3): 531 — 538. 2001 


532 BRIT.ORG/SIDA 19(3) 


del material de herbari lo que: 1) el papo de N. deserticola y N. patagonicum consiste en ca. de 10 


yaleas linear-elipticas, mientras _ en el resto de las especies de Nardophyllum, el papo esta 


— 


compuesto por ca. 30 cerdas y 2) las paleas del receptaculo de eras deserticola y 


Nardophyllum patagonicum son anchas y abrazan las flores, tr | resto de las especies 
de Nardophyllum, las paleas, si estan presentes, son mas nasostae y no abrazan las flores. Las 
discontinuidades observadas entre N. deserticola y N. patagonicum con el resto de Nardophyllum 


apoyan la consideracién de ambas especies como dos géneros monotipicos independientes, 


Oy 
Aylacophora y Paleaepappus, respectivamente. 


Endemic to Argentinean Patagonia, Nardophyllum deserticola (Cabrera) Nesom 
and Nardophyllum patagonicum (Cabrera) Nesom are 2 of the 10 species recog- 
nized inside Nardophyllum Hook. & Arn. by Nesom (1993). 

Nardophyllum deserticola and N. patagonicum were originally described 
by Cabrera under the monotypic genera Aylacophora and Paleaepappus. 
Aylacophora (Cabrera 1953) was characterized by its paleaceous receptacle, scaly 
pappus, and compressed cypselae with 2(-3) ciliate ribs. Paleaepappus (Cabrera 
1969) was defined by its paleaceous receptacle and its pappus of 7-8 paleae. In 
contrast, Nardophyllum sensu Cabrera (1954) has receptacles naked or with only 
3-6(-13) paleae,a pappus of bristles, and terete, more or less pubescent cypselae. 

According to Nesom, the discontinuities between the paleaceous pappus 
of Aylacophora and Paleaepappus and the bristles of Nardophyllum pappus do 
not support their distinction from Nardophyllum because there is a tendency 

for the pappus bristles to be somewhat flattened in Nardophyllum. 

However, according to our observations and the inter pretation of the data 
gathered (see below), Aylacophora and Paleaepapus should be considered as 
independent genera from Nardophyllum as follows: 


— 


Aylacophora Cabrera, Bol. Soc. Argent. Bot. 4:266. 1953. Type Species: Aylacophora 
deserticola Cabrera, Bol. Soc. Argent. Bot. 4:268. 1953. Nardop Pas oe 
(C sees Nesom, cae aa 75. 362. 1993. Fig. 1 A-D. TypE: ARGENTINA. 
PROVINCIA NEUQUEN: Plaza Huincul, 12 Apr 1952, A.L.C. ae 11053 (HOLOTYPE: 
LP ISOTYPE: US). 
Shrub 50 cm high, densely branched; old branches aphyllous, bearing furrows: 
new branches with sparse nodes; leaves linear; capitula discoid, solitary at ends 
of branches; involucres globose; receptacles paleaceous, paleae wide, apically 
pubescent, each palea enclosing a floret; cypselae compressed, 2(-3) nerved, 
pubescent only on the ribs; pappus of 10-11 oblong scales no longer than 1.2 
mim, in | series. 

Ecology.—Aylacophora deserticola inhabits semidesert areas in Patagonia. 
The very few herbarium specimens of this species lead us to regard them as 
very narrowly distributed. According to Ing. Steibel (pers. comm.), A. deserticola 
Cabrera grows on edaphic communities in the Monte biogeographic province 
(Cabrera & Willink 1973), where it is present on sand dunes with very sparse 
shrub cover, associated with Larrea divaricata, Atriplex lampa, Prosopis flexuosa 


meas 
a ye 


4mm 


) | j pt I C. Cypsela.D. Pappus scales. E-H. Paleaepappus 
patagonicus. E. Capitulum. F. Receptacular palea. G. Cypsela with pappus. H. Pappus palae. I-L. Nardophyllum 
bracteolatum.1.Capitulum.J.R | lea. K. Cypsela with Pr Pappus bristle M-O0. Nardophyllum bryoides. 


Fic. 1.A- A.Capitulum.B.R 


M. Receptacular palea.N Pappus bristle 0 Cypsela with pappus. (A—D, based on Cabrera 11053, LP; E-H, based on Rio 
Chico, 1900, Ameghino s.n., LP; I-L, based on Serra 77, LP; M—O, based on Ruiz Leal 27011, LP) 


534 BRIT.ORG/SIDA 19(3) 


var. depressa, Senna aphylla subsp. divaricata, Chuquiraga erinacea, Gutierrezia 
solbrigii, Fabiana patagonica and Larrea cuneifolia. Aylacophora deserticola has 
been collected in Argentina, Prov. Neuquen, Dptos. Confluencia, Cutralcé, Anielo, 
Pehuenches, and Zapala. West of Rio Covunco and south of Rio Neuquén in 
Dpto. Zapala, A. deserticola is the dominant species regarding surface cover. 
Flowering in the fall. 


Paleaepappus Cabrera, Bol. Soc. Argent. Bot. 11:273.1969. Type SPECIEs: Paleaepappt 
patagonicus Cabrera, Bol. Soc. Argent. Bot. 1:273.1969. Nerdopyila patagonicum 
(Cabrera) Nesom, Phytologia 75:362. 1993. Fig. 1 E-H. Type: ARGENTINA. Chubut, 
Rio Chico, “Aut. 1900,” Ameghino s.n (HOLOTYPE: LP!) 


Shrub densely branched, lateral branches sharp ended; leaves oblong to spatu- 
late, coriaceous; capitula discoid, solitary at the ends of branches; involucres 
campanulate; receptacles paleaceous, paleae wide, apically pubescent, each 
enclosing a floret; cypselae terete, densely pubescent; pappi of 9-10 elliptic 
paleae 7 mm long, in 2 series. 

Ecology.—Paleaepappus patagonicus inhabits semidesert areas in 
Patagonia. Knowledge of the ecology of Paleaepappus is scarce and speculative 
because the only record of this species is the type itself, and there is no ecologi- 
cal information on the label. The conservation status of this species could aptly 
be recorded as endangered. 

Aylacophora and Paleaepappusare strongly segregated from Nardophyllum 
by characteristics of the elements of pappus such as number, shape, and num- 
ber of series. The paleaceous pappus of Aylacophora and Paleaepappus con- 
trast with the bristles in Nardophyllum. Intermediate states of pappus shape, 
like narrow paleae, have not been found in Nardophyllum, but occasionally, flat 
bristles have been found in Nardophyllum. 


Nardophyllum circumscription 

Nardophyllum Hook. & Arn. (1836) was described by Cabrera (1954) as shrubs 
densely branched, leaves alternate, small; heads solitary at the end of the 
branches; discoid capitula, involucre campanulate; receptacle small, convex, 
naked or with few paleae; cypselas turbinate, 4-5 ribbed, hairy; pappus com- 
posed of several bristles. The circumscription of Nardophyllum adopted here is 
the presented by Cabrera (1954) in his revision of the genus, where he included 
7 species: Nardophyllum armatum (Wedd.) Reiche, N. bracteolatum Hauman, 
N. genistioides (Phil.) Gray, N. bryoides (Lam.) Cabrera, N. chiliotrichioides 
(Remy) A. Gray, N. lanatum (Meyen) Cabrera, and N. obtusifolium Hook. & Arn. 


Paleaepappus and Nardophyllum 

Contrasting with the pappus of Paleaepappus (Fig. 1 H), the pappus of 
Nardophyllum is composed of ca. 30 bristles 5-10 mm long (Fig. | L), sometimes 
flattened, especially at the apex (Fig. | N), arranged in 2-3(-5) series. 


BONIFACINO AND SANCHO, REEVALUATION OF AYLACOPHORA AND PALEAEPAPPUS 535 


Involucres and shape of Paleaepappus cypselae are similar to those of 
Nardophyllum. Differences in number and shape of receptacular paleae are also 
found in these two genera. Paleaepappus has paleaceous receptacles with wide 
and apically pubescent paleae enclosing each floret (Fig. 1 F). The receptacular 
paleae of Nardophyllum are narrow, a not enclose the florets (Fig. 1 J and M), 
and vary from absent (N. armatum) to 3-6 (rarely more numerous, 9-13 in N. 
bracteolatum). 


Aylacophora and Nardophyllum 

The pappus of Aylacophora (Fig. 1C and 1D) contrast highly with the pappus of 
Nardophyllum (see above). In addition, other c distinguish Aylacophora 
from Nardophyllum: the involucre of Aylacophora is globose (Fig. 1 A); its cypselae 
are re d, 2(-3), and pubescent only on the ribs (Fig. 1 C). The involucre of 
Nardophyllum is campanulate to obconical (Fig. 1 1) and the cypselas are terete, 
(4-)5-7(-8) nerved and uniformly pubescent (Fig. | K and 1 0). 

Aylacophora has a paleaceous receptacle with wide and apically pubes- 
cent paleae enclosing each floret. The number and shape of receptacular paleae 
of Aylacophora (Fig. 1 B) contribute to set this taxon apart from Nardophyllum. 

Aylacophora and Paleaepapus were placed in the Chiliotrichum group 
(Zhang & Bremer 1993; Bremer 1994) that includes shrubs with mostly densely 
set, coriaceous, and often abaxialy tomentose leaves, and for most of the genera, 
paleate receptacles (Bremer 1994). Within this group there are intermediate 
morphotypes for pappus elements that range from terete bristles in 
Chilliophyllum, through narrow paleae in Lepidophyllum (Cabrera 1971), to 
paleae in Aylacophora and Paleaepappus. In addition, the number of receptacular 
paleae vary within the Chiliotrichum group. In reference to this character the 
variation observed within Nardophyllum is also present when comparing other 
genera of Chiliotrichum group, such as Lepidophyllum without paleae, 
Chiliotrichum with few paleae, and Chiliophyllum and Chiliotrichiopsis with 
fully paleate receptacles (Cabrera 1971, 1978). Because the gradation in pappus 
and receptacular paleae is inherent to the Chiliotrichum group itself, this varia- 
tion cannot be an argument against the recognition of both Aylacophora and 
Paleaepappus as distinct from Nardophyllum. 

Genera of Chiliotrichum Group (sensu Bremer 1994) can be identified as 
follows: 


1. Capitulum without ray florets. 


2. Pappus of ca. 30 or more bristles arranged in 2-3(-—5). Nardophyllum 
2. Pappus of 10 or fewer paleae arranged in 1-2 series 
3. Cypselas compressed, with hairs restricted to the ribs. Aylacophora 
3. Cypselas terete, wholly pubescent. Paleaepappus 


1. Capitulum with ray florets. 
4. Receptacles epaleate. 
5. Ray corollas white or violet. Diplostephium 


536 BRIT.ORG/SIDA 19(3) 


5. Ray corollas yellow. 
6. Leaves opposite; pappus composed of a series of unequal paleaceous 


bristles Lepidophyllum 
6. Leaves aerate pappus composed of 2 series of bristles. Parastrephia 
4, Receptacles palea 
7. Ray corollas ae Chiliotrichum 
7. Ray corollas yellow 
8. Pappus of terete bristles. Chiliophyllum 
8. Pappus of linear, acute scales. Chiliotrichiopsis 


Nesom’s reinstatement of Nardophyllum scoparium Philippi (Nesom 1993) is 
not accepted here. Presence of pistillate ligulate florets in the periphery (5-6) 
casts serious doubts about positioning this taxon inside Nardophyllum. The taxo- 
nomic placement of this Chilean species and its possible status as an 
undescribed genus allied to some genus inside Chiliotrichum group as Nesom 
suggests (pers. comm.) is being reviewed by one of us JMB) and will be pre- 
sented as a more comprehensive study of the Chiliotrichum group (in prep.). 

According to Bremer (1994), Aylacophora and Paleae pappusare very similar 
and possibly sister groups. Aylacophora was related to Nardophyllumand Chiliotri- 
chiopsis by Cabrera (1953); Paleaepappus was related to Nardophyllumand Aylaco- 
pho ra (Cabrera 1969). Our opinion, based on observations of cy pselat mot phology, 
number of series of the pappus elements, involucre shape, leaves shape and plant 
habit, favors a closer relationship between Paleaepappus and Nardophyllum. 

Nesom (1993) suggest that the narrow, internally tomentose cauline sulcae 
of Aylacophora could be homologous with those found in some species of Nardo- 
phyllumand so denotinga closer relationship of Aylacophora to Nardophyllum. 
To complicate the matters further, the paleate receptacle of both Aylacophora 
and Paleapapus, would favor a closer relationship of these two genera with 
Chiliotrichiopsis, Chiliophyllumand Chiliotrichum instead, which are,as Nesom 
(1993) points out the closest relatives to Nardophyllum and all of them have a 
paleate receptacle (Chiliotrichiopsis, Chiliophyllum fully paleated, Chiliotrichum 
15-21 paleae). Evidently, as Bremer (1994) states, the elucidation of the phylo- 
genetic relationships among these genera demands a more comprehensive study 
of the Chiliotrichum group that is beyond the scope of this paper. 

Segregation of Aylacophora and Paleaepappus from Nardophyllum on the 
basis of pappus shape is supported by other characters such as the shape of 
both cypsela and capitulum in Aylacophora, and the shape and quantity of 
receptacular paleae in both Aylacophora and Paleaepappus. 

Although Aylacophora and Paleaepappus are closely related to the rest of 
Nardophyllum species, the discontinuities basically observed in the shape of 

the pappus elements, with no defined intermediate states, justify the consider- 


— 
ju 


ation af both species as two distinct genera. 
Our conclusion is supported by Nesom etal. (in press). These authors, based 
on features of the involucre and mainly on pappus morphology, have included 


BONIFACINO AND SANCHO, REEVALUATION OF AYLACOPHORA AND PALEAEPAPPUS 537 


a new Peruvian species of Asteraceae:Astereae inside Chiliotrichiopsis Cabrera 
(Chiliotrichiopsis peruviana Nesom, Robinson & Granda). Nesom et al. (in 
press) found that the pappus morphology is a good character to separate gen- 
era inside Cihiliotrichum group and concluded that Cabrera’s narrow generic 
concept of Nardophyllum was a better description of the diversity inside 
Chiliotrichum group, therefore having independently arrived at the same con- 
clusion we have, regarding the consideration of Aylacophora and Paleaepappus 
as distinct from Nardophyllu 


APPENDIX | 
Additional speci ined of A ticola Cabrera. ARGENTINA. Neuquén, Plaza 
Huincul, 11 Dec 1996, H. ity Se 12503 (SRFA); Plaza Huincul, 11 Dec 1996, H. Troiani et al. 12504 
(SRFA); Paso de Los Indios a ralcd, 5 Feb 1999, P Steibel y H. Troiani 14074 (SRFA); Paso de las Bardas, 


2 Feb 1999, H. Trojani ee ve 13920 (SRFA). Paso de Los Indios, 5 Feb 1999, P Steibel y H. Troiani 
14069 (SRFA); Paso de Los Indios, 20 km hacia Cutralcd, 5 Feb 1999, P Steibel y H. Troiani 14073 (SRFA). 

yllum. Nardophyllum armatum (Wedd.) Reiche. ARGENTINA. San Juan: Iglesia, 
camino a el Fierro, Canada de La Zorra, 24 Jan 1974, Cabrera & al. 24510 (LP); Pampa de Pauacan, 
entre las Aguaditas y Chepical, 12 Dec 1957, Ruiz Leal 18945 (LP).La Rioja: Gral. Sarmiento, Rio del 
Oro, Cordillera, 7 Feb 1947, Hunziker 2197 (LP). Catamarca: Santa Marfa, Campo Arenal, Loerner 8 (LP). 
Salta: San Antonio de los Cobres, 29 Jan 1944, Cabrera ae Jujuy: 1 km al W de la Quiaca, 11 
Feb 1 1960, Meyer & al.21 29] (LP); Rinconada, Cusi Cusi, Mar 197 ris & uloaga 70 (LP): Humah }UdaCa, 
Esquinas Blancas, 22 Ene 1966, Cabrera & al. 17726 (LP); Abr pees Feb 1 1937, Castellanos 20229 


rs 

Nardophyllum bracteolatum Hauman. ARGENTINA. Mendoza: San Carlos, El Pedernal, 25 
Mar 1916, Sanzin 1810 (LP); Tunuyan, Paso del Portillo, Cuesta de los Afligidos, 29 Jan 1934, Ruiz Leal 
2052 (LP); San Carlos, Rincén de los Leones, 18 Jan 1941, Ruiz Leal 7212 (LP); San Carlos, Arroyo de la 
Oda."Casa de Piedra,” 17 Jan 1952, Serra 77 (LP). 

tinea al eee ) Cabrera. CHILE. Magallanes: ja Nacional Torres del Paine, 
Lago Paine, 17 Jan 1999, Bonifacino s.n a 2ee Bee eee NA. Santa Cruz: alrededores de E| 
Chalten, 10 Feb 2000 iene \\V/FA): Guer-Aike, Est ie Cabo Virgenes, 

v 1974, Molina 7 (LP);Rfo alae 5 ae 1932, Coaeleness n. (LP); Est.de las Vizcachas, Cerro de 
ae 17 Jan 1970, Ruiz Leal 27019 (LP); a 2 km de Guarumba, 16 Jan 1970, Ruiz Leal 27011 
(LP); Lago Argentino, Parque Nacional Los Glaciares, Fitz Roy, 14 Feb 1975, Cabrera & al. 25864 (LP).Tierra 
del Fuego. Est. Cullen, 52° 44' S, 68° 33' W, 5 Jan 1972, Moore & Goodall 336 (LP); Bahia Lee. 52° 52'S 
70° 16' W, 6 Nov 1971, Moore 2339 (LP). 

Nardophyllum chiliotrichoides (Remy) A.Gray. ARGENTINA. San Juan: Calingasta, Qda.Los 
avestruces (oeste de Cerro Castano), Feb 1960, Fabris & Marchionni 2354 (LP); Rio Maurique a 
Portezuelo de Potrerillos, 23 Jan 1991, Kiesling hee (Sl). Chubut: a 20 km al E de Cushamen, 31 Dec 

1947, Soriano 2786 (LP); Rawson, 12 km al S de Trelew, 7 Dec 1980, Castroviejo & Lopez 7313 (Sl). 
sas meena aun del Rio San cae Fierro Se Jan ae ening 621 (LP). 

m lanatum (Meyen) Cabrera. ARGENT je, ruta 40,20 km 
al S - eraiae 1 Feb 1963, Boelcke et al. 10420 (SI). iene Laguna Copi Leuquen, al S de 
Calmuco, 15 Feb 1942, Burkart & al. 14425 (LP); Chos Malal, entre Rio Barranca y Buta Ranquil, 8 Feb 
1950, Boelcke 4235 (LP). CHILE. Colchagua: Termas del Flaco, 19 Feb 1966, 76/Iner 833 (LP);Vegas del 
Flaco, al E de la Quebrada de los Rios, falda SE del cerro, 7 Feb 1974, Mahu 9858 (LP). eee 
Cerro ia Cordillera de la Costa, 6 Mar 1966, Zéllner 1402 (LP). Ovalle: Geisse s.n (LP 60 ae 

ardophyllum obtusifolium Hook. & Arn. CHILE. Magallanes: San Gregorio, 26 ae 
ee 33903 (LP). ARGENTINA. Santa Cruz: Pto. San Julian, 1915, Carete s/n (LP); Corpen ne ee 


538 BRIT.ORG/SIDA 19(3) 


km N Piedra Buena, Ruta 3,3 Dec 1971, Boelcke & al. 15339 (LP); Pampa del Castillo, 11 Feb 1936, Scott 
de Brirabén & Birabén 17 (LP); Lago Bs. As., Los Antigu mino a Pto. Moreno, 24 Nov 1965, Correa & 
Nicora 3638 (LP); Puerto Deseado, Jan 1896, Alboff 21 169 (LP); Esquel, 2 Apr 1946, Sco/nik 306 (LP); 
Leleque, 13 Jan 1947, Soriano 2341 (LP); Est. Pepita, Alto Rio Senger, 13 Feb 1947, Soriano 2585 (LP). 
Neuquén: Charahuilla, Arroyo Lapa, Feb 13 1939, Chicchi 123 (LP);Laguna Blanca, 9 Jan 1966;camino 
a Nirihuau, 16 Jan 1935, Cabrera & Job 353 (LP). Santa Cruz: Col. Carlos Pellegrini, Est. La Flora, Dec 
1979, Von Thuingen 2 (LP). 


~ 


ACKNOWLEDGMENTS 


It is a pleasure to acknowledge Liliana Katinas and Jorge V. Crisci for helpful 
critical comments on the final manuscript; Guy Nesom for his helpful critical 
comments on this paper, making available to us the manuscript “A new species 
of Chiliotrichiopsis (Asteraceae: Astereae) from Peru. Guy Nesom, Harold Rob- 
inson, and Arturo Granda Paucar” (Brittonia, in press) and his encouragement 
towards publication; John Strother for his corrections on the final manuscript; 
Ing. Agr. Pablo E. Steibel for the information regarding the ecology of Aylacophora 
and its geographic range; the Consejo Nacional de Investigaciones Cientificas y 
Técnicas de Argentina (CONICET) and Comision Sectorial de Investigacion 
Cientifica (CSIC), Uruguay, for the financial support on this investigation, and 
LP, LPAG, MVE SI, SRFA and US for loans of herbarium specimens. 


REFERENCES 


Bremer, K. 1994. Asteraceae: Cladistics and classification. Timber Press, Portland, Oregon. 

Caprera, A.L. 1953. Un nuevo género de astereas de la Republica Argentina. Bol. Soc. Ar- 
gent. Bot. 4:266-271. 

Casrera, A.L. 1954. Las especies del género Nardophyllum. Notas Mus. La Plata, Bot. 83, 
17:55-66. 

Caprera, A.L. 1969. Compuestas nuevas de Patagonia. Bol. Soc. Argent. Bot. 11:273-275. 

Caprera, A.L. 1971. Compositae. En M. N. Correa (ed.), Fl. Patagdnica, Colecc. Ci. Inst. Nac. 
Tecnol. Agropecu. 8(7):1—451. 

Caprera, A.L.1978.Compositae. In: A.L. Cabrera, ed. Fl. Prov. Jujuy, Colecc. Ci. Inst. Nac. Tecnol. 
Agropecu. 13(10):1-726. 

Caprerd, A.L.and A. Witink. 1973. Biogeografia de America Latina. O.E.A. Serie de Biologia, 
Monografia 13.Washington, D.C 

Grau, J. 1977. Astereae. Systematic Review. |n:V.H. Heywood, J.B. Harborne, and B.L. Turner, eds. 
The biology and chemistry of the Compositae 2. Academic Press. London. Pp. 539-576. 

Nesom, G.L.1993.Taxonomic status of Nardophyllum scoparium (Asteraceae: Astereae) with 
observations on the definition of Nardophyllum. Phytologia 75:358-365. 

Nesom, G.L., H. Roginson and A. Granpa. A new species of Chiliotrichiopsis (Asteraceae: 
Astereae) from Peru. Brittonia (in press). 

ZHANG, X. and K. Bremer. 1993. A cladistic analysis of the tribe Astereae (Asteraceae) with 
notes on their evolution and subtribal classification. Pl. Syst. Evol. 184:259-283. 


GEOGRAPHIC VARIATION AND TAXONOMY OF NORTH 
AMERICAN SPECIES OF MIRABILIS, SECTION 
OX YBAPHOIDES (NYCTAGINACEAE) 


Richard Spellenberg and Sergio R. Rodriquez Tijerina' 


Department of Biology 
New Mexico State University 
Las Cruces, NM 88003, U.S.A 


ABSTRACT 


A revision of Mirabilis, section Oxybaphoides, Nyctaginaceae, in western North America is presented. 
Mirabilis oligantha (Standl.) J.K Macbride, M. oxybaphoides (A. Gray) A. Gray, and M. tenuiloba S. 
Wats. remain as traditionally classified. Mirabilis bigelovii A. Gray, M. californica A. Gray ex Torr, M. 
| ee ) Curran, and M. retrorsa Heller are combined as a single species, M. laevis, and recog- 

M. laevis var. sis (Kellogg) Spellenb. (comb. nov.), M. laevis var. crassifolia 
aa ene me nov.), M. laevis var. laevis, and M. laevis var. retrorsa (Heller) Jepson, re- 
spectively. Distribution maps are presented ‘or ee species, those for the varieties within the M. 

of 


laevis complex abe madlcating & g characters. The inspection of these maps 
was of imy g ic decisi Lists of important collections are provided. Chro- 
1 if£ fi 


the first time for M. laevis var. villosa (2n = 30,,), M. laevis var. retrorsa 
(2n = 31-33,) and M. oxybaphoides (2n = 30,). 


RESUMEN 


Se presenta una revision ae Me igs seccion earn By clagmaceas en el oeste de Norte 


América. Mi J.F Macbride, I A Gray) A. Gray. y M.tenuiloba 


S. Wats. permanecen ‘tal como se clasificaban ma aoa ees Mirabilis bigelovii A. Gray, M. 
californica A. Gray ex Torr, M. sit (Benth.) Curran, y M. retrorsa Heller se combinan como una 
nica especie, M. laevis, y variedades, i.e., M. laevis var. villosa (Kellogg) Spellenb. 


(comb. nov.), M. laevis var. crassifolta snolsy spellen>: (comb. eae M. tdevts var. hata y M. laevis 


im gee | 


y de las variedades. en nel comes Me laevis indieands rambién la distribucion geografica ae los 


caractere importante para tomar las decisiones taxonomicas. Se 
ofrecen re de colecciones ious Se citan por primera vez numeros cromosomaticos de M. 
Qn 


laevis var. villosa (2n = 30,), M. laevis var. retrorsa Qn = 31-33,) y M is 


INTRODUCTION 


Mirabilis L. is primarily a New World genus comprising 45-60 species distrib- 
uted from southern Canada to southern South America, with one native to 
southern Asia (Bogle 1974; Heimerl 1934; Le Duc 1995). Species have been sus- 
pected of hybridization (Shinners 1951). In addition, some are known to be au- 
togamous and even cleistogamous (Cruden 1973). Stamens and style curl tightly 
together in flowers of the species treated here, as in other arid-land Mirabilis in 


‘Present address: Facultad de Agronomia, Universida Autonoma de Nuevo Leon, Apdo. Postal 358, C.P. 66450, 
San Nicolas de los Garza, Nuevo Leon, México. 


SIDA 19(3): 539 — 570. 2001 


540 BRIT.ORG/SIDA 19(3) 


southwestern North America, probably effecting self-pollination as observed 
in Boerhavia (Chaturvedi 1989; Spellenberg 2001), species in other Mirabilis 
sections (Cruden 1973; Hernandez 1990), and several other genera (Spellenberg 
S Delson 1977). Coupling hybridization with autogamy may produce individu- 
ally rather uniform populations, but geographically complex variation patterns 
(Stebbins 1957). 

Such complexes provided fertile ground for the description of numerous 
entities under taxonomic traditions of early in the 20" century, in which, be- 
cause of locally uniform populations but widespread variation across a geo- 
graphic region, taxonomic decisions may be subjective and perhaps utilitarian, 
following a philosophy expressed by Lewis (1963). Here, for example, more than 
40 synonyms apply to our concept of Mirabilis laevis (Benth.) Curran and va- 
rieties. The taxonomic problems associated with Mirabilis were commented 
upon by Shinners (1951, p. 173) (“Mirabilis is surely one of the most trouble- 
some of Southwestern Jae in nomenclature and taxonomy both.”) and by 
Standley (193la, p. 73) after several decades of study in the family (“I know of 
few groups of plants [Neea, Torrubia, Mirabilis] in which specific differences 
are so unstable and so bafflingl;] .. no single character seems to be constant.”) 
Turner (1993), conversely, in a rather refreshing approach to the taxonomy of 
the genus, noted that if emphasis on vegetative variation were minimized, and 
fruit characteristics were emphasized instead, the genus in Texas was taxonomi- 
cally tractable. 

Mirabilis was divided into six sections by Heimerl (1934; translated in part 
and reviewed in Le Duc 1995), one of which, Oxybaphoides A. Gray, was char- 
acterized by slightly accrescent involucres and fruits that are comparatively 
small and unornamented (Fig. 1). Heimerl included in it the North American 
species M. oxybaphoides(A. Gray) A. Gray, M. californica A. Gray (and close al- 
lies), a number of South American species, and one southern Asian species. 

Mirabilis oxybaphoides has presented little taxonomic controversy at the 
species level since its description by Gray (1853) in the genus Quamoclidion. lt is 
sufficiently distinct from other species of Mirabilis (as the genus is now generally 
construed) that it formed the monotypic genus Allioniella of Rydberg (1902). 
This classification was followed by Standley (1909, 1918) in several treatments 
of the family, but he was apparently unaware of its presence in Mexico, as it 
was not included in his treatment of the family for that nation (Standley 1911). 

The remaining taxa of the section in North America were placed in a new 
genus Hesperonia by Standley (1909), who emphasized differences of fruit form, 
shape of the perianth, and number of flowers in the involucre. Standley recog- 
nized eight species and several subspecies, emphasizing shape, color, size of the 
fruit,and vegetative characters such as plant size, leave size and shape, and char- 
acteristics of vestiture. Jepson (1914) treated this as subgenus Hesperonia 
(Standl.) Jepson, including M. californica and M. tenuiloba S. Wats., noting also 


NYCTAGINACEAE 541 


@2O@0F 008 @ 
A B D & FG I J 


laevis -----—- crassifolia ----- ------- villosa ------- -——----- retrorsa ------- 


laevis 


n®@ 


e @©0@080 
T V OW 


U 


@ee., 
N OP Q oR 


------- oligantha ------- ---- oxybaphoides --- ----- tenuiloba —- 


Fic. 1. Variati ay eee fN . rad n Celt. 44 (Mienhilt 


baphoides. in North America 


laevis, M. oligantha, M oxybaphoides, M seniilbbant Tt he line“leavis” refer ieti ithin M. /aevis. Fruit 
Aji 


Nis 7.5 mm longa. Each | fruit f iff Collections, fully cit 
2 identified t isk (*) following herbari fd ition, are: A, Brandegee s.n.; B, Palmer 886: G Blakley 5657; 


D. Spellenbe 91233 *E. Spellenberg 12335 F 4°G 10206; |, Spellenberg 
12332;J na 303; K Spellenber 12342; ‘i. -Spllenber 1 

Gentry & Fox 11731 John Auller 603; R, Waterfall 12142;5, Columbus 637; i; Gorell iohiiston 
24516; U, Powell & B. L Turner 1708: V, Wiggins & Wiggins 15863; W, Brandegee s.n. 


139290-8 c 470972.N WL * Ws, ° 4LOAN 


that M. laevisand M. cedrosensis (Standl.) Jepson were closely related if not the 
same as M. californica. Standley (1931b), upon completing studies of South 
American Nyctaginaceae, noted that characteristics used to distinguish North 
American genera allied to Mirabilis did not do so and also chose to unite all in 
an inclusive Mirabilis, a classification followed by most botanists since then. 
Of those taxa early placed in Hesperoniaas they are recognized here, M. oligantha 
(Stand1.) J.E Macbride remains poorly known, M. tenuiloba has presented very 
few problems, but M. laevis has been a source of a plethora of names as taxono- 
mists have attempted to deal with the variation presented by populations in 
the complex. The high points of these taxonomic meanderings are discussed 
under each of the taxa below. 

Recently Le Duc (1993) described Mirabilis russellii Le Duc from the west coast 
of Mexico, placing the new species in the section Ox ybaphoides because of its suffru- 
tescent nature, campanulate perianth, and ilaginous anthocarp (when wet). 
It rests very poorly in this section primarily because of general habit and antho- 
carp morphology. The single immature anthocarp available to us on the paratype 
at NMC generally resembles anthocarps of several other Mexican Mirabilissuch 
as M. sanguinea Heimerl, M. hintoniorumLe Duc, and M. urbanii Heimerl,as figured 


542 BRIT.ORG/SIDA 19(3) 


in Le Duc’s (1995) plate Il. For the present treatment of section Oxybaphoides 
we exclude the sp d suggest it lies more comfortably in section Mirabilis. 

In this paper we examine “the geographic variation and taxonomy of the 
most complex species in the section in North America, Mirabilis laevis,and we 
provide a key, descriptions, and distribution maps for the other three species in 
the section Oxybaphoides in the United States and Mexico. Mirabilis laevis and 
its component taxa have a tortuous taxonomic history that has resulted in many 
names published at the specific and infraspecific levels (Rodriguez 1992), based 
on differing generic, specific, and infraspecific concepts in the group. Generic 
concepts emphasized primarily the importance of the number of flowers per 
involucre, the shape of the fruit, and the degree of accrescence of the involucre. 
Specific or varietal decisions have primarily emphasized fruit shape and sur- 
face patterns, color of perianth, and nature of pubescence of foliage and stems. 


MATERIALS AND METHODS 


For this study more than 3000 herbarium specimens were examined from A, 
ARIZ, ASU, BRY, CAS, DS, GH, K, MO, NMC, NY, POM, RM, RSA, SD, UC, US, and 
UTC (abbreviations from Holmgren et al. 1990). From these specimens, 256 from 
the M. laevis complex were selected that had information about perianth color, 
possessed ripe fruits and at least midstem leaves, and had adequate data re- 
garding place and date of collection. These specimens represent the morpho- 
logical variation and geographic range of the taxa. They supplied data for mor- 
phological characteristics plotted in Figures 2 and 3 and described in treatments 
of taxa. Taxonomic decisions were made after study of specimens and the in- 
spection of maps generated by plotting morphological characteristics geo- 
graphically. Types or microfiches of types for basionyms were seen insofar as 
possible. From this information taxa were delineated that seemed to have some 
morphological, ecological and geographical reality. Those that showed consid- 
erable intergradation were recognized at the varietal level. The order of taxa in 
the treatment is based on perceived habitat specialization and reduction in num- 
ber of fruits as generally compared to other Mirabilis. 

We h ted deta f variation based on study of speci- 
mens in the Mi rabilis laevis complex from the main taxonomic treatment and 
have included that in Appendix |. Appendix 2 consists of standard citations of 
representative and/or cited specimens., including those that voucher chromo- 
some counts. 

Le Duc (1995) provides a key to the sections of Mirabilis 


TAXONOMY 
Mirabilis sect. Oxybaphoides A. Gray in Torrey, Bot. Mex. Bound. 173. 1895. 
Allioniella Rydberg, Bull. Torrey Bot. Club 29:687. 1902; Hesperonia Standley, Contr 
U.S. Natl. Herb. 12:360. 1909. Type: Mirabilis oxybaphoides (A. Gray) A. Gray. 


rf NYCTAGINACEAE 543 


Herbaceous to suffrutescent or shrubby perennials; root (of North American 
taxa; others unknown) long, cylindrical, cordlike; stems erect to decumbent or 
prostrate, densely to sparsely leafy. Leaves more or less evenly distributed, basal 
leaves larger, petiolate, distal leaves smaller, short-petiolate or sessile, margins 
plane. Inflorescences axillary and terminal in open or congested, few- or repeat- 
edly-branched cymes; involucres bell-shaped, slightly accrescent, with | or 3 
flowers inserted at base. Perianth broadly funnelform, abruptly flared from 
narrow tube, deeply 5-lobed; stamens 3-5. Fruit ellipsoid or obovoid, base not 
or slightly constricted, apex rounded, truncate, or somewhat nipple-like, sur- 
face with 0 or 5-10 indefinite or prominent lines, often somewhat furrowed, 
smooth or very slightly rugose, usually glabrous, mucilaginous when wetted. 

A poorly understood section of about 10-20 species, North America, South 
America, southern Asia. Heimer! (1934) suggested there were about 23 species 
in the section, but considerable redefinition and consolidation of taxa in North 
America has reduced that number. Diversity is greatest in South America 
(Heimer! 1934). 


KEY TO NORTH AMERICAN MIRABILIS, SECTION OXYBAPHOIDES 


1. Involucres 3-flowered. 1.M. oxybaphoides 
. Involucres 1-flowered (very rarely 2 flowers). 
2. Fruits 6-8 mm long; perianth white, 15-18 mm long above the constriction. 
4.M. oligantha 
2. Fruits 2.5-5.5 mm long; perianth white, pink, or magenta, 7-15 mm long above 
the constriction. 
3. Involucre in flower 7-10 mm long, the lobes narrowly lanceolate, at the base 
1/4-1/3 as wide as long; perianth white or pale pink; leaf blades commonly 
4-6 cm long, ascending. 3.M.tenuiloba 
3. Involucre in flower 3-6 mm long, the lobes triangular to lanceolate, at the 
base 1/3 to equally as wide as long; perianth white, pink, or magenta; leaf 
blades commonly 1-3.5 cm long, spreading or ascending. .M. laevis 


ay 


1. Mirabilis oxybaphoides (A. Gray) A. Gray in Torr. U.S.& Mex. Bound. Bot. 173. 
.Quamoclidion oxybaphoides A. Gray, Amer. Woes Seie2 520), 1853. Allionia 
oxybaphoides (A. Gray) Kuntze, Rev. Gen. PI. 22:533. 1891. Allionella 
(A. Gray) Rydb., Bull. Torrey Bot. Club 29:687. 1918. Type: at the foot of mountains 
east of El Paso, in the shade of high rocks, Wright 596 (Lectotype, here designated: 
GHI, right side of sheet). Gray mixed Wright's field numbers 1223 (mountains near 
El Paso, in shade, procumbent, Sep 12, 1849) and 1322 (at the foot and in the shade 
of high rocks, fl. purple, Oct 14, 1849, noted on sheet to be probably Hueco Tanks). 
At this time it cannot be determined from which site either of the two specimens 
on the sheet originated. The one on the right is the more mature and representa- 
tive; a second specimen originally at the Boston Society of Natural History, trans- 
ferred to GH! in 1941, very closely resembles the plant on the right of the type sheet, 
is in a similar stage of maturity, and is a probable isolectotype. 


Oxybaphus wrightii HemslL., Biol. Centr. Amer. 3:3. 1882. TypE: NORTH MEXICO: 
Chiricahui Mountains, Wright (HOLOTYPE: K). Hemsley sites only general locality 


544 BRIT.ORG/SIDA 19(3) 


and collector, without number or date. Gray (1853) cites Wright 1721 (GH), from 
Guadalupe Pass in the “Chiricahui Mountains,” the collection probably seen by 
Hemsley. lf from the present cay ne Pass, the collection originated in the 
Peloncillo Mountains in New M 
Mirabilis Hehe Alea val. glabrata Heimer! Annuaire Conserv. Jard. Bot. Genéve 
180. Alli var. glabrata (Heimerl) Standl.,, Contr. U.S. Natl. 
Herb. . ao 1909. TYPE: NEW MEXICO. LINCOLN Co.: El Capitan Mts., 31 Aug 1900, 
FS. & E.S. Earle 399 (HOLOTYPE: US! ISOTYPE: NMC). 


es 


Plants usually loosely clump-forming, herbaceous basally, the stems often 
intertangled and clambering through other vegetation. Stems ascending, spread- 
ing or decumbent, 0.2-1.2 m long, repeatedly branched, green throughout, pu- 
berulent in lines or throughout, glandular or not, the pubescence denser dis- 
tally. Leaves thin or slightly fleshy; petioles up to 3.5 cm long on basal leaves, 
becoming progressively shorter distally, the distal leaves subsessile or on peti- 
oles to 4 mm long; blades of the basal and midstem leaves broadly deltoid or 
ovate, 1.5-8.0 cm long, 1.0-7.5 cm wide, glabrous or puberulent, and then often 
glandular, the base cordate, the apex usually acuminate or acute, sometimes 
rounded; distal leaves from broadly deltoid to lanceolate, 5-15 mm long, 3-10 
mm wide, the base cordate or rounded. Inflorescence loosely and narrowly cy- 
mose or narrowly thyrsoid. Involucres solitary or loosely clustered at the ends 
of branches, or solitary in forks of branches or axils of leaves, on slender pe- 
duncles up to 17 mm long, glandular-puberulent, widely bowl-shaped in fruit, 
much broader than deep, 5-9 mm long, the 5 bracts united by their margins 
1/3-1/2 their length, the lobes approximately equal, broadly triangular, 4-6 mm 
long, about as wide at the base, the apices acute. Perianth campanulate, purplish 
pink, pale pink, or occasionally white, sparsely viscid-puberulent externally, 
5-9mm long, about as wide, strongly constricted above the indurate base. Fruits 
3 per involucre, olive or dark brown and black-mottled or evenly black, broadly 
obovoid to nearly spherical, ca. 2.5-3.5 mm long, the width ca. 70-90% of the 
length, smooth or very slightly rugulose, sometimes faintly marked with 5 shal- 
low grooves (Fig. 1). 2n = 30); (Spellenberg & Soreng 5858). 

Distribution (Fig. 4).—Southern Nevada, southern Utah, and southern Colo- 
rado, south through Arizona, New Mexico, and western Texas to northern Chi- 
huahua, western Coahuila, and western Nuevo Leon, in open woods, on banks 
in woodland, among brush or boulders, usually where somewhat moist, 1500- 
2600 m. Flowering June-)August-October. 

The species is readily recognized by the distinctive shape of the leaves. At 
the apex of the petiole the base of the blade is broadly cuneate within the sinus 
of the overall cordate base, the curve at each side of the base of the blade revers- 
ing in a sinuate manner before joining the petiole. The apex of the blade is usu- 
ally acuminate. Very glandular-pubescent foes and glabrate plants may oc- 
cur in the same population (Spellenberg et al. 9681). Plants may be sufficiently 
viscid to “catch little birds” (label data, Vestal & Vestal 56). Leaf shape is consis- 


_ 


NYCTAGINACEAE, 545 


tent throughout the range except in Nuevo Leon, where leaves on some plants 
are cordate-truncate at the base, rounded at the tip. On these plants the stems 
are little-branched and apparently ascending. 

Plants were used by Native Americans to help heal “broken or bent” bones 
(label data, Vestal & Vestal 408). 


2. Mirabilis laevis (Benth.) Curran, Proc. Calif. Acad. Sci., ser. 2, 1:235. 1888. 
Oxybaphus laevis Benth., Bot. Voy. Sulphur 44. 1844. Hesperonia laevis (Benth.) 
Standl., Contr. U.S. Natl. Herb. 12:363.1909. Quamoclidion laeve (Benth.) Rydb., Bull. 
Torrey Bot. Club 29:687. 1902. Type: BAJA CALIFORNIA: Magdalena Bay, 1841, 
Hinds s.n. (HOLOTYPE: Ki; photos of holotype NMC). 

Plants few-stemmed and clambering through other vegetation to many 

stemmed and forming clumps as wide or wider than tall; stems from the previ- 

ous year often present and skeletal-white. Stems herbaceous or suffrutescent or 
clearly woody basally, 0.15-0.8 m long, erect or decumbent, repeatedly branched 
and appearing more or less dichotomous, glabrous, glabrate, puberulent, more 
or less scabrous, or viscid-villous, when pubescent, the pubescence denser dis- 
tally, hairs spreading or retrorse; internodes 0.5-11.5 cm long. Leaves more or 
less fleshy, pubescent like the stem; petioles 1-22 mm on basal leaves, becom- 
ing progressively shorter distally, 0-4 mm long on distal leaves; blades of the 
basal and midstem leaves ovate, deltoid-ovate, ovate-rhombic, or subreniform, 
1-4(-5.5) cm long, 0.5-3.5(-5) cm wide, the base cordate, truncate, or broadly 
obtuse, apex acute (occasionally attenuate), obtuse, or rounded, distal leaves 

lanceolate, lance-ovate, or ovate-rhombic, 5-17(-23) mm long, 3-11(-26) mm 

wide, the base cordate, truncate, or rounded. Inflorescences cymose or, in west- 

ern races, more or less thyrsoid by partial suppression of one of the pair of axes. 

Involucres clustered and nearly sessile at the ends of branches, or solitary in 

forks of branches or axils of leaves, on peduncles 3-12 mm long, campanulate, 

3-7 mm long in flower, enlarging about 1.5 x in fruit, the peduncles elongating 

slightly and deflexed; lobes of involucre 5, 1/3as long to equalling the length of 

the tubular portion, slightly unequal, narrowly to broadly triangular or trian- 
gular-lanceolate, the base 1/3 to equal to the height. Perianth widely flared from 

a narrow constriction atop the indurate base, white, white with magenta veins, 

pink, lavender, or magenta, sparingly puberulent externally, 10-16 mm long, in 

full anthesis usually slightly wider. Fruits | (rarely 2) per involucre, gray or dark 

brown to almost black, ovoid, obovoid, or almost spherical, 3-5.5 mm long, 3-4 

mm wide, glabrous, almost smooth to moderately rugose, often faintly mottled 

with darker brown or black, with or without 10 paler, diffuse longitudinal lines, 

becoming mucilaginous when wetted (Fig. 1). 

Distribution (Figs. 2, 3).—United States from central California and east- 
ern Oregon southward through southwestern Utah and central Arizona, south 
to Mexico in west-central Sonora and west-central Baja California Sur. 


— 


BRIT.ORG/SIDA 19(3) 


MIRABILIS LAEVIS 
oO var. CRASSIFOLIA 
O var. RETRORSA 

f @ var. VILLOSA 


Sennett” 


eeecy &> 
i ee L> i involucre lobes 
CALIF. i ‘ m= =or > than tube 
; » <than tube 
pubescence on stems 
ie and inflorescence 
BS < hairs conical or coarse 
DP Py — hairs long, often viscid 
D> - hairs slender, usually short, 
& PLD reflexed 


UTAH 


Fic. 2. Distribution of Mirabilis laevis varieties in western United States. Varieties as recognized in this treatment are 
77 £ 4 £, ras + ts. 4 i JL L | 


indicated by open or closed circles or sq , and ch p y sy 
tuisbhndl + et | n ee er eee 


Vs oe +) AA 4 Ufiad Dade, (1007) 


1 \ J / t t 


We recognize four intergrading varieties based on morphological differ- 
ences that serve to help delineate more or less distinct geographic races. For the 
past century authors have wrestled with the variation presented by these plants, 
some taking a rather conservative view and placing most forms in an inclusive 
Mirabilis laevis, others splitting variants as species or infraspecific taxa. Even 
on one of the syntypes (NY) of Mirabilis californica, the first of the variants to 


( NYCTAGINACEAE 547 


Sette See Zz < var. LAEVIS 

Oo var. CRASSIFOLIA 
o var. RETRORSA 
e var. VILLOSA 


involucre lobes 
m= =or > than tube 
» <than tube 
pubescence on stems 
and inflorescence 
< hairs conical or coarse 
— hairs long, often viscid 
~ hairs slender, usually short, 
reflexed 


118°145' 
GUADALUPE 
ISLAND 


24° 


Fic. 3. Distributi 


by open or closed circles, squares, or triangles, and characters of specimen from that site are indicated by symbols 
rs Bper es - 1 f = Hi ee 1 | 4 J YR 7 fb 94 I £, Dadri, (1003)\ 


J 


be split from M. laevis, the epithet “leavis Benth.” appears on the collecting label 
along with “Oxybaphus glabrifolius Vahl.” Curran (1888), in transferring 
Mirabilis laevis from Oxybaphus noted that plants recently brought from 
Magdalena Bay were “nearly but not quite glabrous, the inequality of the in- 
volucral lobes variable and often not greater than is found in our Californian 


548 BRIT.ORG/SIDA 19 


P< 


3) 


forms.” Wiggins (1964) placed all forms in the complex south of about 27° N in 
Baja California Sur into M. laevis, apparently ignoring populations to the north 
except for Mirabilis californica var. cedrosensis, which he noted to occur from 

San Clemente Island, California, southward on the west side of Baja California 
to the Vizcaino Desert area (ca. 27'N). In that work Wiggins noted the inland 
specimens of M. laevis to be “quite viscid-puberulent to short-villous and often 
have the coarser and almost retrorse scabrous hairs on the upper stems that 
occur in forms of M. californica.” 

What was previously considered Mirabilis californica now comprises M. 
laevis var. crassifolia, which usually has an inflorescence with a more or less 
well defined central axis and shorter lateral branches, the entire shape being 
irregularly conical. We are terming this inflorescence form “thyrsoid.” This con- 
trasts to the much more openly and symmetrically forked inflorescences of 
many of the inland populations. We call these “cymose.” The distinctions are 
not sharp. 


KEY THE VARIETIES OF MIRABILIS LAEVIS 


. Plants glabrous or with a few hairs in distal parts; lobes of involucre lanceolate, 
width of base 1/2—2/3 times the height of lobe; perianth probably magenta. 2a. M. laevis 


var. laevis 
1. Plants pubescent, rarely glabrate; lobes of involucre lanceolate to triangular, width 
of base 1/2-1 times height of lobe; perianth white, pink or magenta. 
2. Perianth pink or magenta, rarely white; width of lobe of involucre at base often 
/2 times height of lobe; plants pubescent, the pubescence usually not 
notably viscid nor retrorse (but hairs often stout and recurved along the coast); 


it iflorescet Ce COMIMNMOFL ily Mal lQVvy and more Or less thyrsoid. 2b. M. laevis 


var. crassifolia 
. Perianth he rarely pale pinewiety lobe of involucre at base 2/3-1 times height 
of lobe; plants commor ily notably viscid- pube scent or sparsely short- pubescent 
ds ae hairs; inflorescences broad, cymose 
. Plants viscid-pubescent, hairs spreading, ascending or sometimes retrorse. 
c.M. laevis var. villosa 
3. Plants puberulent, hairs retrorse. 2d.M. laevis var. retrorsa 


2a. Mirabilis laevis (Benth.) Curran var. laevis. 
Stems glabrous. Leaves glabrous adaxially, with a few short straight hairs 
abaxially; blades of the basal and midstem leaves ovate or deltoid-ovate, 3-4 
cm long, 2-3cm wide, apex acute; distal leaves lanceolate, lance-ovate, or rhom- 
bic-ovate, 7-14 mm long, 2-5 mm wide. Inflorescence narrowly thyrse-like, with 
a long main axis and shorter side branches bearing near their tips involucres 
borne singly or in small clusters. Involucres 7-10 mm long, sparsely short vis- 
cid-villous, the bracts united 1/3-1/2 their length, the lobes lanceolate or ovate- 
lanceolate, acute. Perianth magenta. Fruit almost spherical, 4.5mm long, 4mm 
wide (Fig. 1). 

Distribution (Fig. 3).—Apparently restricted to the vicinity of Magdalena 


NYCTAGINACEAE 549 


Bay, Baja California Sur, Mexico; habitat not recorded, 0-50? m. Flowering late 
winter and spring. 


2b. Mirabilis laevis var. crassifolia (Choisy) Spellenb., comb. nov. Oxybaphus 
glabrifolius Vahl var. crassifolius Choisy in DC., Prodr. 13(2):431. 1849. TyPE: NOVA 
CALIFORNIA: 1833, Douglas s.n. (HOLOTYPE: G-DC [microfiche RSA')). 


Mirabilis californica A. Gray ex cos in W. H. Emory, Rep. US. Mex. Bound. 2(1):169, 173, 
plate 48. 1859. Oxybaphus californicus (A. Gray) Hook. in Benth. & Hook. f., Gen. 
PL. 3:4.1880. Hesperoniaca Eee (ee cn US. Natl. Herb. 12:364. 
1909. TyPE: CALIFORNIA: San Diego, sand hills, 1850, CC. Parry s.n. (LECTOTYPE, here 
designated: NY!). In proposing the name, Gray states that M. californica came from 
“dry hills, San Diego, California,” and then indicates the collectors “Parry, Thurber.” 
Ona sheet from the Torrey Herbarium at NY are three specimens. One, at the top, 
collected in Los Angeles by Mr. Rich is of no further concern. At the bottom of the 
sheet are two portions of plants associated with a single label, indicating the plants 
were collected by C.C. Parry on “sand hills” in San Diego in 1850. Because the state- 
ment on the label most closely matches the habitat described, and the specimen 
at the lower right of the sheet clearly is the source of the illustration for plate 48 
(mistyped as plate 46 in the original description), we are selecting the material at 
the bottom of the sheet as the lectotype. SYNTYPES: San Diego, Wood Valley, May 
1852, Thurber 569 (2 specimens, GH!); Bigelow, Whipple Expedition, on the Colo- 
rado, 1853-54 (GHI, NY! [which represents the var. villosa (Kellogg) Spellenberg, 
as delimited herein]; Bigelow, 21 Mar 1954 (GH!). At the time of publication Gray 
questioned whether Oxybaphus laevis Benth. might be the same as his new spe- 
cies, noting that the species is “commonly more or less pubescent, and rarely 
glabrate.” Torrey (U.S. Rep. Expl. Miss. Pacific 4:131. 1857) assigned Bigelow speci- 
mens incorrectly to Oxybaphus glabrifolius Vahl. Gray, in the protologue of M. 
californica, cites Torrey’s listing, correcting the placement of these specimens. Gray 
later, in the protologue of M. bigelovii alludes to the Bigelow fe esis and indi- 

cates that this new taxon occurs “perhaps in California on the Color 

eT californica subsp. microphylla Standl., Contr. U.S. Natl. eh 12:365. 1909. 

E: MEXICO. LOWER CALIFORNIA: San Martin Island [off the west coast of the 
state of Baja California], 12 Mar 1897, Brandegee s.n. (HOLOTYPE: UC!). PARATYPE: 
LOWER CALIFORNIA: Ensenada, 26 Apr 1893, TS. Brandegee s.n. (UC). 

Hesperonia cedrosensis Standl., Contr. U.S. Natl. Herb. 12:362. 1909. Mirabilis cedrosensis 
(Standl.) Jepson, Fl. Calif. 459. 1914. M. californica var. pay E alae sea 
Contr. Gray Herb. 56:24. 1918. M. laevis var. cedrosensis Standl.) Munz, Man. South. 
Calif. 151. 1935. Type: MEXICO. BAJA CALIFORNIA: Cedros felaiel 3 Apr 1897, TS. 
Brandegee s.n. (HOLOTYPE: UC!). PARATYPES: CALIFORNIA: San Clemente Id., Oct. 
1902, Trask 14 (US). California: San Clemente Id., May 1903, Trask 193 (US)). 

Hesperonia heimerlii StandL., Contr. U.S. Natl. Herb. 13:412. 1911. Mirabilis heimerlii 
(Standley) Macbride, Contr. Gray Herb. 56:24. 1918. Type: BAJA CALIFORNIA: 
Guadalupe Island, S end of island, 3 Mar 1889, FE. Palmer 886 (HOLOTYPE: US! ISOTYPE 
GH!). 

Mirabilis laevis var. cordifolia Dunkle, Bull. S. Calif. Acad. Sci. 40:108. 1941. TYPE: CALI- 
FORNIA: San Clemente Island, Chinetti Canyon, 3 Apr 1939, Dunkle 7234 (HOLO- 
TYPE: RSA #350685! ISOTYPE RSA #464676!). Both specimens at RSA have been 
transferred from LAM; of the two the holotype has a hand written label with the 

ord “type.” Dunkle (1941) noted two variants in the same population, his new 
var. cordifolia intermixed with var. cedrosensis. 


550 BRIT.ORG/SIDA 19(3) 


Stems often glabrous basally, viscid-pubescent or more or less scabrous distally. 
Leaves puberulent, viscid-villous, or more or less scabrous, sometimes becom- 
ing glabrate with age (or occasionally glabrous); blades of the basal and midstem 
leaves ovate-rhombic, subreniform, or deltoid-ovate, 1-4.5 cm long, 0.4-3.5 cm 
wide, the apex obtuse or acute, occasionally rounded; distal leaves lanceolate, 
lance-ovate, or ovate-rhombic, 5-14 mm long, 2-7 mm wide. Inflorescence often 
rather thyrse-like after the first few dichotomous branches, the branches short, 
the involucres in clusters along a main axis. Involucres 5-9 mm long, densely 
short viscid-villous or sometimes slightly scabrid, the bracts united 1/3-2/3 their 
length, the lobes ovate or ovate-oblong, obtuse or acute. Perianth pink, lavender, 
magenta, occasionally white. Fruit ovoid, 3-5 mm long, 2.7-3.7 mm wide, dark 
to pale gray-brown and mottled with dark gray-brown, tan, or red-brown, some- 
times faintly and irregularly pale-striped (Fig. L). 

Distribution (Figs. 2, 3).—West-central California south along the coast, on 
the Channel Islands, and in the Coast Ranges to the Viscaino Desert, Baja Califor- 
nia Sur and the coastal islands, Mexico; coastal bluffs, road banks, coastal scrub, 
grasslands, chaparral, oak woodland, often on rocky outcrops, 0-1830 m. Flow- 
ering most of the year, most vigorously in spring. 

A particularly difficult area with regard to variation is around the southern 
end of the Sierra Nevada in California, where three of the varieties are in contact. 
Howell 38179, from the Lake Isabella region in Kern Co. illustrates very well the 
problematic classification of some specimens. It was first left unidentified in 
Mirabilis, then sometime later placed in an inclusive M. laevis; 9 years later was 
identified as M. retrorsa; 21 years later as M. bigelovii var. bigelovii, and shortly 
later placed ina variable var. crassifolia. The specimen combines the spreading 
leaves and (light) villous pubescence of var. bigelovii, the pointed leaves and 
rather long involucre lobes of var. crassifolia,and some retrorse hairs similar to 
var. retrorsa. Flower color was not feren 2 the cone and) is not evident from 
the specimen, but as judged from Spell from this area, flowers 
were probably white. Nearby, from the entrance to Kern River Canyon, comes 
Howell 38142, a late-season collection showing thyrsoid inflorescences of var. 
crassifolia, and fairly pointed leaves, but in other respects is the var. villosa; a 
similar, later collection (Howell 38675) from the same area notes “calyx rose.” For 
plants from the lower reaches of the Kern River Canyon, Spellenberg’s observa- 
tions note flowers only pink to rose, yet if these plants were collected eastward 
they easily would be placed in more consistently white-flowered var. villosa. 
Twisselman 8391, from the same area, is more or less villous and has blunt leaves, 
in these respects similar to the var. villosa, but it has a more or less accrescent 
involucre with proportionately longer lobes, more reminiscent of var. crassifolia. 
Al mingly intergradient to var. villosa is Bedell 74-5 and Twisselmann 198, 
both from the north end of the Temblor Range in western Kern Co. they have 


os 


NYCTAGINACEAE 551 


thick fleshy leaves that are blunt, short involucre lobes, but thyrsoid inflores- 
cences; [wisselmann notes that flowers are “purple.” 

Mirabilis laevis var. crassifolia may also intergrade with M. oligantha in Baja 
California Sur (Moran ) 

Much has been made of features of pubescence over the taxonomic history 
of this group. Even within var. crassifolia there is considerable variation. Near 
the coast, and particularly on the islands of northwestern Mexico, hairs are stout 
ane coun distinguishing ercoma cedrosensis and subsequent synonyms. 

type is thoroughly dient to finer but still conical-based 
ae common within the var. crassifolia. Fruit characteristics, such as those used 
to distinguish H. heimerlii, also an island population, seem to be completely 
inconsistent from population to population. Other specimen-based discussion 
focuses individually on characteristics that have been used to distinguish species 
in this complex and is found in Appendix 1. 

The label on a specimen collected in Baja California (Moran 12832) notes 
the indigenous name and use “Yerba del Empacho.-bueno para el estomacho.” 
The vernacular name is repeated on Moran 23821 from Baja California Sur. 


2c. Mirabilis laevis var. villosa (Kellogg) Spellenb., comb. nov. Mirabilis californica 
var. villosa Kellogg, Proc. Calif. Acad. Sci. 3:10. 1863. TYPE: CALIFORNIA. MONO 
Co.: Calif. Hwy. 182 10 m from Nevada border, Devil’s Gate, canyon of East Walker 
River, Spellen berg 12326, 12 Jun 1996 (NEOTYPE, here designated: NMC! ISONEOTYPES 
(!: BYU, CAS, F K, MEXU, MO, NY, RM, RSA, UC, US). Kellogg (1863) provides brief 
but clear description of a plant that matches the classic concept of Mirabilis 
bigelovii except that he notes his var. villosa to have a “pink perigonium” (rather 
than white). He notes the taxon tocome “from the interior-Devil’s Gate and Carson 
River...” but he cites no documenting specimens. We found no specimens that were 
seen by Kellogg, all perhaps having been destroyed in the 1906 San Francisco fire. 
During field work involving this complex Spellenberg could not find Mirabilis at 
the well known “Devil's Gate” along U.S. Hwy. 395 north of Bridgeport, Mono Co., 
Calif., nor were plants to be found along the upper portions of the West Walker 
River along this highway. In either place habitat seems incorrect. They do occur, 
however, at a lesser known “Devil’s Gate” along the East Walker River (DeLorme 
Mapping 1990) northeast of Bridgeport, and this is presumed to be Kellogg's Devil's 
Gate. Plants from this site form the neotype series. 

Mirabilis bigelovii A. Gray, Proc. Amer. Acad. 21:413, 1886. Hesperonia bigelovii (A. 
Gray) Standl. N. Amer. FI. 21:235. 1918. TYPE: Grand Canyon, May 1885, A. Gray s.n. 
(HOLOTYPE: GH1!). Gray cites his own collection in the protologue, indicating also 
that the type comes from “below Peach Spring” where the species is “common.” 
The holotype has penned in Gray’s hand on a printed label “Grand Canon” and 
“Mirabilis bigelovii n. sp.,” but does not mention Peach Spring. In the protologue 
Gray alludes to Bigelow specimens, as discussed herein in the nomenclatural sec- 
tion under Mirabilis californica. 

Mirabilis aspera Greene, Erythea 4:67. 1896. Mirabilis californica subsp. aspera 
(Greene) Parish, Muhlenbergia 3:125. 1907. Hesperonia aspera (Greene) Standl., 
Contr. U.S. Natl. Herb. 12:362. 1909. M. californica var. aspera (Greene) Jepson, FI. 
Calif. 458. 1914. M. laevis var. aspera (Greene) Jepson, Man. Fl. Pl. Calif. 340. 1923. 


552 BRIT.ORG/SIDA 19(3) 


M. bigelovii var. aspera (Greene) Munz, Man. S. Calif. 151.1935. Type: CALILFORNIA: 
Mojave Desert, 14 Jun 1895, Parish 3757 (HOLOTYPE: ND-G, photocopy at NMCI; 
ISOTYPES: GHI, UCI). Jepson (1914) indicates the type locality to be Hesperia, San 
Bernardino Co., California. 

Mirabilis glutinosa A. Nelson, Proc. Biol. Soc. Wash. 17:92. 1905 [nomen illeg., later 
homonym of M. glutinosa Kuntze, Rev. Gen. 3(2):265, 1898, a Bolivian plant. 
Hesperonia glutinosa Stand, Contr. US. Natl. Herb. 12:365. 1909. Mirabilis limosa 
A. Nelson, Bot. Gaz. 47:426.1909[a substitute name for the earlier illegitimate name}; 
ESproue ee “ Nesow Standl, Muhlenbergia 5:104. 1909 [a superfluous 

name]. ifornica var. glutinosa Jepson, FL. Calif, 1:458. 1914. M. laevis 
var. eco sates lca Man. FI. Pl. Calif. 340.1923. M. laevis subsp. glutinosa 
1) E. Murray, Kalmia 13:32. 1983. (same combination in Kalmia 12: 22,1982, 
eee on sea basionym). TYPE: NEVADA: Karshaw, Meadow Valley Wash, 27 
May 1902, Goodding 967 (HOLOTYPE: RMI; IsoTYPES: DS!, MO!, NY!, POM!, UC). The 
paratypes cited (NEVADA. WASHOE Co. Pyramid Lake, 9 Jun 1903, G.H. True 758 
[RM!; UTAH: St. George, 13 May 1902, L.N. Goodding 778 [RM!|, duplicates at GH}, 
MO) are the var. retrorsa in the present treatment). 

Hesperonia aspera subsp. villosa Standley, Contr. U.S. Natl. Herb. 12:363. 1909. TyPE: 
CALIFORNIA: Mohave Desert, Cushenberry Springs, 2 Jun 1901, S. B. Parish 4940 
(HOLOTYPE: USk ISOTYPE: NY!). PARATYPES: CALIFORNIA: Providence Mts., 26 May 
1902, T'S. Brandegee s.n. (UC). CALIFORNIA: Argus Mts., Shepherd Canyon, 30 Apr 
1891, Coville & Funston 741 (NY!, US). 

Hesperonia glutinosa subsp. gracilis Standl., Contr. U.S. Natl. Herb. 12:365. 1909. H. 
limosa subsp. gracilis (Stand1.) StandL, Muhlenbergia 5:104. 1909. Typ: ARIZONA, 
Sabino Canyon, 1892, ].S. loumey 471c (HOLOTYPE: US!). PARATYPES: ARIZONA: with- 
out locality, 1876, Palmer 644 (US). ARIZONA: Tempe, 6 Apr 1896, J.S. eee. Sn. 
(UC! - 2 sheets). CALIFORNIA: Colton, Feb 1881, G.R. Vasey s.n. (US! - 3 sheets, 2 
apparently seen by Standley, 1 possibly not). NEW MEXICO: without ae G.R 
Vasey s.n., 1881 (US!) [as Standley notes in the discussion of this paratype, the loca- 
tion is probably incorrect; Standley knew of no isc from New Mexico, and 

ne have been seen in the present study. Ewan and Ewan (1981) indicate that 
ee was in central and northern New ee well aw ay from nearest known 
populations in Arizona, and that specimens were apparently sent back to Wash- 
ington unlabelled, increasing the possibility of error. To Spellenberg, the New 
s very similar to Vasey paratypes from Colton, California, 

eine US #226311 


Stems moderately to densely villous or viscid-villous, often with wavy hairs, or 
puberulent with + retrorse hairs, the pubescence denser and increasingly viscid 
distally. Leaves + fleshy, viscid-villous; blades of the basal and midstem leaves 
reniform-ovate, broadly deltoid-ovate, or suborbicular, 0.5-4 cm long, 1-3.7 mm 
wide. Inflorescence usually cymose, the branches + equal throughout; involucres 
5-7 mm long, the bracts united ca. 2/3 their length, the lobes ovate-triangular or 
ovate-oblong, obtuse or acute. Perianth white or pale pink, occasionally (espe- 
cially in far western part of range) deep pink or purple. Fruit ellipsoid to obovoid 
or almost spherical, 4-6 mm long, 2.5-4 mm wide, gray-brown, dark charcoal- 
brown, or olive, often dark-mottled, often faintly marked with 10 paler longi- 
tudinal lines (Fig. 3). 2n = 30) (Spellenberg 5444) 


PELLENBERG AN { NYCTAGINACEAE 553 


Distribution (Figs. 2, 3).—Southeastern Oregon through Nevada, southwest- 
ern Utah, southern California (primarily southeastern, but extending west as 
far as eastern San Luis Obispo Co.), western Arizona to Baja California and north- 
western Sonora; roadbanks, slopes, open desert, often among brush or in open 
woodland, 35-2200 m. Flowering most of the year, most vigorously in spring. 

In California the var. villosa (as M. bigelovii), has been considered to be from 
east of the Sierra Nevada and the Transverse Range. Nevertheless, around the 
San Joaquin Valley some plants of the var. crassifolia approach the var. villosa 
(e.g., Ewan 10309; Hoover 3170, Raven et al 9240, Twisselmann 8377) or cannot be 
excluded from it as here defined (e.g., Bacigalupi et al. 5205; East wood & Howell 
5839; Ferris & Bacigalupi 10350; Keck 2158). 

A vernacular name in Baja California recorded for this species is “Yerba de 
la Vieja” (Moran 23774). 

From the type locality to the north plants are sporadic along the East 
Walker River and along the West Walker River where it exits from the Sierra 
Nevada and pinon pine vegetation into the Great Basin and its shrub associa- 
tion (Spellenberg 12331, 12332). Other plants in the region have much shorter, 
sparser pubescence and are more readily referable to the var. retrorsa 
(Spellenberg 12327,12329, 12333). No obvious habitat differences were detected 
between the two pubescence phases. Collection 12333 had flowers closed in mid- 
morning that were very slightly pinkish. Otherwise, all plants seen in flower 
had white perianths. 

In discussing M. aspera on the Colorado and Mojave deserts, Parish (1907) 
notes intergradation along edges of range with M. californica, and places the 
former into the latter as a subspecies. Intergradation is particularly evident in 
perianth color - those plants from the zone of contact having pink (rather than 
red-purple or white) perianths. To the west, in the var. crassifolia, perianth color 
is usually red-violet, but white-flowered plants are known. To the east the var. 
villosa usually hasa white perianth, occasionally witha pale pink tube, or rarely 
entirely pale pink. The pattern probably results from selection pressure of pri- 
marily diurnal pon aior in the west and nocturnal eDOUNIAHOTS in the drier 
deserts to the east (Baker [1961] di various polli Mirabilis froebelii 
(Behr) Greene, a species with red-violet Flowers) 

The pivotal nature of the var. villosa in the Mirabilis leavis complex is in- 
dicated by its extensive synonymy. As indicated by the discussion of variation 
as seen in various specimens (Appendix 1), the variety is variable and often inter- 
grades with var. crassifolia and the var. retrorsa. In southeastern California and 
Baja California it is sometimes distinguished with difficulty from M. tenuiloba. 
2d. Mirabilis laevis var. retrorsa (Heller) Jepson, Man. FI. Pl. Calif. 340. 1923. M. 

retrorsd Heller, Muhlenbergia 2:193. 1906. Hesperonia glutinosa subsp. retrorsa 
(Heller) StandL., Contr. U.S. Natl. Herb. 12:365. 1909. H. limosa subsp. retrorsa (Heller) 
Standl., Muhlenbergia 5:104. 1909. Mirabilis californica var. retrorsa (Heller) 


554 BRIT.ORG/SIDA 19(3) 


Jepson, Fl. Calif. 458. 1914; Hesperonia retrorsa (Heller) StandL, N. A. Fl. 21:236. 
1918. M. bigelovii var. retrorsa (Heller) Munz, Man. S. Calif. 151. 1935. Type: CALI- 
FORNIA. MONo Co. near the Southern Belle Mine, 25 May 1906, Heller 8336 (HO- 
LOTYPE: BKL on indefinite loan to NY! lsoTyPEs: DS!, GH!, NY!, MOI, US). 
Stems glabrous or with a few retrorse hairs below, sparsely to densely retrorse- 
puberulent distally, when densely pubescent, then often also + viscid. Leaves + 
fleshy, puberulent with retrorse hairs; blades of the basal and midstem leaves 
reniform-ovate, broadly deltoid ovate, or suborbicular, occasional orbicular- 
reniform, 0.5-3.5 cm long, 1-3.4 cm wide. Inflorescence usually cymose, the 
branches + equal throughout; involucres 5-7 mm long, the bracts united ca. 2/3 
their length, the lobes ovate-triangular or ovate-oblong, obtuse or acute. Peri- 
anth white or occasionally white tinged with pink at the base, rarely entirely 
pale pink. Fruit ellipsoid to obovoid or + spherical, 3.5-5 mm long, 2.6-4 mm 
wide, occasionally slightly wider than long, gray-brown, dark charcoal-brown, 
or olive, occasionally dark-mottled, often faintly marked with 10 paler longi- 
tudinal lines (Fig. 1). 2n = 31-33] (Strother 1256). 

Distribution (Figs. 2, 3).—Southeastern Oregon, western and southern Ne- 
vada, southwestern Utah, northwestern Arizona, southern California, and 
northern Baja California; arid open areas among desert brush or in open wood- 
land, often on banks, 60-2000 m. Flowering in spring, occasionally in winter, 
less frequently at other times. 

In general, plants of the var. retrorsa are smaller, with smaller leaves, and 
apparently are more compact, providing more of a forking, repeating “wish- 
bone” aspect (Bagley 2098, Clemon and Jonsson 1690, Clokey & Templeton 5725, 
Munz 16449), than most of those of var. villosa. Nevertheless, open sprawling 
plants with stems 3.5-4 dm long, with leaves 2+ cm long, and inflorescences + 
thryse-like (Peirson 7180), resemble in aspect either the var. crassifolia or the 
var. villosa. Local environmental factors may also affect the phenotype; e.g., 
Munz & Keck 4754 is a lanky plant with broad thin leaves. It is said to come 
from “among rocks along canyon” and may be a shade form. Plants indistin- 
guishable from the tighter, smaller, northern forms of this variety occur as far 
south as the mountains of southern California (Peirson 9846) and Baja California 
(Moran 14842). 

The variety retrorsa may co-occur with the var. villosa (see two specimens 
at DUD, Train s.n., 30 Apr 1937, both from Darwin Falls Canyon; also Duran 
3455 [retrorsa] and Mooney et al. 132 [villosa], both from Silver Canyon in the 
White Mountains). Munz noted his collections 13036 (var. retrorsa) to be not 
glutinous, 13037 (var. villosa) from the same site to be glutinous. The Duran 
3455 specimen cited immediately above has long internodes and spreading 
rounded leaves more typical of var. villosa, but has very short, mostly retrorse 
hairs; in respect to habit and pubescence it is intermediate between the two 
varieties. Mixed collections of the two are represented by M.& E. Eplings.n.and 


(GUEZ, NYCTAGINACEAE, 555 


Maguire & Holmgren 25193. The two also occur in close vicinity on the east side 
of the Sierra Juarez in northern Baja California (Thorne, Boyd, et al. 61758 = var. 
retrorsa; Thorne et al. 5/784 = var. villosa). 

As discussed for the var. crassifolia, Kern Co., California, is also an area of 
particular difficulty concerning the var. retrorsa. Numerous collections sug- 
gest intergradation with the var. villosa; e.g., very dense pubescence, clearly ret- 
rorse, is present in Eastwood 3200; on Hall and Chandler 6882, a similar plant 
from the same general region, the collectors note that the flowers are pure white 
and the plants are viscid. Further ene the difficulty of satisfactorily clas- 
sifying material from this area, tw cimens collected very near one another 
a week apart in the same year ae represent a different variety; Voegelin 67 is 
nearest the var. crassifolia, whereas Cole and Voegelin 120 is clearly var. retrorsa. 
Another pair of specimens from the same vicinity, in Red Rock Canyon (vicin- 
ity of Red Rock Canyon State Park) are the var. retrorsa (Abrams 11877) and a 
fairly lightly pubescent phase of the var. villosa (Munz 1246). Howell 37115, in 
its fairly dense but downward-flexed pubescence, approaches the var. villosa, 
and in its pointed leaves the var. crassifolia (flowers on the specimen appear to 
have been white). In this region of contact between the three varieties, + typical 
plants of the var. retrorsa occur (Howell 38667). 


3. Mirabilis tenuiloba S. Watson, Proc. Amer. Acad. Arts 17:375. 1882. Hesperonia 
tenuiloba (S. Wats.) Standley, Contr. U.S. Natl Herb. 12:363. 1909. Type: SOUTHERN 
CALIFORNIA: San Bernardino, 1880, W.G. Wright 106 (HOLOTYPE: GH}; photo and 
fragment of holotype at DS!). According to note on labels of Parish 6072, the type 
locality is in West Canyon, western edge of the Colorado Desert, Riverside Co. 

Plants forming leafy clumps 0.3-1 m or more in diameter, usually with many 

stems, herbaceous or somewhat sulfrutescent basally. Stems ascending 0.2-1m 

long, with few to many ascending branches, pale green or white at base, green 
distally, puberulent in lines or throughout, usually glandular-viscid, the pu- 
bescence denser distally. Leaves slightly fleshy; petioles to 2.2(-5) cm long on 
basal leaves, becoming progressively shorter distally, the distal leaves sessile or 
on petioles to 4 mm long and gradually intergrading to the bracts of the inflo- 
rescence; blades of basal and midstem leaves broadly deltoid or ovate, the larg- 
est often wider than long, 2-5(-8) cm long, 1.7-7.0(-12) cm wide, glabrate to 
glandular villous, the base rounded to cordate, the apex usually acute, some- 
times rounded; distal leaves from broadly deltoid to lanceolate, often acumi- 
nate, 1-2 cm long, 7-15 mm wide, the base cordate or rounded. Inflorescence 
usually narrowly thyrsoid. Involucres densely clustered among distal leaves or 
bracts near ends of branches, on peduncles 0-2 mm long, glandular-pubescent, 
narrowly campanulate, deeper than broad, 7- 16 mm long, the 5 bracts united 
by margins 1/3-1/2 their length, the lol ly equal, narrowly lance- 
oblong, 1/5-1/4 as wide at the base, the apices acute or attenuate. Perianth cam- 


556 BRIT.ORG/SIDA 19(3) 


panulate, white (rarely pink), sparsely viscid-puberulent externally, 13-18 mm 
long, about as wide, strongly constricted above the indurate base. Fruits 1 per 
involucre, dull reddish brown to almost black, rarely with 10 inconspicuous 
and very slightly paler lines, broadly ovoid to nearly spherical, 4-6 mm long, 
the width 60-85% of the length, smooth or very slightly rugulose, sometimes 
faintly marked with very shallow grooves (Fig. ). 

Distribution (Fig. 4).—Southern California, southwestern Arizona, and 
northwestern Sonora, south to Baja California Sur, on slopes, canyon sides, cliffs, 
and among rocks, or in gravel or sand in semi-arid and arid areas, 0-400(-900) 
m. Flowering late winter and spring, occasionally other times. 

The species is from east of the mountains in southern California and from 
near the gulf in Baja California and therefore is mostly a desert species. It is 
known in Arizona only from the Tinajas Altas Mountains in Yuma Co., where it 
was collected in 1940 by L. Goodding (s.n., 7 Mar 1940), the collection remaining 
unidentified for more than 50 years. It was rediscovered (Felger & Broyles 92- 
613) and reported from there by Felger (1993). As noted by Felger, the species 
was sympatric with M. bigelovii (= M. laevis var. villosa) (Felger & Broyles 92- 
614). A.and R. Nelson apparently collected M. laevis var. retrorsa (3236, but as 
M. limosa) in sympatry with M. tenuiloba (3236a), perhaps separating the col- 
lections later under the “a” number. The Nelson collection of M. tenuiloba 3236a 
has leaf tips more rounded than usual for the species, plants are less robust, and 
involucres in the shorter portion of the range for the species. It may be an 
introgressed plant. Gander 1301, a robust, more “typical” M. tenuiloba, among a 
number of other collections, is from the same canyon. Sympatry involving such 
similar perennial species provides the opportunity for hybridization. Occasional 
collections such as Moran 8877 have involucres with triangular teeth 3-4 mm 
long, shorter than the tube, also suggesting intergradation with M. laevis. 

MacBride (1918) considered Chandler 5332, from near Escondido in south- 
ern California, to be included in his concept of M. tenuiloba var. polyphylla, the 
only record north of Mexico for this entity. That specimen is here considered to be 
an extreme form of M. laevis var. crassifolia. Other somewhat similar specimens, 
having at maturity rather large involucres for M. laevis var. crassifolia, are from 
the Channel Islands (see Blakley 5238, Clokey 4923, Raven 17655). 

The southernmost collection in Baja California Sur (Wiggins et al. 258) is 
much less pubescent that is characteristic of M. tenuiloba. The specimen was 
originally identified as M. oligantha. 


— 


4. Mirabilis oligantha (Standley) J. Macbride, Contr. Gray Herb. 56:23. 1918. 
Hesperonia oligantha Standley, Contr. U.S. Natl. Herb. 12:363. 1909. Type: BAJA 
CALIFORNIA: Calmalli, Jan-Mar 1898, Purpus 82 (HOLOTYPE: UCI). 

Hesperonia polyphylla Standley, Contr. U.S. Natl. Herb. 12:363-364. 1909. Mirabilis 
polyphylla (Standley) Standley, Publ. Field. Mus. Nat. Hist, Bot. Ser. 8:306. 1931. M. 
tenuiloba var. polyphylla (Standley) J.E Macbride, Contr. Gray Herb. 56:23. 1918. 


CDCIICAIDCD ARKIN DANDICIICT NYCTAGINACEAE 557 


Nev. Utah Colo. 
Calif. ee a 
| | ® 
rT 7 ee a 
z | 
a @ 
= Tex. 
a 
U.S.A. = 
MEXICO n N.M 
: 8 
N 
‘| 
a 
— N.L. 
@ M. OLIGANTHA ct ° 
BM. OXYBAPHOIDES 
OM. TENUILOBA 
Lo 
Fic. 4. Distributi f Mirabilis oligantha,M oxybaphoides, and M A 


TYPE: BAJA CALIFORNIA: San Borga, 6 May 1889, TS. Brandegee s.n. (HOLOTYPE: 
UCI). PARATYPE: LOWER CALIFORNIA: Gulf of California, Los Angeles Bay, 1887, 
Palmer 600 (UC! on same sheet as holotype; duplicate of paratype at GHI, with 
month given as Dec). 
Plants usually dense shrubs or subshrubs. Stems erect, ascending or spreading, 
0.3-1.2 m long, repeatedly branched, with a whitish or gray exfoliating bark on 
older stems, glandular-puberulent, densely so distally, becoming glabrate with 
age. Leaves slightly fleshy; petioles 1-20 mm long, about 1/5-1/3 the length of 
the blade, becoming progressively shorter distally; blades of the midstem leaves 
broadly deltoid-ovate or ovate, about 2-5 cm long, 1.5-3 cm wide, sparsely to 
densely glandular puberulent, the base subcordate, rounded, or broadly cuneate, 
the apex acute, obtuse, or sometimes rounded; distal leaves progressively re- 
duced from midstem leaves, from ovate to lanceolate, those among the flowers 
as small as 5mm long, 2mm wide, witha petiole of 1 mm long, the base rounded 
tocuneate. Inflorescence when well developed widely branching, the main axis 


558 BRIT.ORG/SIDA 19(3) 


zig-zag, or sometimes comparatively dense and thyrsoid. Involucres or solitary 
in forks of branches or axils of leaves, on slender peduncles 4-15 mm long that 
are deflexed after anthesis, glandular-puberulent, narrowly urn-shaped in 
flower, distended by the globose fruit, 7-10 mm long, the 5 bracts united by 
margins aboutl/2 their length, the lobes narrowly triangular or lanceolate, 4-5 
mim long, 1/3-1/2 as wide at the base, the apices acute. Perianth campanulate, 
usually white, less often pinkish or lavender, sparsely puberulent externally, 
especially on the tube, 12-20 mm long, about as wide, strongly constricted above 
the indurate base. Fruits | per involucre, dark brown or nearly black, sometimes 
with 5 faint paler lines, ellipsoid, 6-8 mm long, smooth or slightly rugulose, 
sometimes faintly marked with 5 shallow grooves (Fig. 1). 

Distribution (Fig. +).—Central Baja California and northern Baja Califor- 
nia Sur, on dry rocky slopes among desert shrubs and cacti, 50-600 m. Flower- 
ing fall to early spring, sometimes later. 

Mirabilis oligantha is an endemic to the Baja California peninsula. A puz- 
zling series of collections, mostly identified originally as M. bigelovii, come from 
the mountains of northern Baja California Sur, particularly from the vicinity of 
Picachos de Santa Clara, where M. oligantha has been collected (Gentry 7717). 
These plants may not have been so shrubby and stems may have been sprawl- 
ing. They have rather sparse foliage, the progressively reduced leaves in the in- 


| 


florescence 
singly or in few-flowered clusters. Involucres are small for M. oligantha, but have 
long lobes. Flower color noted on labels is white, pink, or lavender. Mirabilis 
laevis var. crassifolia occurs in the region, and the plants may represent 
intergradient forms. Such specimens have been annotated as that variety, with 
the note that they may be intergradient (Gentry 7697, Moran 18723, Moran & 
Reveal 19671, 19689). 

Standley (1909) distinguished Hesperonia oligantha from other species in 
his key in part by stating that flowers are “purplish red.” He also noted that 
stamens are “long exserted” in M. oligantha. Though the stamens are exserted 
somewhat in the species, as judged from herbarium specimens, the “long ex- 
serted” impression comes from Standley mistaking flowers as Hesperonia that 
actually are from some gamopetalous family, not Nyctaginaceae, attached as 
fragments to the holotype sheet. These flowers appear to have been dark in color, 
In 1911 and 1918 Standley did not mention flower color. Wiggins (1964) explic- 
itly noted that M. tenuiloba hasa white perianth and scarcely exserted stamens, 
and that M. oligantha Gncluding M. polyphylla) has a white (or pink?) perianth, 
but for that species there is no mention of stamens. Of the 15 collections of M. 
oligantha seen, labels of five report the perianth as white or creamy white. One 
reports “white, slightly pinkish” (Moran 23808) and another “pale lavender” 
(Gentry & Fox 11731) 


istic of several M.oligantha specimens, and flowers borne 


ra 


NYCTAGINACEAE, 559 


APPENDIX 1 


Specimen-based discussion on characteristics that have been used to dis- 
tinguish taxa in the Mirabilis laevis complex. Plants vary in many fea- 
tures, and often a collection used to illustrate one point also illustrates 
others. Specimen citations comprise Appendix 2 

Mirabilis laevis var. crassifolia 

Pubescence of var. crassifolia.—The ifoliais variable ithin por 
lations (Werff 422] notes sane snl y ganda ornes not and intergrades with the vars. 
retrorsaand villosa to the eas e\ ifoli 


broader at the 
base, somewhat or consider coarser than ie fines pubescence of the var. villosa or the short 
retrorse ne of the var. retrorsa. Intergra more villous var. villosa occur ee much of 
range and are maintained in var. pipes id eee because of their relatively acute lower leaves, 
often comparatively long involucre lobes, and more or less thyrsoid in orescence (Daniel 1345; 
Gentry 8886; Templeton 11388; Thorne & Tilforth 41536). Such pl common in the 
southern California mountains and in Baja California. Others, Roe See from interior Baja Cali- 
fornia, have blunter leaves and slightly finer pubescence than coastal plants, and in this respect be- 
gin toapproach the var. villosa (Burgess 6095; Carter, Alexander & Kellogg 2522; Moran 18694). These 
are retained in the var. crassifolia because of generally thyrsoid inflorescences and magenta or pink 
flowers. In California, specimen from near the coast in vote Co. ae most of the character istics of 
nae var. eee Le., long internodes, rounded leaves, vill | ly longer 
involucre lobes as in the var. crassifolia and flowers that are magenta to layende: (Thompsom 1857), 
plants to the north in Santa Barbara Co., are also similar (Pollard s.n., 30 Sep 1956). Others from this 
region have more acute leaves and proportionately narrower and longer involucre lobes as in the var. 
crassifolia (Bourell et al. 2938), contrasting with other plants in the on such as Hoover 7644, which 
has rounded lower leaves and proportionately short involucre lobes, but which has strongly taper- 
ing hairs more consistent with var. crassifolia. E ey lightly pubescent plants that have ae 
a and rather long involucre lobes occur on western edge of the California desert (Dunkle 341] 


eee (1909) established Hesperonia cedrosensis in large part on the conspicuous, ee 
recu a hairs, the extreme in this fescue zeal in+ i plants as occur on Cedros Island 
eres 14453). Plants with such hairs are almost entirely coastal but not necessarily insular 
(Standley 1918). Plants of the Viscaino region in central Baja California may have pubescence similar 
to the stout recurved trichomes of plants from Cedros Island (Boyd, Ross & Appleby 8100; Gentry 739D, 
as do plants along the northern Pacific coast of Baja California, which have notably pointed leaves 
(Chisaki & Newcomb 525; Epling & Robinson s.n., 15 Feb 1935). Specimens that have stout recurved 
hairs interspersed with finer hairs occur on the Channel Islands (Brandegee s.n., 25 Aug 1894; Munz 
6645) and in 1 the southern par of the eae ve ane 176). ees oon sea same pees show sous 
recurved h nd] 846 
Sea ata al but are more (Eastwood 6387) or ee (ireadione 2874) viscid-villous. The in i 
rounded lower leaves reminiscent of the var. villosa, as do many plants on the Channel Islands, where 
larger, but not especially recurved, hairs may be mixed with a fine glandular hairs (Raven 17307). A 
very villous plant from these islands, thus similar to var. villosa, ia long involucre lobes character- 
istic of var. ii ae okameniae: Plants with nical curved hairs may occur 
nland to the north, as in Fresno Co. (Boolootian s.n., 6 . 1951). 
Plants var. crassifolia are not completely distinct from the var. retrorsa. Howell 39241, from 
Monterrey Co., isan open sprawling pent with pointed leaves as expected in the var. crassifolia, but 
has short calyx lol ol 


to those of the var. retrorsa. Other collec- 
tions, but from southern California, also well away from the main range ss eu var. ue rsd are Renee 


pubescent and | but have longi 


560 BRIT.ORG/SIDA 19(3) 


the var. crassifolia (Epling & Ellison s.n., 28 Mar 1930). Plants on the west slope of the southern Sierra 
Nevada approach var. villosa in their more or less villous pubescence, the hairs of which may be 
somewhat deflexed, and in their blunt pie A ea 3214, Hoover 3170); similar combinations at 
characteristics are found at the southern end 0 where tl 

the three varieties come into contact Jepson a Thorne 31702). Alene the eee zone of i vat. 
retrorsa with the var. crassifolia in southern California are plants with retrorse hairs and white flow- 
ers, but with pointed leaves and rather long involucre lobes and conical hairs (Kamb 902). 

A sparsely pubescent, lanky, very thin-leaved plant (Munz et al. 2672) appears to be an envi- 
ronmentally modified phase, having been collected on a “damp hillside.” 


Flower color of var. crassi one —Usually var. crassifolia has a deep rose or magenta perianth, 
- DP g | 


5 f tana: vith thi Tit stly white flowered Val. villosa 
Gas tistaciontl ee) Neve silicic ind contrary tothe ke He: ia by Standley (909), within 
y f 


the range of var. crassifolia, sporadic variants with pale or white eens airly frequent. “Flowers 
vary from purple to white” in populations in the northern part of range (Merced Co., Lyon 932) or 
“pale white with rose tint along veins” (San Benito Co., Ewan 10309, a plant very closely approaching 
var. villosa). Toward the southern end of the range plants with white (Gentry 8694, Moran & Reveal 
19671, Orcutt 219a; Reeder & Reeder 7259), almost white (Gray s.n.), white tinged with pink (Moran 
& Reveal 20006), pure white oe red (Trask s.n.), white to light lavender (Henrickson 8940), or pale 
lavender (Moran & Reveal 19870) {1 


— 


sometimes in mixed populations (Ewan 7041 = a 
7042 = pink, Mordn 20414 = pink, 20415 = whe Pale-flowered, white-flowered, or mixed, populations 
in this region and along the eastern edge of the range are probably a response to selection pressures 
from nocturnal pollinators in the desert (e.g. Jepson 6073, 8859). Whitish-flowered plants also occur 
in the coastal scrub of Baja California (Hodgson & Pinkava 3011); white, pink and “red” flowers occur 
in the same population near the coast in southern California (Hastings s.n.16 Apr 1941), and plants 
may occasionally have flowers white with red veins (Trask 193). Trask 14, however, from the same 


general locality, has magenta flowers. The last two specimens are paratypes of H. cedrosensis. 


Leaf shape of var. crassifolia.—In an attempt to distinguish species in the complex, authors have 
reiterated features emphasized by Standley (1918), where leaves of the var. crassifolia (as Hesperonia 
californica) are said to be “.. obtuse or acutish, .. most of them narrowed to the apex and never 
rounded.” (Standley 191] 8). Leaves are eusttenee W ith ae pune apices in Torrey’s (1859) original 
plate (4 48), which we | | 


y us. Atthe northernmost known 
locality in the Coast Ranges, in Alameda Co., plants at are siiuch less pu bescent than is common in the 

yeu, pu in may aoe either alana mest rales av Q 929) © or rather pointed leaves 
( clearly | a eee Plants 
Sere spreading + rounded or bluntly acute leaves, lightly glandular-villous pubes 
cence, and rather cymose inflorescences from the interior coast ranges in San Benito and Merced 
counties are very close to var. villosa; flower color is not indicated on specimens (Be ylik 25; aa 
4309; Spellenberg 12336). 


ed 


Inflorescence of var. crassifolia.—In their extremes, the differences between the thyrsoid inflores- 
cence of western races and the neatly forked inflorescence of some eastern populations from the 
desert are notable. From the western edge of the Colorado Desert, where collectors mostly note rose 
per ao rae white - Munz & Everett 16245), plant pen, sprawling, and leafy, the inflores- 
cences thyrse-like as the var. crassifolia. Nearby, on the sandy deen ae are plants more typical of 
var. retrorsa, less = in appearance, with recurved hairs, shorter erect or spreading stems, and an 


inflorescence that is much less thyrse-like, though it is still not neatly forked (V.& A. Grant 15979). 


Fruits of var. crassifolia.—Various authors have indicated certain fruit shapes or surface pattern are 
distinctive for taxa, particularly at the varietal level. Munz & Keck (1968) indicate considerable varia- 
tion in the fruit of a broadly delineated M. laevis var. crassifolia in sense of this paper), indicating 


peed 


NYCTAGINACEAE 561 


the fruits to be “dark, sometimes mottled or pale-striate, smooth,” but provide specific and limited 
characteristics for the fruit of infraspecific taxa of M. bigelovii. Standley (1918) also maintains lim- 
ited variation for the fruits in his taxa within Hesperonia. i the var. cndssvalig nulls may be o 

} ILy a J 


void, broadly ellipsoid, or + globose, grayish brown, and very tl & How 


2396, Philbrick B68-80, Saliorig 2670); irregularly and a pale ce and indefinitely dae 


mott — Robe ooag spate ery ae, mottled faintly with tan (Wiggins 2054); grayish brown mottled 
with re i brown, faintly mottled dave (Youngberg 7); dark 


eee faintly ledd and bee, taney striped, paler near the apex (Munz & Harwood 3900); 
grayish brown and mottled faintly slightly darker, with faint pale — at each end (Havlik 929). 


’ 


Mirabilis laevis var. retrorsa 
Pubescence of var. retrorsa.—Two collections from the Granite Mountains, eastern San Bernardino 
Co., Calif. vmencate tnteieeadation pelween var. retrorsa vans var. vi ae and perhaps the low signifi- 


cance of j general aspect, being small 
and well branched; Stein 12 is g iebrate with a few aoG nward-oriented hairs on the stems (a “good” 

var, retrorsa), whereas Tilforth & Tilforth 1012 is villous. From the area of contact in southern Cali- 
fornia intermediate plants occur; Gould 2248, from the east base of the Coast Ranges in San Diego 
Co., has glandular villosity, some hairs downward directed, and white flowers as in var. villosa, but 
hairs with conical bases and thyrsoid inflorescences similar to var. crassifolia 


F s of var. retrorsa.—F lowers are usually white, but there occasionally are other color forms, 
oe es ane rose throat (Clemon & Jonsson 1690); rose (Twisselman 7280); white and rose-pink at 
lland Chandler 6882, 6884, respectively). 


Inflorescence of var. retrorsa.—The very neatly forked branching characteristic of this variety is 
illustrated by Peirson 8900, Holmgren & Holmgren 7697,and Twisselman 7280. Near the area of contact 
with var. crassifolia, plants may have thyrsoid inf| B 3136, M. E. Jones s.n., 25 Apr 190 


Winblad s.n., 2 Feb 1937); the Jones specimen also has unusually long involucre lobes for the variety. 
Fruits of var. retrorsa.—F h {surface pattern are variable; + globose, gray brown, not lined 
(Reveal & Reveal 50); + globose with 10 pale lines (Henrickson 18257, Henrickson & Bekey 18288); + 
globose, yellowish brown, faintly darker mottled, not lined (Ferris 7988); broadly obovoid, dark, with 
10 thin, pale lines (Munz & Keck dics Peirson 8900), broadly obovoid, smoky brown, not lined 
(Spellenberg et al. 315D; ellipsoid, dark brown and very faintly mottled, incompletely and faintly 10- 
lined pale posed 33848). One ei has plants occasionally with 2 fruits per involucre 
34 


(Spell enberg 12 
E 


Leaves of var. retrorsa.—Leaves are usually obtuse or rounded at the tip. In the zone au intergr adanion 
to the var. crassifol ida in southern California, intermediate plants may have acute leaves (Henri 

5557). A pair of specimens suggest a strong geneti | t to leaf shape md size, nature ‘= pu- 
bescence, and involucre characteristics. Progeny from a collection with white flowers, small, acute 
leaves, and the pubescence of var. retrorsa from the north end of the Coachella Valley (Munz and 
Everett 16245) has ret alaed these features (flower color not given) when grown in the Rancho Santa 
Ana Botanic Garden in Claremont (Ba 
sented by Hen ee 17348. 


ls 19406). A very acute-leaved phase of open habit is repre- 


Mirabilis laevis var. villosa 

Involucre of var. villosa.—Ordinarily, the involucre lobes of the var. villosa are about 1/2-1/3 the 
length of the tube. Plants in southwestern Arizona | unusually long involucre lobes, equal to, or 
even slightly longer than, the tube. These may have resulted from introgression with M. tenuiloba 
(Reeves & Lehto L20124, Harrison LD. Some eee from te pennineals 2: Bae California have the 


] 


dense viscid-villous pubesene e ue va 
of var. crassifolia, and l he tube (e.g., Carter 5449, Thorne et al. 62452). 


562 BRIT.ORG/SIDA 19(3) 


Flower color of var. villosa.—Flowers are usually white in this variety. All three varieties mix in the 
Sierra San Pedro Martir of Baja alors wenere pink- or purple-t EOWEREE villous Spans occur (Daniel 
414, Moran 24540). Transition from pink to I retains pink 
but the perianth limb is white (M. Baker 4544, Palmer 208). On the west side range of var. villosa a 
number of collections document color variation in noes within populakops or eepetiire from tn 


joey 


usually white perianthso whit 
white to lavender (Munz & Hitchcock : 12046), deep iageadey (will ken & Werner 7485), or ear (Cooper 
2257 - a plant intermediate to var. crassifolia in its acute leaves). In southeastern California collec- 
tions by Hall and Chandler note pwns (70 oe) ane pink ey) f Were to occur in Pees same eu 
Pale pink, pink, lavender, or purp east (Graham 3222. ‘ickson 
14004, Lloyd 2866; Train 1377, Wiggins 9648). In Kern Co., Caiianaia. characteristics of varieties are 
variously combined; Twisselmann 8377 and Keck 2158, both from western part of the county are of 
open habit, have fairly large fruits, rather blunt leaves, and very villous pubescence, similar to “good” 


var. villosa, com in its * magenta sicdibete and dies inflorescence it is more similar to the var. 


crassifolia | “rosy- ee perianths and lighter pubescence, 


witha few retrorse hairs, ing, large aa blunt, overall dagen pees var. villosa, but 
1137226) 


the lighter pubescence and s edael eaves characteristic also of va 

Leaf shape of var. villosa.—Plants with acute leaf apices occur along or near line of contact with the 

var. crassifolia in southern California (Peirson 1853; Roos s.n., 26 Mar 1966; Thorne & Tilforth 40843; 

Tilforth & Dourley 340). Sonoran plants often have + acute leaves (Spellenberg 5444; Van Devender 

& Kearns s.n., 18 Feb 1977). More or Ee i ie 7 ants with rounded leaf apices occur as far west as 
)25). 


— 
= 


the Transverse Ranges of Calif 

Pubescence of var. villosa.—In the mountains of southern California plants often are more sparsely 
pubescent in basal parts but are notably villous in upper parts (Peirson 5356). Lightly pubescent 
specimens in southwestern Arizona shee lesetal. oe approach the var. retrorsa, as do plants from 


oo 


southeastern Californ hel bit and large, broad leaves of var. villosa, but with very short 
pubescence, often sparse on loner parts cainane & Lindner c57). 
Inflorescences of var. villosa.—The dichotomous inflorescence cl teristic of the desert races of 


M. laevis from east of the Sierra Nevada and the southern California cepstal ranges is nicely illus- 
trated by Clokey and Anderson 6603. Thyrse-like inflorescences more characteristics of the var. 
crassifolia occur in var. villosa well away from the renee of os iit in eastern Se ue Desert 
(Charlton & Pitzer 1834), or nearer to range of var.cr g 
(Holmgren & Holmgren 7539) or in Baja California Cae 20832). 


Fruits of var. villosa.—Fruits in this variety vary from nearly globose to ellipsoid or obovoid, the 
surface mottled or striped. Example of variation are: fruits + globose, grayish, with 10 very faint and 
indefinite pale lines (Lloyd 2636, Munz 12465, Spellenberg 10206, Turner 62-2), broadly obovoid, in- 
definitely pale-lined at base (Morefield 4800, Spellenberg 2982), + globose, 10 faint, pale lines alter- 
nating with 10 diffuse darker lines (Parish 3183, Munz 10930); + globose, very dark and dark-mottled, 
without lines J. & L. Roos 4182); ellipsoid or obovoid, ce or brown and black- or dark-mottled (Boyd 
et al. 2112, Felger & Valenzuela L. 86-180, Higgins 6378, Reeves & Lehto L20124, Roos s.n, Spellenberg 
10205), broadly obovoid, unlined, grayish brown (Jepson oD) or faintly lined Gepson 5957, same 


ae) 


time and place). 

APPENDIX 2 
Representative and/or cited specimens. Specimens are cited by taxon, and 
within taxa geographically by country, state, and county, then alphabetically 
by collector. Those specimens that provided fruit for illustration in Figure | are 
indicated by and asterisk (*) following herbarium citation. 


NYCTAGINACEAE, 563 


Mirabilis laevis var. crassifolia 
MEXICO. BAJA CALIFORNIA: 21.9 mi E of El Rosario via Hwy. 1,13 Oct 1981, Burgess et al.6095 (ARIZ, 
SD);0.9 mi N of Rosario, 6 Feb 1953, Chisaki & Newcomb 525 (ARIZ, GH, RM, SD, UC); San Matias Pass of 
Sierra San Pedro Martir, 20 May 1981, Daniel 1345 (ASU); Cedros Id.,ca.2 mi S of lighthouse on E side, 
23 Feb 1977, Davidson 5488 (RSA); 5 mi N of San Quintin, 15 Feb 1935, Epling & Robinson s.n. (ARIZ, 
GH, NY, RM, UC); Mina Desengana,ca..16 mi N of Punta Prieta, 30 Mar 1950, Gentry 8886 (ARIZ); Cedros 
Id.,ca.1 mi S of village at Cabo Norte, 19 Jan 1975, Henrickson 14453 (NMC); 1 km N of San Vicente, 6 
Jan 1984, Hodgson & Pinkava 3011 (ASU); South Todos Santos Id., 7 Apr 1948, Moran 2802 (UC); Sierra 
San Borja, Rancho Carrizo, 20 Mar 1966, Moran 12832 (SD);San Esteban Id., NE peak, 28 42'N, 112°35'W, 
26 Apr 1966, Moran 13051 (SD);San Martin Island, 21 Apr 1970, Moran 17458 (RSA); 7 mi SE of Laguna 
Chapala, 18 Oct 1971, Moran 18694 (ARIZ, RSA, SD); Guadalupe Island, south end of island, 30 Mar 
1889, Palmer 886 (US*); ca. 23 km NW of parador Catavina [Santa IAez], 15 Jun 1980, Reeder & Reeder 
7259 (SD); San Martin Island, 3 mi off cinder cone of San Quintin, 21 Feb 1986, Thorne 61594 (RSA); 
Guadalupe Island, NE Anchorage, 28-29 Mar 1988, Thorne 63015. BAJA CALIFORNIA SUR: NW end 
of Viscaino Peninsula on road from Bahia Tortugas to Punta Eugenia, 2 May 1993, Boyd, Ross & Appleby 
8100 (TEX); 26 km N of San Ignacio, 10 Jan 1948, Carter, Alexander & Kellogg 2522 (ARIZ, UC);8 mi N of 
San Juanico, 8 Mar 1939, Gentry 4314 (ARIZ, GH); E bajada of Sierra Calvario, 10-15 Mar 1947, Gentry 
7391 (ARIZ, RSA, UC); Picachos de Santa Clara, 5-10 Nov 1974, Gentry 7697 (SD); 2-3 mi E of Punta 
Eugenia, 13 Mar 1949, Gentry 8694 (ARIZ); between Volcan Tres Virgenes and Cerro Azufre, 27 29’N, 
112 34'W , 11 Apr 1973, Henrickson 8940 (SD); Rancho la Laguna, Sierra San Francisco, 27 35'N, 
113 02'W , 23 Nov 1976, Moran 23821 (SD); 6 mi N of San Andrés, Arroyo Calvario, 10 Feb 1973 
Moran & Reveal 20006 (SD); Picachos de Santa Clara, 3 Feb 1973, Moran & Reveal 19689 (SD), 19671 
(ASU, POM; SD); Cerro Azufre, 27 30'N, 112 36'W ,20 Oct 1971,Moran 18723 (SD;UC);Volcan las Tres 
Virgenes, 27°29’N, 112°36'W, 11 Apr 1973, Moran 20414 (SD), 20415 (SD, UC); Arroyo Malarrimo 11m 
S of mouth, 6 Feb 1973, Moran cee 19870 (ASU, SD, UC). U.S.A. CALIFORNIA. Alameda Co.: - 
facing slope of Mission Peak, 1 n 1980, Havlik 929 (CAS); E side of Fremont on ease Peak, 16 
Jun 1996, Spellenberg 12335 (F, Ae NMC*, NY, RSA, UC, US). Fresno Co.: Owens Mtn., 6 Apr 1951, 
Boolootian s.n. (JEPS); Owens Mt.,6 mi SE of Friant Dam, 9 May 1953, ee 20 (RSA). Kern Co.: 
Greenhorn Mts., Mt. Breckenridge, 3 Apr 1932, Benson 3214 (UC); Oildale - Woody Road, 17 Apr 1938, 
Hoover 3170 (DS, UC); entrance to Kern River Canyon, 7 Jul 1962, Howell 38142 (CAS); Wofford, 7 Jul 
1962, Howell 38179 (CAS); Kern River Canyon, 21 Sep 1962, Howell 38675 (CAS); Caliente, 15 Apr 1916, 
Jepson 6752 (JEPS); 2.6 mi E of Caliente, 16 May 1963, Thorne 31702 (RSA); Temblor Range, Cedar 
nyon, 1 Jun 1952, Twisselmann 198 (CAS); mouth of Kern Canyon, 11 Jun 1963, Twisselman 8391 
(CAS, RSA); 2 mi NE of Weldon, 5 May 1933, Voegelin 67 (UC). Los Angeles Co.: San Clemente Island, 
10 Jun 1962, Blakley 5238 (SD);San Clemente Is.,25 Aug 1894, Brandegee s.n. (UC); Santa Monica Mts., 
Las Flores Canyon, 28 Mar 1930, Epling & Ellison s.n. (MO, RSA, UC); Los Angeles, 16 Apr 1904, Grant 
791 (ARIZ, CAS, DS, RSA); Los Angeles, May 1885, Gray s.n. (GH); Pacific Palisades, Temescal Canyon, 16 
Apr 1941 Hastings s.n. (NY); Santa Catalina Id.,S of Wilson's Harbor, 2 Mar 1941, Moran 669 (RSA); San 
Clemente Id.,2 mi S of Eel Point, 18 Sep 1958, Moran 6848 (DS, RSA, UC) [same site, 9 Mar 1959, Moran 
7170 (DS, RSA)]; San Clemente Id., £ coast, 9 Apr 1923, Munz 6645 (POM, UC); E of Zuma Beach, 4 Apr 
1959, Raven 13964 (RSA); San Clemente Id., S of Eel Point, 11 Apr 1962, Raven 17307 (RSA, SD); San 
Clemente Id., just N of Guds, 9 May 1962, Raven 17655 (RSA, SD); near isthmus on Santa Catalina Id., 
Templeton 11388,25 Feb 1968 (RSA); Santa Barbara Id., Cat Canyon, 28 Apr 1968, Thorne 37483 (RSA, 
SD); San Gabriel Mts., San Dimas Canyon, 9 Apr 1971, Thorne & Tilforth 41536 (RSA);San Clemente Id., 
May 1903, Trask sn (A).Merced Co.: Mine Canyon near Little Panoche Valley,6 Apr 1940, Hoover 4309 
(DS); Mine Creek 1.5 mi N of junction of Merced, Fresno, San Benito cos.,11 Apr 1935, Lyon 932 (UC). 
Monterrey Co.: 6 mi From King City, 10 May 1936, Eastwood & Howell 2396 (CAS, NY, UC); Redwood 
Gulch, 20 May eee 5795 (RSA); 6 mi N of King City, San Lorenzo Creek, 7 May 1963, Howell 
39241 (RSA);6 mi NE King City, 7 May 1963, J.T. Howell 39241 (RSA); 6 mi NE of King City, 7 May 1963, 
Rose 63030 (CAS, DS, RSA). Riverside Co.: Whitewater Canyon about 3 mi from mouth, 8 Apr 1932 


564 BRIT.ORG/SIDA 19(3) 


Ewan 7041 (POM); San Gorgonio Pass, 25 May 1914, Jepson 6073 fare nary icin near 
Whitewater, 11 Apr 1948, Kamb 902 VJEPS); Dry Morongo Wash, 2 May 1952, Munz & Everett 16245 
(RSA). San Benito Co.:ca.6 mi SE of Panoche School, 12 May 1958, Beylick 25 (RSA); Cherry fl es 
W of Llanada, 25 Apr 1937, Fwan 10309 (RSA); 17.6 mi from New Idria on road to Panoche, 6 May 
1956, Raven et al. 9240 (RSA); Road 107, 14.5 km SE of junction with Little Panoche Rd, 27 km NW of 
New Idria, 17 Jun 1996, Spellenberg 12336 (NMC*, NY, UC). San Bernardino Co.: Dry Morongo Creek, 
6 Apr 1933, Dunkle 3411 (RSA); Santa Ana River Canyon, 3 May 1919, Munz, Street & Williams 2672 
(POM); Hwy. 330 ca. 3 mi E of Highlands, 27 May 1990, Spellenberg 10208 (NMC, NY, UC); Collius Valley, 
aed Canyon, 28 Apr 1920, Jepson 8859 (JEPS); Fallbrook, 15 May 1920, Munz & Harwood 3900 (RSA); 
n Diego, Chollas Valley, 1 Jan 1884, Orcutt 219a (MO). San Luis Obispo Co.: summit of Cotton 
ee Pass, 1 May 1949, Hoover 7644 (CAS); Escondido, 5 Jun 1904, eae ames vata (cited 
in new combination of M. tenuiloba var. polyphylla); 3 mi S San Clemente, 19 Mar 1966, Wallace 176 
(SD); Otay Lake, 12 Apr 1981, Werff 4221 (SD); Cuyamaca Mts.,6 mi below Alpine, 20 Mar 1926, Wig- 
gins 2054 (SD); Camp Kearney Mesa, 7 Apr 1935, Youngberg 7 (POM). Santa Barbara Co.: Santa Bar- 
bara Island, Cat Canyon, 4 May 1963, Blakley 5657 (US*); toward Figueroa Mtn., 4 Apr 1986, Bourell, 
Patterson & Timbrook 2938 (CAS); Santa Cruz Id., 17 May 1962, Breedlove 2874 (DS); Santa Cruz Id., 9 
un 1930, Clokey 4923 (NY, RSA, UC); Santa Cruz Id., 16-17 Jul 1917, Eastwood 6387 (CAS); Santa Bar- 
ue Id.,Cat Canyon, 19 Mar 1968, Philbrick B68-80 (RSA); W of Goleta, 20 Sep 1956, Pollard s.n. (CAS). 
a Co.: 5 mi S of Filmore, 2 Apr 1958, Solbrig 2670 (NY); 2 mi E of Point Mugu, 14 Mar 1959, 
aes 1857 (CAS). 


ae 


wa 


oS 


Mirabilis laevis var. laevis 

MEXICO. BAJA CALIFORNIA SUR: Magdalena Bay, 18 Jan 1889, 1.S. Brandegee s.n. (GH*):E base San 
Lazaro Mt., Santa Maria Bay, 30 Mar 1952, Moran 3530, SD; Magdalena Bay, without date, Dr. Sung 28, 
UC (#101225, mounted on sheet with M. laevis var. crassifolia). 


Mirabilis laevis var. retrors 

MEXICO, BAJA CALIF annie Sierra Juarez, Arroyo el Toruno, 17 Mar 1968, Moran 14842 (ASU, RSA); 
Canon de Guadalupe, 32 09’N 115 48'W___, 23 Mar 1986, Thorne, Boyd, et al.61758 (RSA); San Matias 
Pass, 6.2 mi E of Ejido San Matias, 20 Apr 1985, Thorne and Charlton 60220 (RSA). U.S.A. ARIZONA. 
Mohave Co.: road from Chloride to the river, 13 May 1931, Fastwood 18313 (CAS*). CALIFORNIA. 
Inyo Co.: Panamint Range, Emigrant Springs, 6 Apr 1935, Clokey & Templeton 5725 (POM, NY, UC); 
White Mts., Silver Canyon, 1 Jun 1933, Duran 3455 (CAS, POM, RSA); Panamint Mts., Surprise Canyon, 
13 Jun 1930, Ferris 7988A (DUD, UC); ca. 25 air mi S of Olancha at Little Lake, 8 Jun 1979, Henrickson 
18257 (NMC); ca. 25 air mi SSE of Olancha, 12 Jun 1979, Henrickson & Bekey 18288 (NMC, NY); Death 
Valley, S end, Bradbury Well, 9 Apr 1940, Munz 16449 (POM, UC); Eureka Valley along Big Pine road, 13 
May 1962, Reveal & Reveal 50 (NY); ca. 25 air mi S of Olancha at ae Lake, 8 Jun 1979, Henrickson 
18257 (NMC); Darwin Falls Canyon, 30 Apr 1937, Train s.n. (DUD - 258204); Death Valley Natl.Mon.,25 
Mar 1947, Wiggins 11529 (DUD, UC). Kern Co.: Red Rock aoe May 1927, Abrams 11877 (POM); 
butte S of Mojave, 25 Mar 1932, Benson 3136 (POM); 2 mi E of Weldon, 12 May 1933, Cole & Voegelin 
120 (UC); Mojave, 12 May 1913, Eastwood 3200 (POM); Mojave - Randsburg region, 0.5 mi W of Big 
Bend, 1 Jun 1962, Twisselman, 7280 (CAS); near Searls P.O., 8 May 1906, Hall and Chandler 6882, 6884 
(UC); California City land develo satel land, 10 Apr 1974, Holmgren & Holmgren 7697 (NMC, RSA); 
Kernville, 20 May 1962, Howell 37115 (CAS); NE of Lake Isabella, 12 Jul 1962, Howell 38667 (CAS); Red 
Rock Canyon, 13 May 1930, ne 8900, (P OM, RSA); Sierra Way on N side of Lake Isabella, 5 km E of 
junction with Calif. Hwy. 178 at Bella Vista, 19 Jun 1996, Spellenberg 12342 (NMC*, NY); Cache Creek, 
ca.0.5 mi W of Big Bend, 1 Jun 1962, Twisselman 7280 (CAS).Los Angeles Co.: Palmdale, May 1925,M 
& E. Epling sn, (MOQ); Lovejoy Buttes, 17 Apr 1932, Peirson 9846 (RSA). Riverside Co.: cultivated from 
Munz & Everett 16245, 26 May 1954, Balls 19406 (RSA); Morongo Valley road ca. 1 mi N of Hwy 60, 7 
Apr 1951, V.&A. Grant 15979 (RSA); Indio, 26 Apr 1906, M. E. Jones s.n. (POM); Coachella Valley, 2 Feb 


SPELLENBERG AND RODRIGUEZ, NYCTAGINACEAE, at 


1937, Winblad s.n. (CAS); N base of Eagle Mts., 12 Apr 1949, Munz 13036 (RSA); Dry Morongo Wash, 
NW end of Coachella Valley, 2 May 1952, Munz & Everett 16245 (RSA) (seed source for Balls 19406); 
Shavers Well near Mecca, 9 Apr 1922, Munz & Keck 4754 (POM, UC); ca.4 mi S of Morongo Valley, 3 
May 1964, Thorne 33848 (DUD, RSA); Coachella Valley, 2 Feb 1937, Winblad s.n. (CAS)., San Bernar- 
dino Co.: Kramer Junction, 4 Jun 1987, Bagley 2098 (RSA); S side Ord Mtn.,8 Apr 1988, Boyd et al. 1726 
(RSA); 19 mi E of Banning, 2.3 mi NW of Hwy 62 on rd to Big Morongo Canyon, 29 May 1971, Henrickson 
5557 (MO); 30 mi NNW of Barstow,N of Black Canyon, 28 May 1978, Henrickson 17348 (RSA); Newberry 
Mts., 7 Apr 1924, Munz & Keck 7862 (POM); S of Death Valley Natl. Mon., Avawatz Mts., 18 May 1973, 
eee et al. 3151 (NMC); Granite Mts., 0.5 mi SW of Willow Spring Basin, 14 Apr 1978, Stein 12 
(R mi W of Barstow, Iron Mtn., 21 Oct 1976, Strother 1256 (UC - chromosome count by 
ae renoned on specimen). San Diego Co.: McCain Valley, 15 Apr 1987, Clemons & Jonsson 

1690 (SD); 6 mi E of Banner on Hwy 78,5 Apr 1944, Gould 2248 (UC); E end Santo Rosa Mts., 14 Apr 

1927, Peirson 7180 (RSA). NEVADA. Clark Co.: Las ets Ranch, 10 May 1905, Goodding 2347 
a Sons ee S a Spring Mt. Range, 10 Jun 1938 Train 1932 (NY). Esmeralda Co.: base of 
tezuma Mts.W of Goldfield, 4 Jun 1919, caeien 9755 (RM). Humboldt Co.: Bilk Creek Mts., SW 
side of Black Mts, Bo R33E, 22 May 1987 Tiehm 11048 (CAS). Lincoln Co.: Pahranagat Valley, rd 
from Crystal Springs to Ash Springs, opposite Geer Ranch, 30 Aug 1938, Train 2421 (A). Lyon Co 
East Walker River Rd.,ca.55 air km SE of Yerington, 13 Jun 1996, Spellenberg 12327 (MO, NMC, NY, UC); 
East Walker River Rd.ca.45 air km SE of Yerington, 13 Jun 1996, Spellenberg 12329 (NMC*, NY); 3 km 
W of Wellington on Nev. Hwy. 208 1 km E of Douglas Co. line, 13 Jun 1996, Spellenberg 12333 (MO, 
NMC, NY). OREGON. Harney Co.: 14 mi S on Toole Spring Rd., E of Alvord Lake, T36S R34E, 6 Jun 
1964, Holmgren & Reveal 870 (GH, NY); Pueblo Valley, 8.5 air mi NE of Fields, T37S R34E, 6 Jun 1964, 
Homgren & Reveal 870 [sic] (NY). Malheur Co.: Owyhee River canyon, 9 mi upstream from Adrian, 25 
May 1989, Barneby 18303 (NY*); Owyhee River, T22S R45E $3, 17 Jun 1976, Packard 76-107 (NY); 
Owyhee Canyon, 13 mi below dam, 15 Jun 1942, Peck 21227 (NY). UTAH. Washington Co.: Virgin 
River 12 mi i bal St. George, 18 May 1965, Cronquist 10110 (NY, RSA); St. George, Black Hill, 16 Apr 
1942, Gould 1567 (GH); near Ft. Pierce, 14 May 1986, Higgins 16468 (NY); 5 mi E Washington, Rock 
Cliffs, 20 May 1 i Maguire & Blood 4390 (GH, POM). 


——_ 


S SS 


me 


Mirabilis laevis var. villosa 
MEXICO. BAJA CALIFORNIA: Sierra San Pedro Martir, between Hwy. 3 and Rancho Mike, 20 May 
1981, Daniel 1414 (ASU); San Borja, 28 47'N 113 57'W__, 20 Apr 1946, Moran 1997 (UC);3 mi N of El 
Alamo, 31 38'N, 11601.5'W ,30 May 1970, Moran 17644 (RSA, SD); + 25 km of Tecate, Kumeyaay 
rancho of Ha-a,32 22'N, 116 30’W ,10 Oct 1976, Moran 23774 (SD); Sierra San Pedro Martir, 1 km NE 
of El Socorro, 30 58.5'N, 115 38.5'W ,20 Aug 1977, Moran 24540 (SD); Sierra Juarez, on road to Ojos 
Negros, 18.3 mi SW of Laguna Hanson, 30 May 1983, Thorne et al 55990 (RSA); Cafion de Guadalupe, 
32 09'N 115 47.5'W 18-20 Feb 1984, Thorne et al. 57784 (RSA); W foothills of the Sierra de Judrez, 
near El Bashisha, 26 & 27 May 1987, Thorne et al.62452 (RSA), along road to San Matias Pass and Valle 
Trinidad, San Felipe Desert, 11 Nov 1967, Wiggins 20832 (SD); E of Ensenada, 2 mi W of Coyote along 
rd uae Ojos Negros and Laguna Hanson, 5 May 1969, Wilken & Werner 7485 (UC). BAJA CALI- 
FORNIA SUR: Cerro del Pinto, N of Portezuelo de San Antonio, 24 50.5'N, 110 44’'W ,21 Feb 1970, 
coe on (NMC). SONORA: Ouitobaquito, 19 km W of Sonoita, 10 Apr 1986, Felger & Valenzuela L. 
-180 (ARIZ); ca. 5 mi S of Puerto Libertad, 1 Apr 1980, Spellenberg 5444 (ASU, CAS, ENCB, K, MEXU, 
es NY, RSA, UC, UNM, WTC); Punto Cirio, ca. 7 mi S of Puerto Libertad, 27 Apr 1962, Turner 62-2 
pe Sct Bacha, Punto Cirio near Libertad, 18 Feb 1977, Van eile Kearns s nn. (ARIZ); 19 mi 
NW of San Ignacio, 26 Feb 1979, Walker fle (ARIZ).U.S.A.-- ARIZONA. 0 Co.: Red Lake, 
17 May 1969, Cazier s.n. (ASU). Graham Co.: Camp Grant, 2 nae 867, sat ae (MO). Maricopa 
Co.: 33 32'30"W,111 27'N,9 Apr 1983, a Baker 4544 (ASU); Hassayampa Plain, Coyote Wash, 14 Mar 
1979, Fischer 5969 (ARIZ, ASU); Sierra Estrella Regional Park, 31 Mar 1968, Pinkava 4739 (ASU, NMC); 
Sacaton Mts., 14 Oct 1925, Peebles, Harrison & Kearney 463 (ARIZ). Pima Co.: Organ Pipe Natl. Mon., 


566 BRIT.ORG/SIDA 19(3) 


Ajo Mt., 11 Mar 1983, Daniel 2586 (ASU, RSA); Sabino Canyon, Santa Catalina Mts.,2 Apr 1928,Graham 
3222 (DS); Tucson Mts., Picture Rocks Pass, 17 Apr 1977, Van Devender et al.sn (ARIZ); ca.6 mi NW of 
Sells, 10 Apr 1973, Spell enveig 298? (NMC*, NY): Pinal Co.: SE Sierra Estrella, 23 Feb 1983, Rea 290 
(ARIZ, SD). Yuma Co.: W side Plomosa Mts.,27 Mar 1981 Butterwick & Hillyard 7043 (ASU, CAS); SE side 
of Tinajas Altas Mts., Borrego Canyon, 16 Jun 1992, Felger & Broyles 92-614 (ARIZ); Palm Canyon,8 Oct 
977, Harrison 11 (ASU); Tule Tank, 23 Mar 1935, Kearney & Peebles 10890 (ARIZ, GH); Kofa Mts., Palm 
Canyon, 19 May 1976, Reeves & Lehto L20124 (ASU). CALIFORNIA. Fresno Co.: Alcalde Canyon, 12 
Jun 1938, Fastwood & Howell 5839 (CAS); Coalinga-San Lucas Rd. 2.5 mi W of Coalinga, Alcalde Can- 
yon, 28 May 1941, Ferris & Bacigalupi 10350 (UC). Imperial Co.: ca. 20 mi NW of Winterhaven, 26 Mar 
1973, Higgins 6378 (NMC, NY); Jacumba Mts., 29 Mar 1974, Holmgren & Holmgren 7539 (NMC, NY); 
Chocolate Mts.,8 Apr 1949, J.& L. Roos 4182 (RSA); E side Chocolate Mts., 11.3 mi NW of Beal Well, 25 
Mar 1941, Wiggins 9648 (GH, UC). Inyo Co.: Panamint Canyon, 15 May 1906, Hall & Chandler 7023, 
7024, (UC); Westgard Pass, 18 Jun 1963, Lloyd 2636 (NY, UC); White Mts., Silver Canyon, 27 Jun 196 
Lloyd 2866 (NY);White Mts., Silver Canyon, 7 Jun 1961, Mooney, Andre & oe 132 (DS); White Mts., 
Cottonwood Creek, 18 Jul 1988, Morefield 4800 (NY); Death Valley Natl.Mon., 1 mi SE White Top Mtn., 
3 Jun 1982, Peterson 566 (RSA); Darwin Falls Canyon, 30 Apr 1937, Train s.n., (DS-258167). Kern Co.: 
Cedar Canyon, Bedell 74-5,11 Jul 1962 (CAS); E side of Walker Pass, 21 May 1962, Howell 37226 (CAS); 
McClure Valley (near Kings Co.line),4 May 1933, Keck 2158 (DS);9 mi N of Ricardo, 5 May 1932, Munz 
12465 (UC); Temblor Range, Ross Ridge, 4 Jun 1963, Twisselman 8377 (CAS, RSA). Los Angeles - : 
mi S of Gorman, 21 Jun 1978, Gustafson hen (RSA); San Gabriel Mts., Arraster Creek, 10 May 19 
ee 1853 (RSA); Mint Canyon, 16 Jun 1918, Peirson 5356 (RSA); San Gabriel Mts., Little soe 
round, 30 Jun 1971, Thorne & Ti A oe 3 (RSA).Mono Co.: ca. 3 mi W of Benton Station, 31 
May os imate aed Riverside Co.: Ne a Palen Mts.,10 Mar 1988, Boyd et al. 2112 
(RSA); Banning, 6 May 1945, Cooper 2257 (RSA); N base of Eagle Mts., 12 Apr 1949, Munz 13037 (RSA); 
2 mi SE Desert Center, : Mar 1966, Roos s.n. (RSA). San Bernardino Co.: Providence Mtns, 9 mi E of 
Mitchell Caverns Rec. Area, 7 May 1988, canes & Pitzer 1834 (RSA); di Sere 5 Jun 1935, 
Clokey & Anderson 6603 (NY, POM, RSA, UC); N Kingston Mts. 2 mi SW of Tecopa Pass, 13 May 1974, 
Alaa 14004 (RSA); Shay’'s Well, Mojave Desert, 14 May 1941, Jepson 5957, aie rae Cushenberry 
1927, Munz 10930 (DS, POM); San Bernardino Mts. and E base, 16 Jun 1894, Parish 
3183 (NY); ce 1812 mi SWof Victorville, 27 May 1990, sai 10205 (NMC,NY, UC); near Hesperia, 
27 May 1990, Spellenberg 10206 (NMC*, NY, RSA, TEX, UC); Morongo Valley, 14 Apr 1971, Tilforth & 
Dourley 340 (ASU, RSA); Granite Mts., Snake Spring area, 18 May 1975, Tilforth & Tilforth 1012 (RSA). 
San Diego Co.: Vallecito Wash 30.5 airline mi NW of Ocotillo, 26 Mar 1973, ety & Holmgren 
6535 (NY); walls of Box Canyon, W. Colorado Desert, 2 Apr 1932, Munz & Hitch 6 (MO, UC). 
San Luis Obispo Co.: just E of summit of Cottonwood Pass on St. Hwy. 41 24 hay ae che upl 
et al. 5205 (DS). NEVADA. Clark Co.: Sheep Mts., Hidden Forest, Deadman’s Canyon, 30 Jun 194 
Alexander & Kellogg 1811 (GH);43 air mi S of Mesquite, mts.S of Virgin Mts., Hell's Kitchen, 8 ‘ 1975, 
Holmgren & Holmgren 7926 (NY); Newberry Mts., Hiko Spring, 11 Apr 1938, Train 1377 (NY). Lyon Co.: 
East Walker Road E., ca. 24 air km SE of Yerington, 13 Jun 1996, Spellenberg 12331 (NMC); Nev. Hwy. 
208 13 km ENE of Smith at E entrance to Wilson Canyon along West Walker River, 13 Jun 1996, 
sei pape. ie Me NMC*, NY). Nye Co.: 10 mi SW of Beatty above Buck Springs, 27 May 1945, 
25913 (GH,NY). UTAH. Washington Co.: Bulldog Knolls, T43S R18W $28, 30 Apr 
1986, \f/oleha eg Rrird 23 06 (NY). 


TEED. sae ntha 

BAJA CALIFORNIA: Catavina arroyo ca.5 km N of Santa Ynez, 6 Jun 1974, Carter & Demp- 
ster pret McC); Catavina, 23 Mar 1932, Harvey 5017 (US); Catavina Mesa, 22 Apr 1952, Gentry & Fox 
11731 (LL*);Catavinacito, 29 44’N, 114 45'W_ 21 Nov 1976, Moran 23808 (SD*); 1 mi S of Las Arrastras, 
25 Mar 1960, Wiggins & Wiggins 15940 (ARIZ, TEX*); E of El Marmol on trail to Gulf, 14 Feb 1935, Shreve 
6845 (ARIZ); 10. 7 mi (by road) N of Bahia San Luis Gonzaga, 6 Oct 1967, Hastings & Turner 67-10 (ARIZ, 


— 


NYCTAGINACEAE 567 


SD); Sierra de Volcan 4 mi E of El Marmol, 13 Feb 1935, Wiggins 7571 (UC); Rancho Catavina, 35 mi S 
of El Marmol, 8 Mar 1930, Wiggins 4406 (UC); San Francisquito Wash, 18.7 mi (by Road) SW of Bahia 
San Luis Gonzaga, 12 Oct 1963, Hasting & Turner 63-158 (ARIZ). BAJA CALIFORNIA SUR: Picachos 
de Santa Clara, 5-10 Nov 1947, Gentry 7717 (ARIZ). 


Mirabilis oxybaphoides 

MEXICO. CHIHUAHUA: Ca. 23 air mi ENE of Villa Anumada,12 Sep 1973, Henrickson 12849 (NMC). 
COAHUILA: Arteaga, C.Los Camargos, 4 Aug 1980, Hinton et al. 17926 (CIIDIR); Sierra del Pino, west- 
ern ridge, W of camp at La Noria, 24 Aug 1940, Johnston & Muller 603 (GH*, LL, TEX);Canion de Calabasa, 
N wall of Sierra Mojada,27 Oct 1941, Stewart 2209 (LL,GH). NUEVO LEON: Hacienda Pablillo, Galeana, 
5 Aug 1936, Taylor 93 (TEX); Dist. Zaragoza, Puerto Pino, 19 Jun 1979, Hinton 17556 (TEX); Dist. Arteaga, 
Canyon de Los Amargos, 4 Aug 1980, Hinton 17926 (CIIDIR, TEX); Mcpio. Galeana, W slope Potosi, 29 
Jun 1983, Hinton et al. 18491 (GH, TEX). U.S.A. ARIZONA. Apache Co.: Canyon de Chelly Nat'l. Mon., 
2 Sep 1972, Halse 811 (ARIZ);ca.1 mi N of Nelson Reservoir (SE Eager), 20 Sep 1975, Lehto et al. 19057 
(ASU, NMC); Little Colorado River, 0.15 mi S of jct AZ Hwy. 260 along AZ Hwy 273, around bridge 
over river, 30 Aug 1988, Ricketson & Raechal 4364 (ASC, MO, NY, RSA, TEX, UC). Cochise Co.: Cedar 
Gulch, Paradise, 30 Sep 1907, Blumer 2241 (GH); Dragoon Mts.,N side of Mt.Glenn,9 Sep 1983, Daniel 
3148 (ASU). Coconino Co.: Colton Ranch, field 1,24 Aug 1957, McDougal s.n. (ARIZ); Sycamore Can- 
yon Wilderness Area, 11 Oct 1969, Pinkava et al. 5855 (ASU); 6 mi S of |-40 at Meteoh Crater, 3 na 

1981,R.& D. Sauleda 6438 (ASU).Navajo Co.: Chuska Mts., E of Fort Defi ridge W of Coal Min 
Wash,12 Sep 1977, Spellenberg 4893 (NMC). Yavapai Co.: Mingus Mountain, 3 Sep 1968, panes 
(ASU). COLORADO. Chaffee Co.: near junction of highways 285 and 291 NW of Salida,20 Aug 1954, 
Waterfall 12142 (TEX). El Paso Co.: W side of Colorado Springs at entrance to Queen's Canyon, $27 
T13S R65W, 6 Jul 1996, Kelso & Maentz 96-32 (NMC); W of entrance to Garden of the Gods,14 Aug 
1954, Waterfall 12040 (TEX). Montezuma Co.: lower Spruce Canyon near jct. with Navajo Canyon, 
17 Sep 1947, Weber 3629 (ARIZ, NMC, TEX). Montrose Co.: Dolores River Canyon, 7.1 mi S of Sinbad 
Valley Rd., T49N, R18W, S34, 29 Aug 1985, Franklin 2489 (GH). Ouray Co.: W of Ridgeway, 20 Aug 1920, 
Payson 2308 (GH).San Miguel Co.: Norwood Hill, 20 Aug 1912, Walker 508 (GH). NEVADA. Clark Co.: 
Charleston Mts., Little Falls, 3 Aug 1935, Clokey 5454 (GH). NEW MEXICO. Catron Co.: 14 mi SW of 
Horse Springs, vic. Bat Cave, 19 Aug 1948, Smith 208 (ARIZ, GH); Forest Rte. 95, Whitewater, 24 Sep 
1972, lays GT-18 (ARIZ). Cibola Co.: Ramah Navajo, 22 Aug 1939, Vestal & Vestal 56 (ECON), 10 Sep 
1939, Vestal & Vestal 408 (ECON). Colfax Co.: Philmont Scout Ranch, near Cimarron, South Ponil Can- 
yon, 1 mi E of Pueblano Camp, 5 Aug 1968, Hartman 2605 (TEX). Dofia Ana Co.: S end Organ Mts., 24 
Oct 1971, Spellenberg 2738 (NMC); Organ Mts.,4 Sep 1897, Wooton 587 (ARIZ).Grant Co.: Bear Moun- 
tain, near Silver City, 15 Sep 1903, Metcalfe 696 (ARIZ,GH, NMC); ca. 5 air mi NW of Silver City on Bear 
Mountain, 6 Sep 1980, Spellenberg & Soreng 5858 (NMC, NY) (chromosome count by D. Ward). Lin- 
coln Co.: near Gray, Aug 1898, Skehan 103 (GH); Oscura Peak, 26 Aug 1988, Spellenberg et al. 9681 
(MO, NMC, NY). Luna Co.: summit of Cooke's Peak, 20 Sep 1986, Columbus 637 (NMC*). Otero Co.: 
ca. 5 air mi ENE of Alamogordo at High Rolls, 16 Sep 1988, Spellenberg & Ward 9728 (NMC, NY). 
Sandoval Co.: Sky Village, S22 T14 R1W, Oct 1974, Blankenhorn 214 (ARIZ). San Miguel Co.: 18 mi E 
of Las Vegas, M. E. O'Connor Trust Ranch, 2.8 mi S of Rte. 104, Mogote Trap, near gate to Crystal 
Pasture, 21 Aug 1982, Hill & Levandoski 12161 (GH); near Pecos, 18 Aug 1908, Standley 5063 (NMC). 
nta Fe Co.: north of Glorieta, 24 Aug 1908, Standley 5255 (NMC). Sierra Co.: Kingston, 5 Oct 19 

Metcalfe 1459 (GH, NMC); W face of Caballo Mts. 8.6 mi by winding road E of Caballo Dam on Rio 
Grande, 8 Sep 1974, Spellenberg 3936 (LL, NMC). Taos Co.: between Amalia and Ute Springs, 14 Aug 
1973, Holmgren & Holmgren 7169 (ASU, NMC). TEXAS. Brewster Co.: Chisos Mts.,23 Aug 1931, Muel- 
ler 7994 (GH); Big Bend National Park,Lost Mine Peak, 12 Sep 1961, Correll & Johnston 24516 (GH*, LL). 
Chaffee Co.: near junction of hignways 285 and 291,20 Aug 1954, Waterfall 12142 (TEX*). Culberson 
Co.: Guadalupe Mts. Nat'l. Pk., Shumard Canyon, 29 Sep 1973, Burgess 1631 (ASU); Guadalupe Mts. 
Nat'l Park, Bear Canyon trail to The Bowl, 16 Sep 1974, Burgess 2657 (ARIZ). Presidio Co.: Sierra Tierra 


568 BRIT.ORG/SIDA 19(3) 


Vieja,ca. 1/2 mi S of Vieja, 4 Oct 1941, Hinckley 2184 (ARIZ, GH).UTAH. Wayne Co.: Aquarius Plateau, 
Utah 117 _ mi S of Grover, T30S, R5E, $2, 19 Aug 1965, Holmgren et al. 2548 (TEX) 


plea tenuiloba 
EXICO. BAJA CALIFORNIA: along trail from Guadalupe Cyn to Laguna Hanson, 32 10'N, 
15 : W ,13 Mar 1988, Clemons & Jonsson 1999 (SD); canyon 3 mi from Bahia de Los Angeles vil- 
toward San Borja, 17 Feb 1963, Cowan 2321 (CAS, GH, SD); Cocopa Mts., 22 Apr 1949, Gentry 
8712 (ARIZ, RSA, SD); first large canon W of Punta Diablo, 25 Mar 1959, Moran 7251 (DS, SD); 29 mi N 
of San Luis Gonzaga, 30 08'N, 114 40’W ,20 Apr 1960, Moran 8211 (SD);S end of North San Lorenzo 
Island, 24 Mar 1962, Moran 8877 (CAS, SD); ca.6 km SE of Puerto Refugio, 17 Mar 1977, Moran 23949 
(SD);Los Angeles Bay, Dec 1887, Palmer 600 (GH, UC) (paratype of Hesperonia polyphylla);41.6 mi S of 
Mexicali, 22 Mar 1970, Powell & Turner 1708 (TEX, US*); Puerto Refugio, Punta Norte de la Isla Angel 
de la Guarda, 7 Feb 1986, Tenorio L.& Romero de T. 10836 (RSA, TEX); Bahia de los Angeles, 12 Feb 1962, 
Wiggins & Thomas 238 (US); 3/4 mi S of Puertocito, 21 Mar 1963, Wiggins & Wiggins 15863 (ARIZ, DS, 
GH, TEX, US*); Arroyo la Bocana near Rancho Santa Ynez, 13 Mar 1991, Van Devender et al. 91-410 
(ARIZ, NMC). BAJA CALIFORNIA SUR: San Marcos Island, 23 Apr 1952, Moran 3975 (UC); 29 Mar 
1962, Moran 9005 (SD); Carmen Id., Marquer Bay, 5 Apr 1962, Moran 9199 (RSA, SD, UC); 1 mi S of 
Mission Los Dolores, 25 05'N, 110 54’W ,4 Dec 1959, Wiggins, Carter, & Ernst 258 (UC). SONORA: Isla 
San Esteban,N side, 10 Apr 1968, Felger et al. 17573 (ARIZ, RSA, SD, UC); Sierra de Rosario, Gran Desierto, 
10 Mar 1973, Felger 20652 (ARIZ, SD).U.S.A. ARIZONA. Yuma Co.: SE side of Tinajas Altas Mts., Borrego 
Canyon, 16 Jun 1992, Felger & Broyles 92-613 (ARIZ, ASU, MO, RSA, TEX, UC); Tinajas Altas Mts., 7 Mar 
1940, Goodding s.n. (ASU). CALIFORNIA. Imperial Co.: Colorado Desert, Coyote Wells, Apr 1905, 
ok (US*); Coyote Wells, 16 Apr 1983, Jonsson & Clemons 472 PD) Painted Gorge, 8 Apr 
1941, Peirson 13075 (DS, RSA); In-Ko-Pah Mtns. along Hwy. 98, 5 1966, "\ ce & Wilkin 110 (RSA). 
Riverside Co.: Devil's Canyon above Coral Reef Ranch, 23 Feb 1931, Ewan 4036 (CAS); West Canon, 
western edge of the Colorado desert, 18 Apr 1907, Parish 6072 NMC, TEX) (acc. to Parish’s note, 
otype); Deep Canyon Wash, 11 Apr 1922, Peirson 2917 (RSA); Deep Canyon drainage, S side 
ie Pipistrelle Canyon wash, 29 Mar beewe Zabriskie and Zabriskie 594 (RSA). San Diego Co., Palm 
Sire Borego Valley, 17 Mar 1940,R . (POM); Borego Desert, canyon, toward the 
Palms, 24 Mar 1939,A.& R. Nelson 3236a (D5): Berean Palm Canyon, 14 Apr 1936, Gander 1301 (SD). 


= 


ACKNOWLEDGMENTS 


We thank the curators of herbaria listed for providing specimens, and for their 
patience in their delayed return. John Strother and Alan Smith were very helpful 
with nomenclatural details and questions regarding typification in the M. laevis 
complex. We also thank Strother for the use of his chromosome number estimate 
in Mirabilis laevis var. retrorsa. Tom Lammers and Don Pinkava provided excel- 
lent help through their careful reviews. Rodriguez Tijerina is particulary grateful 
to his wife, Leticia, and his daughters, Jasmin and Violeta, for their support and 
patience during his studies at New Mexico State University. He also expresses 
gratitude to his parents, Salvador Rodriguez Mireles and Ramona Tijerina de 
Rodriguez, for their unconditional support, and to his brothers, Salvador and 
Rafael, for their moral support. 


REFERENCES 
Baker, H.G. 1961.The adaptation of flowering plants to nocturnal and crepuscular pollina- 
tors. Quart. Rev. Biol. 36:64—73. 


( NYCTAGINACEAE 569 


Boate, L.A. 1974.The genera of Nyctaginaceae in the southeastern United States. J. Arnold 
Arbor. 55:10-20. 

CHATURVEDI, S.K. 1989.A new device of self pollination in Boerhavia diffusa L.(Nyctaginaceae). 
Beitr. Biol. Pflanzen 64:55-58. 

Crupen, R.W. 1973. Reproductive biology of weedy and cultivated Mirabilis (Nyctaginaceae). 
Amer. J. Bot. 60:802-809. 

Curran, M.K. 1888. Botanical notes. |. Plant from Baja California. Proc. Calif. Acad. Sci. ser. 2, 1: 
22/-237. 

DeLorme Mapping. 1990. Southern & Central California Atlas & Gazetteer, 2nd ed. DeLorme 
Mapping, Freeport, Maine. 

Dunkte, M.B. 1941. No. 23. Two new plants from the Channel Islands of California. Bull. S. 
Calif. Acad. Sci. 40:108-109, 

Ewan, J.and N.D. Ewan. 1981. Biographical dictionary of Rocky Mountain naturalists, 1682- 
1932. Regnum Veg. 107:1-252. Dr.W Junk, The Hague / Boston. 

FeLcer, R. 1993. Noteworthy collections - Arizona. Madrono 40:1 78-186. 

Gray, A. 1853. New genera and species of Nyctaginaceae, principally collected in Texas 
and New Mexico. Amer. J. Sci. Arts, ser. 2, 25:259-263, 319-324 

Hernandez, H.M. 1990. Autopolinizacion en Mirabilis longiflora L.(Nyctaginaceae 
Mex. 12:25-30. 

Heimert, A. 1934, Nyctaginaceae.|n:A.Engler and K. Prantl, Die nattirlichen pflanzenfamilien, 
ed. 2, 16c:86-134. 

Houmeren, P.K., N.H. Hotmeren, and L.C. Barnett. 1990. Index Herbariorum, part |: The Herbaria 
of the world. New York Botanical Garden, Bronx. 

Jerson, W.L. 1914. A flora of California, vol. 1, pt.4. Associated Students Store, University of 
California, Berkeley 

KeLLoaG, A. 1863. Descriptions of two new species of plants from Nevada Territory. Proc. 
Calif. Acad. Sci. 3:9-12. 

Le Duc, A. 1993. A new species of Mirabilis (Nyctaginaceae) from Jalisco, Mexico. Sida 
15:585-587. 

Le Duc, A..1995. A revision of Mirabilis section Mirabilis (Nyctaginaceae). Sida 16:613-618. 

Lewis, H. 1963. The taxonomic problem of inbreeders or how to solve any taxonomic prob- 
lem. Regnum Veg. 27:37-44. 

Maceripe, J.F.1918.A revision of Mirabilis, subgenus Hesperonia.Contr.Gray Herb. 56:20-24. 

Munz, PA. and D.D. Keck. 1968. A California flora, with supplement. Univ. of California Press, 
Berkeley. 

Parish, S.B. 1907. Notes on the flora of Palm Springs. Muhlenbergia 3:121-128. 

Rooricuez T.S. 1992. Taxonomy of the Mirabilis california Gray complex (Nyctaginaceae). 
Master's thesis, New Mexico State University, Las Cruces. 

RypserG, PA. 1902. Studies on the Rocky Mountain flora — IX. The Nyctaginaceae of the 
Rocky Mountain region. Bull. Torrey Bot. Club 29:680-693. 

Suinners, LH. 1951. The north Texas species of Mirabilis (Nyctaginaceae). Field & Lab. 19: 
173-182. 


Acta Bot. 


eer 


570 BRIT.ORG/SIDA 19(3) 


SPELLENBERG, R. 2000. Blooming “behavior in five species of Boerhavia (Nyctaginaceae). Sida 
19:311-323. 

SPELLENBERG, R., and R.K. Detson. 1977. Aspects of reproduction in Chihuahuan Desert 
Nyctaginaceae, pp. 273-287.|n:R.H.Wauer and D.H.Riskind, eds. Transactions - Sympo- 
sium on the biological resources of the Chihuahuan Desert Region, U.S.and Mexico. U. 
S. Dept. Interior, National Park Service Transactions and Proceedings, no. 3 

STANDLEY, PC. 1909. The Allioniaceae of the United States, with notes on Mexican species. 
Contr. U.S. Natl. Herb. 12:303-389. 

StanvLeY, PC. 1911. The Allioniaceae of Mexico and Central America. Contr. U.S. Natl. Herb. 
13:377-430. 

Stanotey, PC. 1918. Allioniaceae. N. Amer. Fl. 21:171-254. 

STANDLEY, P.-C. 1931a. The Nyctaginaceae and Chenopodiaceae of northwestern South 
America. Publ. Field. Mus. Nat. Hist, Bot. Ser. 11(3):73-126. 

Stanbtey, PC. 1931b. Studies in American plants — V. Publ. Field. Mus. Nat. Hist, Bot. Ser. 
8(5):295-469 (Nyctaginaceae 304-310). 

Sreseins, G.L. 1957. Self fertilization and population variability in the higher plants. Amer. 
Midl. Naturalist 91:337-354. 

Torrey, J. 1859. Botany of the boundary, Vol. II, pt. 1. In: W.H. Emory, Report on the United 
States and Mexican Boundary Survey, 2 vols. Washington, D.C 

Turner, B.L. 1993. Texas species of Mirabilis (Nyctaginaceae). Phytologia 75:432-451. 

Wicains, LL. 1964. Flora of the Sonoran Desert. Part Il, pp. 189-1740 (Nyctaginaceae, pp. 
472-489). In: F. Shreve and |.L. Wiggins, Vegetation and flora of the Sonoran Desert (2 
vols.). Stanford University Press, Stanford, CA. 


A REVISION OF SALVIA SECTION HETEROSPHACE 
(LAMIACEAE) IN WESTERN NORTH AMERICA 


Jay B. Walker’ Wayne J. Elisens 
Department of Botany and Microbiology Department of Botany and ed 
University of Oklahoma University of Oklahom 
orman, OK 73019, U.S.A. Norman, OK 73019, ton 
ABSTRACT 


A revision of the western North American members of Salvia L. section ee Benth. is pre- 


eee ae Species are here Becoenieet S. roemeriana Scheele, S. henryi A. Gray, and S.summa A 
Nel inder S. henryi. We present eee of sub- 


generic relationships and se pase saaih variation of ‘the group, as well as keys to the species, distri- 
bution maps, illustrations, and a of typification and synonymy for each species. 


RESUMEN 
os revisan taxonomicamente los bepresentanics) norteamericanos ogcdentaies de Salvia L. seccion 
S. henryiGray y S.summa 


L t 


A. Nels o trata Salvia davidsonii indnimode S ee Presentamos una discusion 

de las relaciones SRNEENENCAS y variacion morfoldgica del grupo, claves para las especies, oc de 

la distribucion, ilustraciones y un informe completo de la tipificacion y sinonimia de cada esp 
INTRODUCTION 


The genus Salvia L. (Lamiaceae) is the largest of the mint genera, consisting of 
over 900 species worldwide and nearly 500 species in the New World (Alziar 
1988). The present taxonomic treatment revises the western North American 
members of section Heterosphace Benth., a group of closely allied species of 
Salvia native to southwestern United States and northern Mexico placed by 
Briquet (1897) in subgenus Leonia (La Llave & Lex.) Benth. Prior to this treat- 
ment, four species were recognized in this group: Salvia roemeriana Scheele, S. 
summa A. Nels., S. henryi A. Gray and S. davidsonii Greenm. All are restricted 
generally to limestone substrates and to north- or east-facing cliffs or slopes. 
These species produce chasmogamous, tubular flowers that are pink, red, or 
purple-colored and are presumably adapted for hummingbird pollination in 
the spring and fall, and produce cleistogamous flowers that are self-pollinated 
throughout the growing season. 

There has been considerable variance surrounding proposed species rela- 
tionships within this group of Salvia (Epling 1944, 1960; Spellenberg 1993). 
Much of the taxonomic confusion has related to character polymorphisms 
within the species and to unclear specific boundaries of S. davidsonii. Three 


‘Current address: Department of Botany, University of Wisconsin-Madison, 132 Birge Hall, 430 Lincoln Drive, 
Madison, WI 53706, U.S.A. 
SIDA 19(3): 571 - 589, 2001 


572 BRIT.ORG/SIDA 19(3) 


species are recognized in the present treatment: Salvia summa, S. roemeriana 
and S. henryi. Salvia davidsonii is synonomized with S. henryi. 


Infrageneric relationships within Salvia 
Despite its widespread distribution and the attention the genus has received 
horticulturally, there is no modern comprehensive taxonomic treatment of spe- 


cies or of infrageneric relationships within Salvia. The most recent classifica- 
tion of supraspecific taxa of Salvia (Hruby 1962) lists eight subgenera and 17 
sections. Many of the subgenera and sections proposed in Hruby (1962) and 
proposed in the only two comprehensive treatments of the genus (Bentham 
1848; Briquet 1897) are generally viewed as artificial (Hedge 1974). Over 400 
new species of Salvia have been described since the last generic treatment in 
1897 by Briquet. 

The species studied in this treatment have been classified in Bentham’s sec- 
tion Heterosphace by most previous workers (Briquet 1897; Fernald 1900; Neisess 
1984). Section Heterosphace is notable in that it is the only one of the five sec- 
tions of New World Salvia with both New and Old World members. The sec- 
tion is represented in th. Old World by nine species native to South Africa, and 
one in central and eastern Africa (Hedge 1974). Heterosphace is represented in 
the New World by the southwestern USA species reported here and by Salvia 
lyrata L. of the southeastern United States. Other sections of Salvia native to 
the New World are: sects. Audibertia Benth. (15 species) and Echinosphace Benth. 
(4 species), both restricted to the California Floristic Province and adjacent 
deserts; sect. Salviastrum Scheele G species), restricted to Texas and northeast 
Mexico; and sect. Calosphace Benth. (ca. 470 species) which occurs primarily 
in Central and South America. 

In all species of Salvia, only two stamens are functional; the other two sta- 
mens typical of the majority of Labiates are reduced to staminodes. In sect. 
Heterosphace, the upper two stamens are reduced to staminodes, and the lower 
two are fertile with the two thecae of each anther separated by an elongated 
connective. Whereas the majority of New World Salvia have sterile posterior 
anther thecae, sect. Salviastrum and sect. Heterosphace are unique in the New 
World because their posterior thecae consistently produce viable pollen. Sec- 
tion Salviastrum is distinguished from sect. Heterosphace by a dense annulus 
in the calyx and by simple leaves. Although the presence of an annulus was not 
noted in the initial description of sect. Heterosphace (Bentham 1832-1836), S. 
henryiand S.summa have a thin annulus, which may indicate a close relation- 
ship between sects. Heterosphace and Salviastrum. Section Salviastrum has been 
described as “..nearest Heterosphace, from which it differs in habit and in the 
calyx closed by hairs” (Torrey 1859). Further investigations into the relation- 
ship between these two groups are necessary based on their sharing of an an- 


en 


— 


— 


WALKER AND ELISENS, REVISION OF SALVIA SECT. HETEROSPHACE 573 


nulusand the fertile posterior anther thecae. The relationship of species in these 
two sections to the Texas endemic Salvia penstemonoides Kunth & Bouche, 
which shares characters of both sections, has not been resolved. 


MORPHOLOGY AND CHARACTER ANALYSES 


Morphological investigations were conducted on 537 herbarium specimens 
from 13 herbaria (ARIZ, ASU, GH, MEXU, MO, NMC, NY, OKL, RM, RSA, SRSC, 
TEX, UTEP; Holmgren et. al. 1990) and on live plants collected from 29 native 
populations (listed with specimens examined). Each population collection was 
based on material from 5-10 individuals. Analyses of morphological variation 
were conducted on individuals or collections representing a total of 59 locali- 
ties: 13 localities for Salvia summa, 22 localities for S. roemeriana, and 24 locali- 
ties for S. henryi. Localities were selected to represent the geographic range and 
extent of morphological variation of a species. The specimens examined 
are indicated in the list of representati xamined for each species. 

Morphological data were scored for fee characters that had been 
used previously to classify and identify species in the group. Results were tabu- 
lated and morphological characters evaluated for their utility to discrimi- 
nate among species in the group. Representative characters employed by past 
authors to distinguish among species include: height of plant, petiole length, 
leaf length, leaflet number, leaf shape, calyx length, calyx lip length, calyx 
pubescence, calyx tube length, corolla color, corolla length, corolla lip length, 
corolla shape, corolla throat width, exsertion of stamens, and symmetry of style 
branches (Epling 1960; Peterson ined; Correll & Johnston 1970). The characters 
we found most useful in differentiating among species in this treatment are leaf- 
let size, calyx length, calyx tube length, corolla length and color, and corolla lip 
length (Table 1). 


Habit 

Each species isa mildly aromatic perennial herb from a woody caudex; the stems 
generally die back to the rootstalk or to basal leaves in the winter. Height of the 
plant ranges from 1-9 dm; number of stems ranges from one to many. Observa- 
tions among natural populations and plants cultivated in greenhouse conditions 
suggest that habit is variable and affected by environmental factors such as 
exposure to sun and availability of water. It is not unusual to find two plants of 
the same population demonstrating distinctly different growth habits, one with 
simple leaves and flowers in the axils of leaves, and the other with compound 
leaves and a distinct inflorescence. Such differences usually are associated with 
occurrence at the edge of a stream or on a cliff face above it. Individuals dis- 
playing such characteristics that are transplanted and grown in a greenhouse 
under equivalent environmental conditions assume similar growth forms. 


574 BRIT.ORG/SIDA 19(3) 


Tasie 1. Variation among diagnostic morphological characters in three species of Salvia sect. 
Heterosphace. 


5. roemeriana (N=22) S. henryi (N=24) S$, summa (N=13) 
mean (range) mean (range) mean (range) 
Calyx Length (mm) 11.8(8-15) 10.2(7-13) 8.5(8-11) 
Calyx Tube Length (mm) 6.8(5-9) 4,2(3-6) 3.1(2-5) 
Corolla Length (mm) 33(24-47) 34(28-39) 39(26-48) 
Corolla Lower Lip Length (mm) 7.1(6-9) 4.4(2-6) 8.3(7-12) 
Ratio of Lateral Leaflet Length/ — 0.23(0.05-0.31) 0.61(0.44-1.16) 0.54(0.43-0.75) 


Terminal Leaflet Length 


Corolla Color Red/Scarlet Red/Scarlet Pink/Purple 


Leaf shape, division and texture 

The plasticity of leaf characters also has been examined in experiments con- 
ducted under uniform greenhouse conditions. Individuals of S. summa with 
compound leaves of 5-7 leaflets in natural populations, consistently produce 
simple leaves when grown in the greenhouse with a daily watering regimen. 
Salvia roemeriana exhibits less plasticity than the other species, and consis- 
tently produces either simple leaves or leaves with 2-4 small lateral leaflets. 
Regardless of growth and environmental conditions, the terminal leaflet of S. 
roemeriana maintains a reniform to cordate shape; if lateral leaflets form, they 
are consistently less than 1/3 the length of the terminal leaflet (Table 1). The 
leaves of S. henryi and S. summa are rarely simple, and the lateral leaflets are 
greater than 1/3 the length of the deltoid terminal leaflet. 

Salvia roemeriana is distinctive among the species examined due to the 
close eee of the basal bg cauline leaves (Fig. 3). Salvia henryi and S. 
phic leaves; the basal leaves are compound, 
and the ‘cauline leaves often simple and graded into the bracts of the inflores- 
cence. In all three species, pubescence type varies between cauline and basal 
leaves. This variation is particularly evident in S. summa where the compound 
basal leaves generally have a thick vestiture of grayish trichomes and the simple 
cauline leaves are more sparsely pubescent. Whereas the basal and cauline leat 
margins of S. roemeriana are generally crenate, those of 5S. henryi and S.summa 
are irregularly-toothed. 


Pubescence 

Pubescence is always present on vegetative and reproductive structures in these 
species, although indumentum is variable and includes pilose, setose, hirsute, 
densely rusty pubescent, white tomentose, and spatsey pubescent. The only 
pubescence character we find to be reliable in distinguishing species is that of 
the calyx; consistently hirsute in S. henryi and S. summa contrasted to a 


WALKER AND ELISENS, REVISION OF SALVIA SECT. HETEROSPHACE 575 


puberulous vestiture in S. roemeriana. Resinous dotting is always represented 
on stems, leaves and calyces, although it varies considerably between popula- 
tions. Vegetative pubescence varies similarly and is not used to differentiate 
species in this treatment. 


Inflorescence 

The inflorescence of S. roemeriana is well-defined and raceme-like, with bracts 
to nearly 1.0 cm long and with 2-6 flowers/verticillaster. Salvia henryi and S. 
summa occasionally exhibit a well-defined inflorescence, but more often there 
is a gradation from solitary flowers in the axils of cauline leaves to a raceme- 
like inflorescence with 4-6 flowered verticillasters and bracts of 0.5 cm in 
length. Flowers borne in the axils of cauline leaves always are cleistogamous, 
whereas fully chasmogamous flowers are only borne in the verticillasters. In 
all species, the persistence and size of bracts is variable. 


Cleistogamy and Polymorphism in Corolla Size and Shape 

Two of the most taxonomically significant characters in sect. Heterosphace are 
corolla polymorphism and cleistogamy. Cleistogamy has been documented in 
the species studied here, as well as in the closely related S. lyrata of the south- 
eastern USA (Uttal 1963) and in African species of the genus (Hedge 1974). In- 
florescences of each of the species observed throughout a growing season un- 
der greenhouse conditions produce fertile flowers ranging from fully closed 
cleistogamous flowers less than 0.5 cm long, to barely open flowers of 1.0 cm in 
length, to fully developed flowers of 3.5 cm or more. With the initial flush of 
growth in the early spring, fully developed chasmogamous flowers are produced. 
As the season progresses, progressively smaller flowers are produced from the 
same indeterminate inflorescence. This was observed among S. henryi, S. 
roemeriana, and S.summa, and consisted of chasmogamous [lowers grading to 
smaller flowers continued until exclusively cleistogamous flowers were pro- 
duced in late spring and summer. In the fall, the sequence is often reversed; cleis- 
togamous flowers grade into small chasmogamous flowers until eventually 
fully developed chasmogamous flowers are produced. In wild populations of S. 
henryi, S. reemeriana and S. summa, the only time of year when plants were 
not observed with cleistogamous flowers was in spring. All flowers, regardless 
of size, have fertile pollen and can produce four viable nutlets, although pre- 
liminary observations found a lower number of nutlets produced in fully 
chasmogamous flowers than in cleistogamous flowers. Despite variation in size 
and shape of the corolla, calyx and nutlet characters remain relatively constant. 


lyx 
The calyx provides reliable characters by which to distinguish species (Fig. 1, 
A,D,G). The upper lip of the calyx is two or three-lobed and the lower is two- 
lobed. The calyces of all species are persistent, and the previous year’s calyces 
are often found on the dried inflorescences adjacent to flowering inflorescences 


576 BRIT.ORG/SIDA 19(3) 


Fic. 1 AA hal ate { + Narth A A H £Calnt : Lot, h Cl | hal £ 


Salvia henryi (A-C), Salvia roemeriana (D—F), and Salvia summa (G-l): A,D,G. Oblique view of calyx. B,E,H. Lateral view of 
corolla. C,F. Frontal view of corolla. 


Salvia summa always has a shorter calyx tube than S. roemeriana, but calyx 
length and calyx tube length overlap between S.summa and S. henryi (Table 1. 
Salvia roemeriana and S. henryiare not significantly different (p > 0.05) in ca- 
lyx length, but these species exhibit consistent differences in calyx pubescence 
and structure. 

The calyces of both Salvia summa and S. henryiare hirsute pubescent, with 
ciliate nerves and a bearded sinus resulting from a thin annulus present in the 
calyx throats. The calyx of Salvia roemeriana is uniformly puberulous on the 
exterior of the calyx and naked within. 

Salvia roemeriana has a repressed (0-0.2 mm) middle upper lobe of the 
calyx, compared to the larger middle upper calyx lobe in S. summa (average = 
L.2mm)and S. henryi (average = 1.0 mm). In addition, the apices of each calyx 
lobe of S. henryi and S. summa are generally rigid and spine-tipped, whereas 
the apices of S. roemeriana are less rigid. 


Corolla 

The corollas of these species are tubular and bilabiate; the upper lip has two 
lobes which are often folded together at the apex, and the middle lobe is the 
largest of the three lower lobes. Cleistogamy and polymorphism in corolla size 
and shape complicates the use of corolla characters in identification and clas- 


WALKER AND ELISENS, REVISION OF SALVIA SECT. HETEROSPHACE 577 


sification. Because of size variation among corolla characters, only fully devel- 
oped, chasmogamous flowers provide reliable morphological characters by 
which to classify and identify species. 

The corolla characters most useful in distinguishing S. summa from the 
other species are differences in corolla length and color. Fully chasmogamous 
flowers of S. roemeriana and S. henryi average 3.4 cm long and are scarlet or 
red, whereas those of S. summa average 4.2 cm and are purple or pink (Table 1, 
Fig] B,E,H). Additionally, the corolla throat of S. roemeriana and S. summa is 
taller than wide, compared to S. henryi which is wider than tall (Fig. 1, CED. 
Whereas S. henryi has a straight corolla, the corolla of S. roemeriana and S. 
summa is arcuate. Finally, although the length of the upper lip of the corolla 
does not significantly differ among the three species, the lower lip of S. 
roemeriana (avg. 7.1mm)and S.summa (avg. 8.3 mm) is equal to or longer than 
their respective upper lips, and the lower lip of S. henryi (avg. 4.4 mm) is shorter 
than its upper (Table 1, Fig. 1). With fresh material, when the two lobes of the 
corolla are pressed together, only in S. henryi will the upper lobe exceed the 
lower. 


Androecium 

One of the diagnostic characters of Salvia section Calosphace is the fusion of 
the sterile posterior anther thecae into a rudder which blocks the throat of the 
corolla. In sect. Calosphace this rudder acts as a lever, which is pushed by the 
pollinator as it attempts to access the nectary at the base of the corolla (Serna 
& Ramamoorthy 1993). This action deposits the pollen on the head or back of 
the pollinator; the fulcrum of the lever is the junction of the filament and the 
elongate connective. A similar floral mechanism is shared by members of sect. 
Heterosphace except that the posterior thecae are fertile and not fused. In 
chasmogamous flowers, the posterior thecae block the throat of the corolla in S. 
henryi and S. summa, but do not block the throat of S. roemeriana, thereby af- 
fecting the lever mechanism of pollen deposition in the latter. The difference in 
thecal placement and mechanism is only visible in fresh material and is not 
observable on herbarium specimens. In fully chasmogamous flowers of S. 
roemeriana, the two staminodes develop to 5mm in length; anthers were never 
observed to develop on the staminodes. 


Gynoecium 

Style characters, such as exsertion from galea and symmetry of stylar branches, 
have been used in previous treatments (Peterson, in ed.) to differentiate species, 
but our observations indicated these cl were not informative taxonomi- 
cally. Whereas the upper style branch had previously been reported to be ab- 
sent or less than 0.5 mm in S. henryi (Peterson, in ed.), our observations found 
considerably more variation (0 to 4.1 mm) in the upper branch of the style both 
in fresh and dried material. Symmetry of stylar branches was not found to vary 


578 BRIT.ORG/SIDA 19(3) 


significantly among the species included in this treatment. Variation in the 
length of the style was observed in S. roemerianaand S. henryi. For example, style 
lengths in fully developed, chasmogamous flowers on the same inflorescence 
of a plant representative of S. henryi ranged from 2.5 cm and included within 
the galea, to 4.1 cm and exserted 1.2 cm from the galea. Because heterostyly has 
been documented in sect. Audibertia (Neisess 1984), the role the variation in 
style length plays in S. henryi and S. roemeriana needs further investigation, 
As with all species of Salvia, the bi-loculed, bi-carpellate ovaries of sect. Hetero- 
sphace divide to produce a maximum of four nutlets. A comparative study of 
nutlet characters for species in sect. Heterosphace and other sections of Salvia 
J. Walker, unpublished data) showed uniformity in nutlet size and pericarp 
anatomy among S. roemeriana, S. henryi, and S. summa. Additional studies by 
Hedge (1970), Ryding (1995), and Wojciechowska (1958) indicate that pericarp 
anatomy is potentially useful for resolving supraspecific relationships in Salvia 
and for testing proposed taxonomic relationships among New World and Old 
World species of the genus. The uniformity in nutlet characters in the three 
species in question further support the closely related nature of these species. 


CHROMOSOME NUMBERS 


Mitotic counts made by the senior author yielded sporophyte chromosome 
numbers of 2n= 28 among populations of S. wemeriana, S. henryi,and S.summa. 
The specimens used in chromosome counts are indicated in the list of repre- 
sentative specimens [or each species. Published diploid counts of 2n = 28 are 
represented in only three other species of Salvia (Hedge 1974), all of which are 
native to northern Africa and none of which is placed in sect. Heterosphace: S. 
aegyptiaca L. (also 2n = 12, 26 reported), S. chudaei Battand. & Trab, and S. 
taraxicifolia Hook. (also 2n = 26 reported; Goldblatt 1981). Of the ten African 
species currently placed in sect. Heterosphace, chromosome counts are known 
from only two, S. nilotica Juss. ex Jacq. (2n = 30)and S.aurita Lf. (2n=18)(Hedge 
1974). Radford et al. (1964) reported 2n = 18 for S. lyrata L., the only other New 
World representative of sect. Heterosphace, although this count was not veri- 
tied through a literature reference. Further investigations of chromosome num- 
ber variation are needed to elucidate base numbers and the extent of polyp- 
loidy and aneuploidy in sect. Heterosphace and in the genus Salvia. However, 
Hedge (1974) reported base numbers x = 7,8,9,10, and 11 for African species of 
Salvia and concluded that chromosome numbers shed “. little light upon spe- 
cies affinities or evolutionary developments in the genus 


GEOGRAPHIC DISTRIBUTION 
Salvia roemeriana is the most widely distributed of the species investigated 
here; it ranges from south-central Texas to central Mexico (Fig. 2). In Mexico, 
the species is found throughout the state of Coahuila and is restricted to the 


WALKER AND ELISENS, REVISION OF SALVIA SECT. HETEROSPHACE 579 


ft Tt T | ; 
13s 110° 107.5° 105° Figure 2 
in Distribution of species 
Salvia henryi - 4 
. Salvia roemeriana - * 
= | Salvia summa- 
35° = 
A 
32.5° 
ARIA) 
AA A is 
A e 
a | 32.5° be 
a A 
o Ld s 
& fA ° Py 
eb é we | * Te ag 
A Ss? 30° = 
we Ft bial fe 2 | 
md i e . . 
oe We 
e ee 
1] 
o . 27.59 
~ ae2 S* 
es 
te - 
es e e 
e re 
° 9 5° bee 
2G 25 
e 
e 
e 
: EL See | 
400 km ; ) 22.5 whe 
110° 107.5° 105° 102.5° 100° 97.5° 
i yi ae, | 24 nos 
| | | | | 
Fic. 2. Distrit ra £Cal L Cal Ico. 


Sierra Madre Oriental to southern regions of the state of Tamaulipas. The range 
of S. henryi is centered in south-central New Mexicoand southeastern Arizona, 
with disjunct populations in the central and southern parts of the state oFGhi- 
huahua and as far west and northas the Grand Canyon region in Arizona. Salvia 
summa is restricted to limestone outcrops in mountain ranges of southeastern 
and south-central New Mexico. 

Salvia summa, S. henryi and S. roemeriana are morphologically and geo- 
graphically distinct and overlap in range with one another in only two regions 
(Fig. 2). In both of these areas of sympatry, hybridization, which is well docu- 
mented in Salvia (Emboden 1971), may be present. The Oscura Mountains of 


580 BRIT.ORG/SIDA 19(3) 


south central New Mexico isa zone of sympatry between S.summaand S. henryi. 
Several collections from this area (Spellenberg & Anderson 10865, Anderson & 
Morrow 5085) exhibit intermediacy in vegetative and calyx characters between 
these species, which suggests interspecific hybridization. A similar example of 
sympatry and possible hybridization is the Big Bend region of Texas and Mexico, 
continuing northwest along the Rio Grande. This is the only region where the 
distributions of S. henryi and S. roemeriana overlap, and a number of collec- 
tions demonstrate morphological intermediacy (Hinckley 1669, Worthington 
4425). Plants from this area possess the calyx pubescence and spine-tipped ca- 
lyx lobes of S. henryi, but have a suppressed middle upper lobe of the calyx 
anda red-tinged calyx, both of which are similar to S. roemeriana. Experimen- 
tal studies are needed to test these hypotheses of hybridization. 


TAXONOMY 
Key to the Species 
1. Corolla red; fully chasmogamous corolla less than 3.8 cm lonc 
2. Lower lip of corolla shorter than upper; corolla straight; ely sinus bearded; 


lateral leaflets greater than 1/3 the length of the terminal. 1. Salvia henryi 

2. Lower lip of corolla equal to or longer than upper; corolla arcuate; oe SINUS 
naked; lateral leaflets less than 1/3 the length of the terminal. 2. Salvia roemeriana 
1. Corolla pink to purple; fully chasmogamous corolla 3.8 cm to 4.8 cm long. ___ 3. Salvia summa 


1. Salvia henryi A. Gray (Figs. 1 A,B,C), Proc. Amer. Acad. Arts 8:368. 1872. Typr: 
U.S.A. NEW MExico: on the Mimbres, May 1851, Thurber 245 (LECTOTYPE, here des- 
ignated: GH! ISOLECTOTYPE: NY!). U.S.A. NEW MEXICO: Mimbres, Henry s.n. 
(SYNTYPE: GH! #1552). U.S.A. 1849. Wright s.n. (SYNTYPE: GH! #1551). U.S.A. NEW 
MEXICO: River Mimbres, Bigelow s.n. (SYNTYPE: NY! #7479). 

Thurber 245 is selected as lectotype due to the complete nature of the collection, 
the presence of fully chasmogamous flowers, and its distribution at two major 
herbaria. 


— 


Salvia davidsonii Greenm., Proc. Amer. Acad. Arts 41:246. 1905, TYPE: U.S.A. ARIZONA: Chiricahua M 
, Lemmon 3077 (LECTOTYPE, here designated: GH!). U.S.A. ARIZONA: Southern Ari- 
SYNTYPE:GHI) U.S.A. ARIZONA: Metcalf, Oct 1900, Day) idson 615 (SYNTYPE: GHI, 


tains, Sep 188 
zona, Lemmon 497, 
RSA!) 


— 


Lemmon 3077 is selected as lectotype due to its being the most complete specimen, although 
aul ieee only seats ag flowers, 

areene nom. nud. U.S.A. ARIZONA: Chiricahua Mts., Paradise, Cedar Gulch, 24 Sep 1907, 
piabeat (GH! MO)). 


Perennial to 6 dm from woody caudex. Stems erect to trailing, generally un- 
branched, one to several. Stem puberulous to plore or hirsute, sparsely resinous 
dotted. Basal leaves generally pinnately compound, 3-5(-7) foliate, terminal leaflet 
deltoid, lateral leaflets greater than 1/3 the length of the terminal. Cauline leaves 
often simple, deltoid or less frequently cordate/reniform and reduced upwards 
and grading into bracts. Margins of all leaves irregularly toothed and lobed. 
Leaf size and shape quite variable depending on environment and time of year. 


— 


WALKER AND ELISENS, REVISION OF SALVIA SECT. HETEROSPHACE 581 


Inflorescence raceme-like to 25cm, verticillasters 5-10, each bearing 1-6 flowers. 
Lower-most bracts most often indistinguishable from uppermost leaves, the 
flowers thus appearing axillary in some instances. Bracts rarely deciduous. 
Calyx 8-11(-13) mm long, the tube 3-6 mm, hirsute outside, particularly on veins, 
puberulous inside with thin annulus, sinuses bearded. Lobes of calyx firmly 
mucronate, middle upper lobe 1 mm (occasionally suppressed). Calyx green 
throughout. Corolla red, puberulous, generally not arched, 3.0-3.8 cm, often 
reduced and cleistogamous. Upper lip of corolla galeate, 6-8 mm, lower middle 
lobe 2-6 mm, the lower lip of the corolla shorter than the upper. Corolla throat 
5-6 mm in height, wider than tall. Upper anther thecae exserted as much as 6 
mm from the galea, the lower anther thecae bent downward into throat of co- 
rolla. Style bifid, exserted from galea, top branch 1-2 mm, lower branch 2-6 
mim. 2n= 28. 

Producing fully developed, chasmogamous flowers April-May. Cleistoga- 
mous flowers produced May-October. 

Distribution and habitat.—Preferring limestone talus or cliffsides, in canyons 
or north-facing slopes; 800 m-2000 m. Texas along Rio Grande and canyons in 
Brewster, Presidio, Jeff Davis, Hudspeth and El Paso counties; New Mexico in 
Otero County west to Arizona and north to northern Socorro County; Arizona 
in eastern Pima County, north to Maricopa County and in disjunct populations 
northwest to Grand Canyon. Mexico along Rio Grande (Rio Bravo) and west to 
Santa Eulalia Mountains in Chihuahua, with a disjunct population in southern 
Chihuahua. 

Comments.—The three individuals which serve as syntypes for S. davidsonii 
were plants collected late in the flowering season (September and October) and 
had cleistogamous flowers with corollas 1 cm long or less. Subsequent collections 
made earlier in the growing season by other botanists at the type localities ex- 
hibited fully chasmogamous flowers with corollas to 3 cm long (see discussion 
under corolla characters). The foliar and inflorescence characters outlined in 
Greenman’s description were all commonly found in populations of S. henryi. 
Greenhouse-grown plants propagated by seed from New Mexico populations 
of S. henryi and from a type locality of S. davidsonii (Chiricahua Mts., north of 
Paradise, AZ) were similar morphologically and appeared to be conspecific. For 
the above reasons, S. davidsonii is treated asa synonym with S. henryi, the latter 
having priority by 33 years. 

Disjunct populations of S. henryi in Arizona occur along Fish Creek in 
Maricopa County, Peoples Spring in the Arrastra Mountains in Yavapai County, 
and various locations along the Grand Canyon of the Colorado River. Each of 
these disjunct populations have plants with slight morphological differences. 
Most of the differences observed, such as simple leaves, larger growth habit, and 
pink or purple flowers, may be the result of environmental conditions; these 
characters are apparently phenotypically plastic and not suitable to delimit 


582 BRIT.ORG/SIDA 19(3) 


infraspecific taxa. Collections from populations growing along an environmen- 
tal gradient, from mesic and shaded to xeric and exposed, showed variation in 
leaf shape from simple to compound, and in corolla color from red to pink to 
purple. Additional investigation is required to determine the basis for the ob- 
served morphological variation. 

Common name.—Henry’s Sage. 


Representative Specimens: * = Specimens used in analyses of morphological characters. t = Speci- 
din chromosome counts. UNITED STATES. ARIZONA. Cochise Co.: Chiricahua Mts, 2mi 

N of Paradise, 2 1935, Maguire 11134 (NY)*; Little see Mts, NE of Benson, E of San Pedro 
River, 5 May 1993, Van Devender 93-632 (ARIZ)*.Coconino Co.: Havasupai Cyn, 23 May 1941, Whiting 
a (ARIZ)*; ae River, Matkatamiba Cyn, 26 Oct 1990, Scott 882 (NYBG)*; Matkatamiba Cyn, 
48 of Colorado River, 18 May 1973, Phillips s.n. (ARIZ)*. Gila Co.: Pinal Mts, S of Globe, Jun 1995, 
aes 1971 (OKL)t. Graham Co.: Upper Gila River drainage, 20 Apr 1978, McGill 2376 (ASU, NYBG, 
TEX)*; Bonita Creek between Midnight Cyn and San Carlos Indian Reservation, 21 Apr 1978, McGill 
2388 (ASU, RSA)*. Greenlee Co.: 15mi N of Clifton, 7 Jun 1935, Maguire 11854 (NY)*; 14 mi N of 
Clifton, 1 Apr 1960, Crosswhite 803 (ASU)*. Maricopa Co.: Fish Creek, 1 Apr 1926, Peebles 5233 (ARIZ)”. 
Mohave Co.: along Colorado River, 1/4mi below Matkatamiba Cyn, 4 May 1970, Holmgren 15536 
(ARIZ, GH, NY); Frasiers Well, off hwy 93, between Wickenburg and Kingman, 2 Nov 1968, Niles 906 


ee 


(ARIZ)*. Pima Co.: Empire Mts, 31 May 1987, Montgomery s.n. (ARIZ)*. Pinal Co.: Superstition Mts, 
Fremont Pass, 24 Mar 1972, McGill 433 ae .Santa Cru z Co.: Santa Rita Mts, 25 May 1884, Pringle 
5.n.(GH, NY); west end of Canelo Hills, ab Monkey Springs, 11 Jun 1978, Kaiser 1193 (ARIZ). Yavapai 


Co.: Southern Arastra Mts, People’s Cyn, South People’s Spring, 4 Jun 1979, Fischer 6628 (ARIZ, ASU)*. 
NEW MEXICO. Dona Ana Co.: 43 air km NNE of Las Cruces on W side of San Andres Mts, 15 May 
1993, Spellenberg 11799 (BRIT, MT, NMC); East Portrillo Mts, 25 Apr 1992, Worthington 20581 (UTEP)*; 
11 air mi NW of Las Cruces, N end of Robledo Mtn, 27 Apr 1983, Spellenberg 7007 (NMC)*.Grant Co.: 
10mi NE of Red Rock, 21 May 1935, Maguire 11552 (ARIZ); Little Hatchet Mts, Howell's Ridge, 14 May 
1992, Worthington 20802 (UTEP). Hidalgo Co.: Big Hatchet Mts, 2 air mi NNE from top Big Hatchet 
Peak, 19 May 1992, Worthington 20891 (UTEP). Lincoln Co.: 12 mi E of Carizozo, 22 May 1987, Barneby 
18236 (NY)*;WSMR, Oscura Cottonwood Spring, 3 Jun 1993, et 6205 (TEX)*. Luna Co.: 
Florida Mts, Mahoney Park, 18 Apr 1982, Worthington 8117 (ASU)*. 2 Sacramento Mts, Dog 
Cyn, 3 Jun 1979, Van fete s.n. (UTEP ARIZ)*; Sacramento ene " 7 Fresnal Cyn Tunnel, 11 
May 1980, Worthington 5898 (ARIZ, UTEP)”. Sierra Co.: Fra. Cristobal Range, cliffs of S side of Chalk 
Gap, 16 Jun 1981, Van Devender 16191 (UTEP); Bear Den Cyn, WSMR, 28 Aug 1991,Anderson 5085(NMC). 
Socorro Co.: White Sands Missile Range, N end of Oscura Mts, 19 Aug 1991, Spellenberg 10865 (NMC, 
UNM,NY,MO,RSA,ID, UC, TEX)*. TEXAS. Brewster Co.: 4mi NE of Lajitas, 18 Apr 1973, Jump s.n. (ARIZ). 
El Paso Co.: Franklin Mts, 6 May 1983, Worthington 10287 (UTEP)*; Franklin Mts, 8 Apr 1978, 
en 2508 (UTEP)*: Hueco Pass, Hueco Mts, 27 Apr 1976, Butterwick 2619 (TEX)*. Hudspet 

ntral Hueco Mts, along hwy 180 near Hueco Inn, 26 Apr 1975, Everitt 75255 (ARIZ). Jeff Le 
Co.: N of Alpine in Fern Cyn, 20 Apr 1938, Warnock 1399 (TEX). Presidio Co.: at base of south bluff in 
Bracks eae 11 Jun 1941, Hinckley 1669 (GH, NY, SRSC)*. 

XICO. Chihuahua: Santa Eulalia Mts, Apr 1886, Pringle 704 (MEXU, TEX)*;Canon del Rayo at 
ae end of the Sierra del Diablo, 25 Jul 1941, Stewart 907 (TEX); 23 air mi ENE of Villa Anumada 
in northwestern cyn of Sierra de la Alcaparra NE of Rancho El Palmar, 12 Sep 1973, Henrickson 12853b 
(TEX); SE of Lajitas, TX, 5 May 1979, Worthington 4425 (UTEP)*. 


— 


— 


2. Salvia roemeriana Scheele (Figs. 1 D,E,F, Fig. 3), Linnaea 22:586. 1849. Tyr: 
US.A. TEXAS: fertile soil in shady woods on the upper Guadalupe, Lindheimer 145 
(LECTOTYPE, here designated: GH; ISOLECTOTYPE: MON). 


583 


WALKER AND ELISENS, REVISION OF SALVIA SECT. HETEROSPHACE 


1854). 


Fig. 3. Habit of § 


BRIT.ORG/SIDA 19(3) 


Scheele described this species from a Roemer collection (In silva prope 
Neubrauntfels leg. Roemer. Aprili). However, no Roemer collection of S. roemeriana 
is present in any of the herbaria surveyed within this project. Neither are any at 
the Munich herbarium (M) where many of Scheele’s specimens are deposited. 
Roemer was in the vicinity of New Braunfels, Texas both in April of 1846 and 1847. 
During the April of 1846, Roemer’s journal states he collected with Lindheimer in 
the New Braunfels area (Mueller, 1935). As is recounted by McKelvey (1955) 
“Lindheimer and Roemer made many botanical excursions together during 
1846..At the end of the season they appear to have exchanged a set of the collec- 
tions make by each during the year and Roemer, on his return to Germany, placed 
Lindheimer’s with his own botanical specimens in the hands of Adolph Scheele 
who..published the descriptions in Linneae from 1848 to 1852. Not only did he 
publish the ‘new species’ of Roemer’s collecting, but also those found among 
Lindheimer’s duplicates, though he knew that Englemann and A. Gray had al- 
ready undertaken to describe these collections in their Plantae Lindheimerianae, 
and so industriously did he continue his work that he soon completely outdis- 
tanced his American competitors and left little for them to describe.” Owing to 
- lack of any Roemer Soe . S. roemeriana, the authors of this paper as- 
ne that the type specimen was not a Roemer collection as suggested by Scheele, 
a the Lindheimer collection ey designated as lectotype. 
Salvia porphyrantha Decne, Rev. Hort. ser. 4, 3:301. 1854. Type: This species was de- 
scribed from specimens grown from seed at the Paris Museum. The source of the 
seed is unknown. Salvia porphyrat d Hook, Bot. Mag. t.4939. 1856. (orth. var.) 


— 


Perennial to 9 dm from woody caudex. Stems erect to trailing, generally un- 
branched, one to several. Stem puberulent to white tomentose or densely rusty 
pubescent, most often pilose-setose, sparsely or rarely densely resinous dotted. 
Basal and cauline leaves similar, petiolate, simple or 3-5 foliate with lateral leaf- 
lets not greater than 1/3 the length of the terminal leaflet. Lateral leaflets sessile 
(rarely petiolate), orbicular and irregularly toothed, occasionally reduced to 
tooth-like appendages. Terminal leaflet 1-5 cm, cordate to reniform, crenate or 
less often irregularly toothed, often denticulate. Leaves often somewhat rug- 
ose, pubescence variable and generally reflecting that of the stem. Inflorescence 
an raceme-like, 5-15(-30) cm, verticillasters 4-10(-17), each bearing 1-6 flow- 
ers. Bracts generally distinct from cauline leaves and reduced upwards, early 
deciduous or persistent. Calyx puberulous and sparsely resinous dotted out- 
side, naked inside, (8-)L0-12(-15) mm long, the tube 3-6 mm long. Middle up- 
per lobe of the calyx generally suppressed (occasionally expressed and 0.1-0.3 
mm) lower lip 2-lobed, softly mucronate. Calyx green blending to red at the 
apex of the lobes. Calyx sinuses naked. Corolla red, puberulous, arcuate, 2.8- 
3.6 (-4.7) cm, of ten reduced and cleistogamous. Upper lip of corolla weakly gale- 
ate, 0-8 mm, lower middle lobe 6-9 mm, the lower lip equal to or longer than 
the upper. Corolla throat 5-7 mm in height, taller than wide. Two staminodes to 
5mm in fully chasmogamous flowers. Upper anther thecae included or exserted 
as much as 6 mm from the galea, the lower anther thecae bent outward or rarely 


WALKER AND ELISENS, REVISION OF SALVIA SECT. HETEROSPHACE 585 


somewhat downward. Style bifid, exserted from galea, top branch lmm, lower 
branch 2-3 mm. 2n= 28. 

Producing fully developed, chasmogamous flowers March-May. Cleistoga- 
mous flowers produced May-October. 

Distribution and habitat.—Preferring limestone cliffs and talus and juni- 
per or oak/pine woodlands; 500 m - 2000 m. Texas along Edwards Plateau and 
north to Bell County, west to Presidio County. Mexico along Rio Grande (Rio 
Bravo) from Big Bend National Park and south along Sierra Madre Oriental to 
southern Tamaulipas. 

Comments.—Several local variants characterize S. roemeriana. For example, 
a densely rusty pubescent form of S. roemeriana is found in the vicinity of 
Melchor Muzquiz in Coahuila and a densely white pubescent form is found 
along the coastal plain northeast of Ciudad Victoria in Tamaulipas. Individu- 
als of S. roemeriana found in the Sierra Madre Oriental south of Monterrey are 
considerably larger than their northern counterparts and reach a meter in 
height, with flowers of 4.7 cm long. Because there exists a continuous grada- 
tion of size and pubescence types among each of these local variants and the 
more northern members of the species, discrete morphological and geographic 
groupings are not apparent and subspecific rank is not warranted. 

Salvia roemeriana is distinguished from S. henryi by a larger lower corolla 
lip compared to the upper lip, a corolla throat that is taller than wide, the lack 
of an annulus or bearded sinus in the calyx, the lack of a middle upper calyx 
lobe, and lateral leaflets less than 1/3 the length of the terminal leaflet. 

Common name.—Cedar Sage. 

Representative Specimens:* = Specimens used in analyses SA ees snes t = Speci 
mens used in chromosome counts. UNITED STATES. TEXAS. Andrews Co. W of Andrews, 10 
May 1958, Scudday 222 (SRSC). Bandera Co.: just NE of Can Creek, 18 Jun 1 on oe ith 684a (TEX)*. 
Bell Co.: Tennessee Valley area, 17 Apr 1954, nes 54156 (TEX)*. Bexar Co.: 18mi SW of San Antonio, 
Helotes Creek, 15 May 1932, Metz 256 (NYBG, RSA). Blanco Co.: at“The Narrows’ of the Blanco River, 
19 Apr 1969, Correll 37019 (TEX)*. Brewster Co.: 28 mi E of Marathon, 23 Apr 1949, Warnock 8522 
(SRSC)*. Comal Co.: W edge of New Braunfels, 17 Apr 1966, Correll 32508 (TEX)*; Coryell-Bell Co.: 
line:8.8mi NE of Killeen, 23 May 1979, Sherwood 554 (OKL).Edwards Co.: 1/2mi S of the dam on the 
west bank of Hackberry Creek, 21 Jul 1974, Smith 253 (TEX). Hays Co.: College Camp, 8 April 1963, 
Pete Abrigo s.n.(TEX).Jeff Davis Co.: NE Davis Mts, Nations Ranch, eastern edge of Timber Mtn, 3 Jun 
1987, Larke 777 (SRSC). Kendall Co.: below Edge Falls, 3 May 1947, Tharp 17T151 (TEX)*. Kinney Co.: 
Anacacho Mts., Anacacho Ranch, 19 Apr 1966, Correll 32539 (TEX).Pecos Co.: Near Sheffield, Owens 
Ranch, 18 Apr 1953, Souddy 48 (RSC), Presidio Co.: Bofecillos Mts., Bofecillos Cyn.,at Aqua Adentro 
and Cuevas Amarillas, 25 Mar 1994, Worthington 23073 (UTEP)*. Real Co.: Smi N of Leakey, along Frio 
River, 24 Jun 1946, Correll 12865 (NY, TEX). Terrell Co.: 30 mi. NE of Sanderson on Sheffield road, 10 
Jul 1950, Surratt 277 (SRSC); 3mi W of Austin, 20 Oct 1996, Walker 1962 (OKL)t. Travis Co.: 5mi W of 
Austin, 7 April 1949, Rogers 6736 (TEX)*. Uvalde Co.: along Rio Frio, 22 Jun 1963, Correll 28018 (TEX). 
Val Verde Co.: on Fawcett Ledge 20-30 mi up Devil's River, 3 Apr 1953, Warnock 11178 (SRSC,TEX)*. 
Williamson Co.: 3mi W of Georgetown, along north fork of Gabriel River, 28 Sep 1958, Correll 20479 
(TEX). 


586 BRIT.ORG/SIDA 19(3) 


MEXICO. Chihuahua: 23 air mi ENE of Villa Anumada in northwestern cyn of Sierra de la 
Alcaparra NE of Rancho El Palmar, 12 Sep 1973, Henrickson 128536 (TEX); 3mi W of Santa Elena 
picnic Area of Big Bend NP, Fern Cyn, side cyn of Santa Elena Cyn, 2 Nov 1973, Wendt 82 (TEX). 
Coahuila: Muzquiz, 1935, Marsh 152 (TEX)*; Mcpo. aint 1km W of Rancho Lagunita, 10 Apr 
1976, Riskind 1904 (TEX)*; Sierra - la Madera NW of Cuatro Cienegas, in Canada la oa 14 May 
1992, Mayfield 1348 (TEX); W of Buenavista, Saltillo, Canon de San Lorenzo, 9 Apr 1976, Marroquin 
3557 (MEXU)*; Mcpo. Villa Acuna, 80km SE of Big Bend NP on SW margin of Serranias - Burro (part 
of the Sierra del Carmen), Rancho El Rincon, 22 Jun 1991, Ruiz 37 (SRSC)*; 28 air mi WSW of Cuatro 
Cienegas, Canon Los Pozos, 5mi W of Rancho Cerro de la Madera towards Canon Desiderio, 2 May 
1977, Henrickson 15989a (TEX)*; Mcpo. Ramos Arizpe, Sierra S. J. de los Nuncios, 2 Apr 1993, Hinton 
22769 (TEX)*. Nuevo Leon: S of Monterrey on road W of Horsetail Falls, Huesteca Cyn, 13 Nov 1989, 
Starr s.n.(TEX)*; Mcpo. Galeana, 10 km NE of Pacitos, 26 Aug 1984, Hinton 18768 (TEX)*; Mcpo. San- 
tiago, San Isidro, 16 Jun 1994, Hinton 24366 (MEXU, TEX)*; SE of Casablanca-Villa de Garcia, Canada 
Cortinas, 20 Apr 1983, Villarreal 2046 (MEXU)*. Tamaulipas: Mcpo. Hidalgo, Los Mimbres, 5 Jun 1994, 
Hinton 24189 (MEXU, TEX)*; Mcpo. San Carlos, Sierra de San Carlos, Cerro de! Diento, 17 km S of San 
Carlos, 22 May 1988, Hernandez 2271 (TEX)*; Ejido Ricardo Garcia o La Presita, km 66 carretera Victo 
ria-Tula, 26 Apr 1985, Mahinda 314 (MEXU). 


3. Salvia summa A. Nels. ne : G,H,D, Amer. J. Bot. 18:432. 1931. Type: U.S.A. NEW 
MEXICO: Carlsbad Cave ock ere near springs, May 1930, G. Convis 59 (HO- 
LOTYPE: RM! 135094; IsoT YPE: FRM 13806 

Perennial to 40 cm from woody caudex. Stems erect to trailing, generally un- 

branched, one to several. Stem pubescent, often densely pilose at base, resinous 

dotted. Basal leaves thickened, generally pinnately compound with 5-7 leaf- 
lets, terminal leaflet deltoid, 10-25 mim, lateral leaflets variable in shape, greater 
than 1/3 the length of the terminal. Cauline leaves grading from compound 
near base into simple and deltoid (rarely reniform). Leaves somewhat rugose, 
generally pubescent with thick vestiture of appressed hairs, particularly be- 
low, margins irregularly toothed and lobed. Inflorescence raceme-like to 20cm, 
verticillasters 4-8, each bearing 1-4 flowers. Lower-most bracts often indistin- 
guishable from uppermost leaves, the {lowers thus appearing axillary in some 
instances. Bracts rarely deciduous. Calyx 7-8 mm long, the tube 2-4 mm, hir- 
sute outside, particularly on veins, puberulous inside with annulus; sinuses 
bearded. Calyx lobes firmly mucronate, the middle upper lobe 1mm (rarely 
suppressed). Calyx dark green throughout. Corolla purple, often with blue dots 
in throat, puberulous, arcuate, 3.0-4.8 cm long, often reduced and cleistoga- 
mous. Upper lip of corolla galeate, 7-9 mm, lower middle lobe 7-10 mm. Co- 
rolla throat 5-7 mm in height. Upper anther thecae exserted from galea as much 
as 2mm, lower anther thecae bent downward into the throat of the corolla. Style 

bifid, exserted from the galea, top branch Imm, lower branch 1-3 mm. 2n = 28. 

Producing fully developed, chasmogamous flowers April-May. Cleistoga- 
mous flowers produced May-October. 

Distribution and habitat.—Preferring north-facing limestone cliffs, also in 
canyons along water courses; 1500 m-2000 m. Known only from six mountain 
ranges. Texas in Culberson and E] Paso Counties, New Mexico in western Eddy, 


WALKER AND ELISENS, REVISION OF SALVIA SECT. HETEROSPHACE 587 


Otero, southern Chaves, and eastern Dona Ana counties. Mexico in Sierra Juarez 
mountains southwest of Ciudad Juarez, Chihuahua. 

Comments.—Salvia summa has the most restricted distribution of the spe- 
cies studied. Described from Carlsbad Caverns in New Mexico, it was thought 
to be endemic to the Guadalupe Mountains until recently when new locations 
were described from mountain ranges farther west (Worthington 1982). Within 
the Guadalupe Mountains, S. summa is the only representative of sect. 
Heterosphace, whereas in the Oscura, San Andres and Franklin mountains, S. 
summa coexists and possibly hybridizes with S. henryi (see discussion under 
Geographic distribution and interspecific sympatry). The most distinctive fea- 
ture of S.summais the long, purple corolla, which is unmistakable compared to 
other members of the group. Its habit is more diminutive than other members 
of the group and its leaves are thicker. The calyx also is distinctive because of 
its small size and thicker annulus. 

Common name.—Summa Sage. 


Representative Specimens:* = Specimens used in analyses of morphological characters. tf = Speci- 
chromosome counts. UNITED STATES. NEW MEXICO. Chaves Co.: 30mi E of Elk, 18 
Apr 1949, Goodman 5005 (OKL); 9mi W of Hope, 30 May 1997, Walker 2160 (OKL)*. Dona Ana Co 
30mi NE of Las Cruces on E side of San Andres Mts, NE side of Black Mtn, 28 Apr 1990, ere 
10170 (NMC)*; Organ Mts, Rattlesnake ridge, 31 May 1980, Worthington 6050 (NMC, UTEP)*. Eddy 
Co.: Carlsbad Caverns, May 1930, Convis 59 (RM); North Rocky Arroyo, 20mi W of areas oy May 
1997 Walker 2147 (OKL)*. TEXAS. Culberson Co.: Guadalupe Mts, McKittrick Cyn, 21 May 1984, Free- 
man s.n. (UTEP)*; Guadalupe Mts, McKittrick Cyn 30 Apr 1961, Warnock 18244 (SRSU)*; Beira 
Mts, Bear Cyn, 26 May 1979, Warnock 21601 (SRSC)*; Guadalupe Mts, above Pine Springs camp- 
ground, 27 Oct 1996, Walker 1964 (OKL)t; Guadalupe Mts, W side of mts, Goat Spring, 29 May 1997, 
Walker 2157 (OKL)*; Delaware Mts, 9 Sep 1994, Worthington 23532 (UTEP)*; Guadalupe Mts, Pine 
Spring, 1 May 1971, Weston 86 (SRSC)*; 9mi N of Van Horn, 24 Apr 1961, Correll 23808 (MO, NY, RSA, 
TEX)*. El fe Co.: Franklin Mts, 30 May 1981, Worthington 7135 (UTEP)*. 
ICO. Chihuahua: Mcpo. Juarez, in the Sierra Juarez in middle of range, 3 air km SW of Cd. 
Juarez center, 7 May 1993, Spellenberg 11795 (BRIT, MEXU, MT, NMC)*. 


ACKNOWLEDGMENTS 


We wish to thank Guadalupe Mountains National Park and Lincoln National 
Forest, both of which permitted the collecting of specimens integral to this 
project, and the curators of the several herbaria listed in the text whose loans of 
specimens made this study possible. Cal Lemke provided tremendous help in 
the germination and care of live collections. We are grateful to Roger Sanders, 
Barney Lipscomb, and John Nelson for their helpful comments and review of 
the manuscript. We also thank Richard Spellenberg for his encouragement and 
sharing his extensive knowledge on this group of Salvia. We are grateful to Mike 
Powell and to the Rancho Santa Ana Botanical Garden for help in procuring 
seeds of the plants studied, and to Olof Ryding for his help in developing meth- 
odology for pericarp sectioning. 


BRIT.ORG/SIDA 19(3) 


REFERENCES 
Aiziar, G. 1988. Catalogue synonymique des Salvia du Monde |. Biocosme Mesogeen 5 
(3-4):87-136 
Aiziar, G. 1989a. Catalogue synonymique des Salvia du Monde II. Biocosme Mesogeen 
6(1-2):79-115. 
Aiziar, G. 1989b. Catalogue synonymique des Salvia du Monde Ill. Biocosme Mesogeen 
6(4):163-204. 
Aiziar, G. 1990. Catalogue synonymique des Salvia du Monde IV. Biocosme Mesogeen 
7(1-2):59-109. 
Aiziar, G. 1992. Catalogue synonymique des Salvia du Monde V. Biocosme Mesogeen 
9(2—-3):413-497. 
Aiziar, G. 1993. Catalogue synonymique des Salvia du Monde VI. Biocosme Mesogeen 
10(3-4):33-117. 
BentHam, G. 1832-1836. Labiatarum genera et species. 260-698. Ridgeway, London. 
BenTHAM, G. 1848. Labiatae. In: De Candolle: Prodromus 12:262-358. 
Briquet, J. 1897. Salvia. In: Engler-Prantl: Die naturlichen pflanzenfamilien 4(3a):2 70-286. 
Corre, D.C.and M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research 
Foundation, Renner 
Decaisne. 1854. Salvia porphyrantha. Rev. hort. ser. 4 (3):301. 
Empopen, W.A.1971.The role of introgressive hybridization in the development of Salvia: 
Section Audibertia. Contr. Sci. Mus. Nat. Hist. Los Angeles, No. 208:1-15. 
EpLina,C. 1939.A revision of Salvia subgenus Calosphace.Repert. spec. nov. regni. Veg. Beih. 
110: 1-383. 
EPLING, C. 1944. Supplementary notes on American Labiatae Ill. Bull. Torrey Bot. Club 71: 
488-489, 
EpLING, C. 1960. Labiatae. In: Kearney, TH. and R.H. Peebles, Arizona flora. University of Cali- 
fornia Press, Berkeley and Los Angeles. Pp. 731-748 
FerNALD, M.L. 1900.A synopsis of the Mexican and Central American species of Salvia.Proc. 
Amer. Acad. Arts 35(25): 489-556. 
Gotpstatt, P. (ed.) 1981. Index to plant chromosome numbers 1975-1978. Missouri 
Botanical Garden, St. Louis, MO. 
Heoce, |.C. 1970. Observations on the Mucilage of Salvia Fruits. Notes Roy. Bot. Gard. 
Edinburgh 30:79-95 
Heoae, |.C. 1974. A Revision of Salvia in Africa. Notes Roy. Bot. Gard. Edinburgh 33:1-121. 
Houmeren, P.K., NH. Houmoren, and L.C. BarneTT. 1990.Index herbariorum, Eighth Edition. New 
York Botanical Garden, Bronx. 
Hrusy, K. 1962. Key to the supraspecific taxa of the genus Salvia. Preslia 34:368-373. 
Martin, W.C.and C.R. Hurctins. 1981.A flora of New Mexico. J.Cramer, Vaduz,West Germany. 
McKetvey, S.D. 1955. Botanical exploration of the trans-Mississippi west, 1790-1850. The 
Arnold Arboretum of Harvard University, Jamaica Plain. 
Muetter, O. 1935.A translation from German of Roemer’s “Texas, with particular reference 


WALKER AND ELISENS, REVISION OF SALVIA SECT. HETEROSPHACE 589 


to German immigration and the physical appearance of the country. Described through 
personal observation by Dr. Ferdinand Roemer.” Standard Printing Company, San 
Antonio, Texas. 

Neisess, K.R 1983. Evolution, systematics and terpene relationships of Salvia section 
Audibertia. PhD Thesis. University of California, Riverside 

Neisess, K.R 1984. Notes and news, heterostyly in Salvia brandegei. Madrono 31:252-254. 

Peterson, K.M. (in editing). Salvia. Flora of the Chihuahuan Desert 

Raprord, A.E., H.E. AHLEs, and C.R. Bett. 1964. Manual of the vascular flora of the Carolinas. 
The University of North Carolina Press, Chapel Hill. 

RyoinG, O. 1995. Pericarp structure and phylogeny of the Lamiaceae-Verbenaceae com- 
plex. Pl. Syst. Evol. 198:101-141. 

Serna, A.E. and T.P. RAMAMOooRTHY, 1993. Revision taxonomica de Salvia seccion Sigmoideae. 
Acta Bot. Mex. 23:65-102. 

SPELLENBERG, R. 1993. Noteworthy collections, Salvia davidsonii. Madrono 40:137-138. 

StaFLeu, F.A.and R.S. Cowan. 1983. Taxonomic literature. A selective guide to botanical pub- 
lications and collections with dates, commentaries and types. Vol V.Bohn, Scheltema & 
Holkema, Utrecht, The Netherlands. 

Torrey, J. 1859. Botany of the boundary; Emory’s report of the U.S.-Mexican boundary 
survey 2(1):132. 

Urtat, LJ. 1963. Cleistogamy in Salvia lyrata. Castanea 28:162-163. 

Wojciechowska, B. 1958. Taxonomy, morphology and anatomy of fruits and seeds in the 


genus Salvia. Monogr. Bot. 6:3-55. 
WorTHINGTON, R.D. 1982. Noteworthy collection, Salvia summa. Madrono 29:217. 


590 BRIT.ORG/SIDA 19(3) 


Book REVIEW 


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7. Minor Crops [currants and gooseberries, kiwi, amelanchiers, highbush cranberry] 


= Appendix: North American Nursery Sources for Berry Plants 
= Glossary 

« References and Other Resources 

» Plant Name Index 

* Subject Index 


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4060, U.S.A, 


SIDA 19(3): 590, 2001 


TAXONOMY OF STENARIA (RUBIACEAE: HEDYOTIDEAE), 
A NEW GENUS INCLUDING HEDYOTIS NIGRICANS 


Edward E. Terrell! 


Research Collaborator 
Department of Botany 
Smithsonian Institution 


Washington, D.C. 20560, U.S.A. 


ABSTRACT 
Five North American species formerly in the genus He I ; Rubiaceae) are revised 
and placed in the newly recognized genus Stenaria, a name fee by Ralinesaie for a subgenus in the 
genus Houstonia. Keys, descriptions, distributions, and E ided. The principal 
cies is Stenaria nigricans, a wide-rangin i i t central, and south- 


western United States and ee and central Mente. The remaining species are S. butter wickiae, 
known only from Brewster Co., Texas; S. mullerae of Coahuila, Mexico, and Brewster Co., Tex 

rupicola of southwestern Texas and northern Coahuila, Mexico, and S. umbratilis, rare and sur at 
in northern Mexico south to Veracruz. Varieties are recognized within S. mullerae, S. nigricans, and S. 


umbratilis. 


RESUMEN 


Se revisan cinco especies de Norte América anteriormente incluidas en el género Hedyotis 
(Hedyotideae; Rubiaceae) y se never en el ae nuevo y reconocido Stenaria, un nombre usado 


or Rafinesque para un del género Houstonne. Se presentan eave es, jar Nciones, 
fs 
distribucién, y sinonimos. | inci 


r I 


fo) L 
2 centrales loeste de los Estados Unid i | 


ry 

la parte norte y central de México. Las otras especies son S. butterwickiae, conocida unicamente de 
Brewster Co., Texas; S. mullerae de Coahuila, México y de Brewster Co., Texas; S. rupicola del sudoeste 
de Texas y norte de Coahuila, México; y S. umbratilis, especie rara y disyunta de la parte norte de 
México al sur de Veracruz. Se reconocen variedades de S. mullerae, S. nigricans, y S. umbratilis. 


Stenaria, newly recognized at generic rank, includes the widely distributed 
polymorphic species, Hedyotis nigricans, and four other related species that 
occur in the southwestern United States or Mexico: Hedyotis butterwickiae, H. 
mullerae, H. rupicola, and H. umbratilis. This group of five species has been 
called the Hedyotis nigricans group (Terrell 1996a). 

The choice of a type species for the genus Hedyotis has been the subject of 
some controversy regarding which of two Linnaean species, H. auricularia or 
H. fruticosa, should be selected. Hedyotis auricularia has oldenlandioid seeds, 
although they differ slightly from typical species of Oldenlandia. Seeds of H. 
fruticosa are quite different from those of H. auricularia. Conservation of the 
name H. fruticosa as type would avoid disruption in Oldenlandia nomenclature 


iCurrent address: Edward E. Terrell, 14001 Wildwood Dr, Silver Spring, MD 20905, U.S.A. 
SIDA 19(3): 591 — 614. 2001 


592 BRIT.ORG/SIDA 19(3) 


Whichever name is chosen, the genus name Hedyotis does not apply to the 
Hedyotis nigricans group, which has seeds differing entirely from the seeds of 
either H. auricularia or H.fruticosa; consequently a new name is needed for the 
H. nigricans group. Rafinesque’s name for a subgenus of Houstonia, subg. 
Stenaria, is here elevated in rank as the name for the new genus. 

Seed characters are very important in the classification of the tribe 
Hedyotideae (Terrell et al. 1986; Terrell 1991, 1996a). The species of Houstonia 
were grouped into subgenera and sections based mainly on seed morphology 
and chromosome number (Terrell 1996a). Certain Houstonia species (e.g., H. 
rubra Cav.and H. subviscosa (Wright ex A. Gray) A. Gray) differ considerably in 
corolla size and somewhat in corolla morphology, but nonetheless have similar 
seed morphology and the same chromosome number, and are classified in the 
same section. Thus, corolla morphology by itself may be misleading as a taxo- 
nomic character. It is likely that research on other tribes of the Rubiaceae will 
demonstrate the importance of seed morphology; for example, Diodia and 
Richardia in the Spermacoceae differ greatly in seed morphology from the 
Hedyotideae. Robbrecht (1989) successfully utilized pyrene and seed charac- 
ters in studying Chazaliella in the African Psychotrieae. 

Hedyotis nigricans differs from Houstonia in having a chromosome num- 
ber of x=9 or 10 (chromosome data in this group are known only for H. nigricans) 
and non-crateriform seeds, which are somewhat pressed and ellipsoid with 
acentric punctiform hilum (Fig. L). In contrast, the 20 species of Houstonia have 
basic chromosome numbers of 6, 7,8, and Ll, and crateriform seeds (witha ven- 
tral depression occupied by a linear hilar ridge or having a ventral subglobose 
cavity without a hilar ridge); these were discussed and illustrated by Terrell 
(1996a). The genus Oldenlandia has a chromosome number of x=9, but differs 
entirely in having very small, numerous, 3-angled seeds. Hedyotis nigricans has 
been seen growing naturally in the same habitat or near species of Houstonia 
and Oldenlandia without any evidence of hybridization, nor have any hybrids 
been found in herbaria. Hedyotis nigricans var. nigricans is widely distributed 
in the United States and Mexico, and differs conspicuously from Houstonia and 
Oldenlandia species by having elongate capsules instead of globose ones, as 
well as by having seeds differing as already described. 

A related group is the presently recognized Hedyotis mucronata group, 
mostly native to Baja California. These eight species have a chromosome num- 
ber of x=l3 and ellipsoid or dorsally strongly ridged seeds with punctiform hila. 


— 


SYSTEMATIC TREATMENT 


Stenaria (Ral.) Terrell, stat. nov. TYPE SPECIES: Stenaria nigricans (Lam.) Terrell. 
Houstonia subg. Stenaria Raf, Ann. Gen. Sci. Phys. sac 1820. LECTOTYPE, here 
designated: Houstonia rupestris Raf., Ann. Gen. Sci. Phys. 5:226. 1820 [=Hedyotis 
nigricans (Lam.) Fosb|] 


TERRELL, TAXONOMY OF STENARIA 593 


ing centric hilum, tilted 50°. b. 5. nigricans, ventral view, 
e. $. rupicola, ventral view. f. 5. umbratilis, ventral view. Bars=0.1 
mm. 5. nigricans, Terrell 3923 (US), Manitee Co., FL; S. mullerae, Chiang et al. 9421 (TEX), Coahuila; S. rupicola, Tharp 43- 
866 (TEX), Pecos Co., TX; $. umbratilis, Johnston et al. 12032B (TEX), Coahuila. 


Fic. 1.Seeds of S i ies.a.5. nigricans lvi I 


not tilted. c, d. 5. mullerae dd 


Houstonia [unranked] Angustifoliae Small, Man. S.E. FL. 1254. 1933. TyPE: Houstonta 
angustifolia Michx. [=Hedyotis nigricans (Lam.) Fosb|] 


Rafinesque included his new species Houstonia rupestris as the first species in 
his new subgenus Stenaria. He also included four other species now in 


594 BRIT.ORG/SIDA 19(3) 


Houstonia. Three (H. heterophylla Raf., H. oblongifolia Raf., and H. obtusifolia 
Raf.) of these are nomina dubia, and the fourth is H. tenuifolia Nutt. (=H. 
longifolia Gaertn. in Terrell 1996a). Steudel (Nom. Bot. ed 2, 1:776. 1840) listed 
Stenaria (asa nomen nudum) in smaller letters under Houstonia, consequently, 
he apparently intended Stenaria asa taxon ranked below Houstonia or else asa 
synonym of Houstonia. 

Perennial herbs or low shrubs with or without woody tap root. Stems 2-62 
cm tall, often from woody base, stout to slender, erect, spreading, or prostrate, 
glabrous to pubescent. Leaves 2-40 mm long, 0.3-8 mm wide, sessile or on short 
petioles, filiform or linear to ovate-lanceolate, elliptic, or oblanceolate, glabrous 
to pubescent. Stipules 0.5-3.5 mm long and wide, scarious, ovate, deltate, or lan- 
ceolate, sometimes acuminate or caudate, 0-few toothed or entire. Inflorescence 
of few-many cymes, dense to diffuse, flowers heterostylous or apparently 
homostylous in S. umbratilis, sessile or pedicellate. Hypanthium (calyx cup) 
hemispherical or cup-shaped, glabrous to hirsute. Calyx lobes 0.5-3 mm long, 
0.2-1 mm wide, narrowly lanceolate to ovate-lanceolate, margins glabrous to 
ciliate, apices obtuse, acute, or mucronate. Corollas 2-8.5 mm long, salverform 
or funnelform, white, purple, pinkish, or lavender; tube 1.5-5.5 mm long, 0.4-1 
mm wide at base, 0.5-2.5 mm wide at throat, densely puberulent or pubescent 
within; lobes 4(-5), 1-4.5 mm long, 0.5-2 mm wide, lanceolate or ovate, densely 
pubescent or puberulent on inner faces, apices glabrous or hairy. Pin form with 
stigmas subglobose, broadly elliptic,or oblong, their lobes 0.2-1.3 mm long, 
pubescent, elliptic or linear, exserted 0.5 to 3.5 mm beyond corolla throat; an- 
thers 4(-5), 0.8-1.8 mm long, linear or narrowly oblong, inserted at midpoint or 
at distal end of corolla tube or with tips exserted at corolla throat. Thrum form 
with anthers 0.6-2 mm long, narrowly oblong or linear, usually blue or white, 
straight or curved, on slender or flattened filaments ca. 0.5-2.5 mm long, lo- 
cated at mouth of corolla tube or somewhat exserted; stigmas ca. 0.3-0.8 mm 
long, included, extended to near midpoint or to distal end of corolla tube. Ma- 
ture capsules l-4.5 mm long, l-2.7 mm wide, 1/2-3/4(-4/5) inferior, dehiscing 
loculicidally, somewhat compressed, turbinate, obovoid, ellipsoid, or subglobose, 
glabrous to hirsutulous. Seeds several to 26 or more per capsule, 0.4-1.15 mm 
long, 0.3-0.6 mm wide, black or brown, often shiny, slightly to somewhat 
dorsilaterally compressed, non-crateriform, ellipsoid, in outline elliptical, ob- 
long, or slipper-shaped, ventral face with punctiform centric hilum on flat, 
slightly convex, or slightly ridged surface, testa finely reticulate. Chromosome 
numbers: x=9, 10 for S. nigricans; other species unknown. 


4 


ay 


KEY TO SPECIES 
1, Plants creeping, herbaceous, not woody or stems slightly woody toward base; 
Mexico, rare and disjunct. 5.S.umbratilis 
1. Plants not creeping, stems or bases woody. 


TERRELL, TAXONOMY OF STENARIA 595 


2. Plants shrublets or woody herbs, densely matted, prostrate, leaves overlapping, 

Coahuila, Mexico and Brewster Co., Texas. 2.S.mullerae 
2. Plants not shrublets, woody at base or on lower stems, not densely matted, erect 

or spreading, leaves not or only ag overlapping. 
and diffi) | 


3. Inflorescence ery small or lac 
ing, ere: oe ics Guttepuicise 
3. Inflorescence not ae or if somewhat diffuse then having apparent leaves 
in inflorescence. 
4. Stem leaves narrowly lanceolate to ovate-lanceolate or somewhat elliptic, 
leathery, cuspidate, margins scabrid or ciliate; capsules subglobose; south- 
western Texas and northern Coahuila. 4.S.rupicola 
4. Stem leaves various, not leathery, not cuspidate, margins not scabrid or 
ciliate; capsules longer than wide or subglobose only in S. nigricans var. 
floridana. 3.S. nigricans 


1. Stenaria butterwickiae (Terrell) Terrell, comb. nov. (Fig. 2) Houstonia butter- 
wickiae Terrell, Brittonia 31:164. 1979. Hedyotis butterwickiae (Terrell) Nesom, Syst. 
Bot. 13:434. 1988. TYPE: U.S.A. TEXAS: Brewster Co.: numerous in shallow pockets or 
crevices of limestone bedrock along ridgetop of Bullis Range, Bullis Gap Ranch, 
ca. 20 miS of Sanderson, 2-3 mi NW of Rio Grande River, 29° 47' 30" N, 102° 32' 30" 
W, assoc. with Penstemon baccharifolius, Phyllanthus ericoides, Polygala 
maravillasensis, Cirsium turneri, 27 Aug 1977, M. Butterwick, E. Lott,& S. Kennedy 
3893 (HOLOTYPE: US! ISOTYPES: GH! LL! MICH! MO! NY!). PARATYPE: Type locality, 16 
May 1977, M. Butterwick & E. Lott 3588 (LL). 


Perennial herb with a thick woody tap root and woody crown. Stems to 25cm 
tall, often numerous, slender, wiry, glabrous, diffusely branched from upper nodes, 
the internodes longer than leaves, the nodes often tinged black. Basal leaves to 
7 mm long, to 1 mm wide, somewhat clustered, sessile, linear, shorter, slightly 
wider, and thicker than cauline ones, glabrous; cauline leaves toca. 20 mm BONS 
0.5-1 mm wide, sessile, rather rigid, stiffly erect, filiform or linear, z at 
apex, glabrous or scaberulous, the midrib thick, the margin revolute. Stipules to 
1 mm long and wide, scarious, whitish, subtriangular, acuminate or truncate. 
Inflorescence with small, few-flowered cymes, the pedicels to ca. 1 cm long, fili- 
form, the inflorescence very diffuse, leaves lacking or greatly reduced, flowers 
heterostylous. Hypanthium glabrous; calyx lobes 0.8-3 mm long, ca. 1/2-3/4 
as long as corolla tube, linear-lanceolate, with thick midribs, sharply acute or 
acuminate, stiffly erect. Corolla 2.3-4.7 mm long, funnelform, white with sev- 
eral dark nerves, glabrous externally; tube 13-3 mm long, gradually widened 
distally, densely puberulent within; lobes 1-17 mm long, 0.5-1 mm wide, ovate- 
lanceolate, puberulent within. Pin flowers with stigma lobes 0.3-0.5 mm long, 
exserted 0.5-1.5 mm beyond corolla throat, anthers 0.8-1 mm long, included, 
attached near midpoint or 3/4-point of corolla tube. Thrum flowers with an- 
thers ca. | mm long, whitish, subsessile at sinuses of corolla lobes, stigmas in- 
cluded in tube. Mature capsules 1.5-2 mm long, equally wide, 1/2-3/4 inferior, 
dehiscing loculicidally, slightly or somewhat compressed, subglobose, glabrous, 
much exceeded by erect calyx lobes. Seeds ca. 10-26 per capsule, 0.4-0.7 mm 


596 BRIT.ORG/SIDA 19(3) 


Fig. 2. Stenaria butterwickiae. Drawing of holotype reprinted from Brittonia 31:64. 1979. A. Habit, x .45. B. Detail of 
inflorescence, pin form, < 7.C. Capsule, x 5.5. D. Corolla with anthers, pin form,  5.5.E. Style and stigma, 5.5. 


long, 0.3-0.4 mm wide, black or brown, ellipsoid, in outline elliptic or oblong, 
slightly or somewhat compressed, the hilum punctiform on ventral face, testa 
finely reticulate. Chromosome number unknown. 

Phenology.—Flowering August. 

Distribution —Brewster Co., Texas. Known only from type collection. 


TERRELL, TAXONOMY OF STENARIA 597 


Stenaria butterwickiae is distinctive in its very diffuse inflorescences with- 
out leaves or with leaves bract-like. The cauline leaves are small and stiffly erect. 
These characters are evident in Fig. 2. 


2. Stenaria mullerae (Fosb.) Terrell, comb. nov. Hedyotis mullerae Fosb., Lloydia 4:288. 
1941. Houstonia mullerae (Fosb.) Terrell, Brittonia 31:169. 1979. Type: MEXICO. 
COAHUILA: Mpio. de Cuatro Cienegas, Canon del Agua, Sierra de la Madera, 9 Sep 
1939, C.H. Muller 3234 (HOLOTYPE: US!, where transferred from NA; ISOTYPE: GH). 

Shrublets or perennial herbs, roots woody or with woody rhizomes to 6 mm 
thick. Stems to ca. 10 cm tall, often prostrate, matted, woody toward base, tet- 
ragonal, often obscured by the numerous small leaves, glabrous, internodes 1-5 
mm long. Leaves 2-8(-10) mm long, 0.7-2 mm wide, sessile, thick, often gray- 
green, sometimes shiny, lanceolate, elliptic, narrowly elliptic, or oblanceolate, 
hispidulous to glabrous above, glabrous or with midrib hairs beneath, midribs 
often thickened and conspicuous below, margins thickened and revolute, con- 
spicuously ciliate to scabrous, apices obtuse, acute, or apiculate. Stipules to ca. 
2 mm long, adnate basally, deltate, margin often ciliate, apices 0.2-0.4(-2) mm 
long, acute, acuminate or apiculate. Inflorescence with flowers heterostylous, 
axillary, usvanya in groups “i 3-5, subsessile or on pedicels to 1 mm long. Hy- 
panthium hi ;calyx lobes 1-2.5 mm long, lanceolate, oblan- 
ceolate, or ae sometimes incurved, ciliate to glabrous, apices obtuse or 
acute. Corollas 2.5-6(-8) mm long funnelform, white, “sometimes tinged rose,” 
hirtellous within distally; tube 2-4(-5) mm long, 0.7-1.2 mm wide at throat; 
lobes 1-3 mm long, I-1.5 mm wide, ovate or lanceolate, densely pubescent to 
glabrous on inner faces, apices sometimes with hairs. Pin flowers with stigma 
lobes 0.2-0.4 mm long, broadly elliptic to linear, exserted on filiform styles to 2 
mm beyond throat, anthers 0.8-1.4 mm long, sessile, partly exserted at throat. 
Thrum flowers with anthers 0.7-1.5 mm long, linear or oblong, “greenish blue,” 
attached near or at sinuses, exserted on filaments 0.5-1 mm long, stigma in- 
cluded, styles to 2.5 mm long, glabrous. Mature capsules 1-2.2 mm long and 
wide, |/2-4/5 inferior, compressed-subglobose, dehiscing loculicidally, apices 
broadly rounded or retuse. Seeds 0.6-0.8 mm long, 0.3-0.5 mm wide, black, 
rather shiny, somewhat compressed, in outline elliptic or angular-elliptic, hi- 
lum punctiform, on flat or slightly concave ventral surface, testa finely reticu- 
late. Chromosome number unknown. 


KEY TO VARIETIES OF S. MULLERAE 


1. | | late, ta gradually t p point;stipules cuspidate with as 
1-2 mm long. b.var.pooleana 
1. Leaves elliptic, ly elliptic, oblanceolate, or narrowly oblanceolate, obtuse, acute, 


or apiculate; stipules abruptly pointed, apiculate, or scarcely pointed, apical points 
0.2-0.4 mm long. 2a. var. mullerae 


598 BRIT.ORG/SIDA 19(3) 


2a. Stenaria mullerae var. mullerae (Fig. 3). 


Leaves 2- B(- 10) mm long, usually gray- green, elliptic, nar cone SMpHG or cen 
hispidulous to glabrous above, 
apices obtuse, acute, or apiculate; stipules with apices scarcely ae to 
abruptly short-pointed (more or less apiculate), apical points 0.2-0.4 mm long. 

Phenology.—Flowering May to September, 

Distribution —Mexico. central (including Cuatro Cienegas Basin) and western 
Coahuila; rock crevices, steep slopes, ridge crests, cliff walls, usually on limestone, 
1250-3000 m. 


Selected representative specimens examined: MEXICO. Coahuila: ca. 35 km W of Cuatro vias 
n Canyon de la Hacienda, Sierra de la Madera, near 27° 3' N, 102° 24' W, 7300 ft, Henri kson & Wendt 
11889 (ASU); ca. 31 air mi WNW of Cuatro Cienegas, Sierra de la Madera, S of Canyon Desiderio, 27° 

06' N, 102° 32'W, 2700-2800 m, Henrickson & Prigge 15275 (LL); SW end of Sierra de la Fraqua, 1-2 km 

N of Puerto Colorado, M Johnston 8741 (GH, LL);W side of Potrero de la Mula, about 20 km NW of 

came: elas as 5A (GH); Sierra San Marcos, N part jutting into Cuatro Cienegas basin, 26° 

26° 51'N, 102° 102° 07' W, 1250-1800 m, M.C.Johnston et al. 10925 (LL, TEX, US); Mina El 
painere ‘ side of ee de la Paila, 26° 05' 30"— 06' N, 101° 36' W, 1700-2200 m,M.C Johnston et al. 

11692 (LL); Sierra de San Marcos, opposite Los Fresnos, Pinkava et al. P-6011 (ASU, ENCB, LL, NY). 


ceolate 


a 


Stenaria mullerae is variable in leaf size and shape, leaf vestiture, and compact- 
ness. Certain collections are somewhat intergradient with S. nigricans, suggesting 


possible hybridization. 


2b. Stenaria mullerae var. pooleana (B.L. Turner) Terrell, comb. et stat. nov, 
Hedyotis pooleana B.L. Turner, Phytologia 79:93. 1995. TyPE: U.S.A. TEXAS. Brewster 
Co. Brushy Canyon Preserve (Nature eau of Texas), Dead Horse Moun- 
tains (Sierra del Caballo Muer . ca. 0.7 mi SW of adobe house on Brushy pies 
divide 29° 27' N, 102° 58' W), 4900 [t, N to E-facing vertical limestone cliff 
with Agave lecheguilla, Rhus vire a. rcus pungens, | May 1987, Jackie Poole ro 
(HOLOTYPE: LE 

Leaves usually 5-7 mm long, lanceolate, bright green, glabrous, scabrous on 

margins, at apex tapering gradually intoa long sharp point (more or less cuspi- 

date). Stipules with long narrow caudae 1-2 mm long, sometimes irregularly 

toothed. 

Phenology.—Flowering May. 

Distribution.—United States. Texas, Brewster County. Dead Horse Moun- 
tains (Sierra del Caballo Muerto), 29° 27' N, 102° 58’ W. Known only from type 
collection. Also seen by the collector (but not collected) about one mile from 
the type locality on 27 November 1987, along the same ridge at about 4840 ft 
elevation within the boundary of the Big Bend National Park. 

Additional collection: Type locality, 25 May 1985, Poole 2527 (SRSC, TEX!). 

Turner (1995b) in the abstract of his paper described H. pooleana as closely re- 
lated to H. mullerae, but differing in having lanceolate, markedly thickened 
leaves with glabrous surfaces and apiculate apices. In addition, he provided 


TERRELL, TAXONOMY OF STENARIA 599 


[Metric 
AE 


Fic. 3.5 j Hl mullerae 


D £halnt Hic) 
vt 


measurements of other organs. Terrell (1996b) advocated sinking of H. pooleana 
into H. mullerae, based on previous study of the considerably variable H. 
mullerae in loans from ASU, ENCB, GH, NY, TEX, and US. Turner (1997) de- 
fended specific status. Recently, | have re-examined specimens and types from 
US and TEX (on current loan) and I now advocate varietal status for H. pooleana. 

There are two clear differences between the two taxa: in leaf shape and 
stipule shape. Hedyotis pooleana has lanceolate leaves which taper gradually 
to their tips, whereas in H. mullerae the leaves vary from elliptic or oblanceolate 
to narrowly elliptic or narrowly oblanceolate, with the apices varying obtuse, 
acute, or apiculate. (That Turner attributed apiculate apices to H. pooleana ap- 
pears to have been a lapse.) I accept these terms more or less as pictured by 
Lawrence (1951, p. 744, Fig. 307). (Also, the leaves of H. mullerae range from 
densely pubescent to glabrous). Turner did not mention stipular differences, 
however, these are at least as well marked as the leaf characters: var. mullerae 
has stipules with very short, abruptly narrowed apices, whereas var. pooleana 
has stipules more or less cuspidate with narrow caudae 1-2 mm long. 

I believe that these differences in vegetative characters represent varietal, 
rather than specific differences in the variable S. mullerae. The two varieties 
are about 150 miles apart. Turner (1995b) stated that the Dead Horse Mountains 
(Brewster Co., Texas) are an extension of the Sierra del Carmen range “across 
the Rio Grande in Coahuila.” Thus, it seems possible that the two varieties may 


600 BRIT.ORG/SIDA 19(3 


have come from similar original stock, but have been isolated from each other 
for many millenia. 
3. Ss nigricans (Lam.) Terrell, comb. nov. Gentiana nigricans Lam., Enc 

64 


ycl. 
5. 1788. Houstonia nigricans (Lam.) Fern., Rhodora 42:299. 1940. Hedyoris 
P; photo, 


nigr icans(Lam.) Fosb., Lloydia 4:287. 1941. Type: Herb. Jussieu (HOLOTYPE: 

Fernald 1940) 
Perennial herb with woody tap root toca. 1.5cm thick, foliage sometimes black- 
ened when dried. Stems to 6.2 dm tall, often many-stemmed from broad woody 
base, stout to slender, erect, spreading, or decumbent, tetragonal and slightly 
winged, glabrous, scabridulous, papillose or densely puberulent or pubescent 
with gland-tipped hairs, occasionally with localized groups of densely aggre- 
gated white hairs. Leaves 7-40 mm long, 0.3-5(-8) mm wide, often fasciculate, 
sessile or tapering to short petioles, strongly revolute, filiform, linear, narrowly 
elliptic, elliptic, narrowly lanceolate, or narrowly oblanceolate, acute or obtusish, 
glabrous, scabridulous, densely puberulent or pubescent. Stipules to ca. 3.5mm 
long and wide, scarious, deltate, ovate or lanceolate, sometimes acuminate or 
caudate, O-few toothed. Inflorescence of few-many dense to loose cymes, often 
many-flowered (sometimes hundreds), flowers heterostylous, sometimes central 
or oldest flowers sessile or subsessile, other flowers on pedicels to ca. lO mm 
long. Hypanthium glabrous to hirsute; calyx lobes 0.5-3(-4.4) mm long, 0.2-1 
mm wide, slightly longer to slightly shorter than capsules, lanceolate or trian- 
gular-lanceolate, glabrous to hirsute, margins glabrous to ciliate, apices acute 
or mucronate. Corollas (2-)3.5-8.5 mm long, salverform to funnelform, quite 
variable in shape and size, white, light purple, pink, or lavender, buds elliptic, 
becoming obovate or oblanceolate and tapering to truncate or obtuse apices 
with or without hairs; tube (1.5-)2-4(-5.5) mm long, 0.4-1 mm wide at base, 
0.5-2.5mm wide at throat, often densely puberulent to hirsutulous within, tube 
length/lobe length ratio usually 11 to 2:1; lobes 4(-5), 1-3.5(-4.5) mm long, 0.5-2 
mm wide, lanceolate or ovate, usually densely puberulent, pubescent, or hir- 
sutulous on inner faces with white often gland-tipped hairs to ca. 0.6 mm long, 
apices also sometimes hairy. Pin form with stigmas bilobed, 0.3-1(-L3) mm long, 
subglobose or broadly elliptic, papillose or pubescent, oblong, elliptic or linear, 
exserted to 3.5mm beyond corolla throat on styles 4.5-5 mm long, anthers 4(-5), 
0.8-1.8 mm long, linear or narrowly oblong, included at distal end of corolla 
tube or with tips exserted at corolla throat. Thrum form with 4(-5) anthers, 
0.6-2.0 mm long, narrowly oblong or linear, usually blue or white, straight or 
curved, exserted on slender or flattened filaments ca. 0.5-2.5 mm long, attached 
near ends of anthers, stigmas ca. 0.3-0.8 mm long, included, linear, extended to 
near 2/3-point or to distal end of corolla tube on styles 1.5-2.5 mm long. Mature 
capsules 1.5-4.5 mm long, 1-2.7 mm wide, (1-)1.3-2 times longer than wide, (1/2-) 
3/4(-4/5) inferior, somewhat compressed, turbinate, obovoid or broadly ellip- 


— 


TERRELL, TAXONOMY OF STENARIA 601 


soid, quite variable in shape and size on an individual plant, glabrous or scabrous 
to hirsutulous, apices retuse, truncate, or broadly rounded, dehiscing widely 
loculicidally, then septicidally through septum, eventually forming two or four 
terminal segments. Seeds 0.45-1.15 mm long, 0.3-0.6 mm wide, black or dark 
brown, often shiny, somewhat compressed, ellipsoid, in outline elliptical, oblong, 
or slipper-shaped, ventral face with punctiform hilum more or less centered on 
flat, slightly concave, or slightly ridged surface, testa finely reticulate. 


KEY TO VARIETIES OF S. NIGRICANS 


. Leaves more or less elliptic, 2.5-3.5(-4) times longer than wide; corollas eae 4- 


6 mm long; Mexice and southwest Texas d. var. gypsophila 
. Leaves usually fi n, linear, narrowly lanceolate, or oblanceolate, more Tee ee 

eer nice ae 2-8 mm long. 

2. Capsules subglobose, 1.2-2.5 mm long, 1.2-2.2 mm wide, 1-1.3 times longer 


than wide; leaves 0.2-1.2 mm wide; corollas 3—5(-6) mm paeeounen Florida, 

Bahamas. 3c. var. floridana 
2. Capsules longer than wide, 1.5-4.5 mm long, 1-2.7 mm wide, 1.3-2 times longer 

than wide; leaves 0.3-8 mm wide; corollas 2-8 mm long. 

3. Corollas 2-4(-4.5) mm long; leaves 0.7—2(-3.2) mm wide; Mexico. ___ 3b. var. breviflora 

3. Corollas (2.5-)3-8 mm long; leaves 0.3-8 mm wide; wide-ranging. ____ 3a. var.nigricans 


W 
+) 


. Stenaria nigricans var. nigricans (Fig. 4). Houstonia angustifolia Michx., Fl. Bor- 
Amer. 1:85. 1803. Non Hedyotis angustifolia Cham. & Schlecht., 1829. Oldenlandia 
angustifolia (Michx.) A. Gray, Pl. Wright. 2:68. 1853. Chamisme i aaa ia (Michx.) 
NieuwL, Amer. re Natitalst 492.1915. Hedyotis stenophylla Torr. & A. Gray, FI. 
N. Amer. 2:41.1841.(Houstonia an gustifolia listed as synonym). ee 
Nutt. ex A. Gray, ‘ Wright. L81. 1852, as syn. of H. stenophylla Torr. & A. Gray, 
invalid name. (Specimens labelled H. lasiantha in BM, K, PH). TYPE: U.S.A. FLORIDA: 


“submaritimis Floridae” (HOLOTYPE: P!). 


— 


Houstonia rupestris Raf., Ann. Gen. Sci. Phys. 5:226.1820. Non Hedyotis rupestris Swartz, 

1797. TyPE: US.A. KENTUCKY: rocks borde ring Kentucky River, specimen lost? 

Oldenlandia angustifolia (Michx.) A. Gray var. filifolia Chapm.,, Fl. S. US. 181. 1860. 
Houstonia angustifolia Michx. var. filifolia (Chapm.) A. Gray, Syn. Fl. N. Amer. 
1(2):27. 1884. Houstonia filifolia (Chapm.) Small, Fl. S.E. U.S. 1109, 1338. 1903, as 
“(A. Gray) Small.” Hedyotis nigricans var. filifolia (Chapm.) Shinners, Field & Lab. 
17:168. 1949. TyPE: U.S.A. “S. FLORIDA,” Chapman s.n. (LECTOTYPE: US-83375! desig- 
nated by Terrell 1986). 

Houstonia angustifolia var. scabra S. Watson, Proc. Amer. Acad. Arts 18:97. 1883. 
Hedyotis nigricans var. scabra (S. Watson) Fosberg, Lloydia 4:288. 1941. TyPE: 
MEXICO. COAHUILA: Caracol Mountains 21 miles southeast of Monclova, 19-22 Aug 
1880, Palmer 410 (LECTOTYPE, here designated, GH}; isotypes, LL! NY! PH! US! VT). 
Isotype at LL designated by Turner 1995a 

Houstonia angustifolia var. rigidiuscula A. Gray, Syn. Fl. N. Amer. 1(2):27. 1884. 
Houstonia rigidiuscula (A. Gray) Wooton & Standley, Contr. U.S. Natl. Herb. 16:175. 
1913. Hedyotis nigricans var. ne gTeTSCUNG (A. Gray) Shinners, Field @& Lab. 17:168. 
ae SYNTYPES: “S. and W. Texas, Palmer, Havard, & c. Coast of E. Florida, Rugel. 

x.).” Two sheets in GH include a poselbic lectotype. 
Westend salina A. Heller, Contr. Herb. Franklin and Marshall Coll. 1:96, pl. 9. 1895. 


BRIT.ORG/SIDA 19(3) 


aha ae fey say RG - 
Me ht WME ALA “f, A - 
ty m*” MIKE, Ihe pe “yt HY, , 
Peas eg ten: Be) hale: AY 
MAC my t A 


> 
te twa) 
Pia ei a lb 

om dal 


Fig. 4 


aris herbarium, lef 


to shia similarity with type. a 625 from Fernald 1940. 


TERRELL, TAXONOMY OF STENARIA 603 


Hedyotis salina (A. A. Heller) Shinners, Field & Lab. 17:169. 1949. Hedyotis nigricans 
f. salina (A. Heller) WH. Lewis, Rhodora 63:222. 1961. TYPE: U.S.A. TEXAS. Nueces 
Co.: shell deposit along beach, Corpus Christi, 31 May 1894, A. A. Heller 1812 (LEC- 
TOTYPE designated by Terrell 1986, GHI; IsoTYPES, BM! ILL! K! MO! NY! PH! US-3). 

Houstonia pulvinata Small, Bull. New York Bot. Gard. 1:289. 1899. Hedyotis nigricans 
var. pulvinata (Small) Fosb., Castanea 19:37. 1954. Houstonia nigricans var. 
pulvinata (Small) Terrell, Phytologia 59:79. 1985. TyPE: U.S.A. FLORIDA. St. Johns 
Co.:sandy soil, St. Augustine, Jul 1876, Mary C. Reynolds s.n. (LECTOTYPE designated 
by Terrell 1986: NY! Ses NAI). PARALECTOTYPES, same locality and date, 
A. P. Garber s.n. (NY! US-2!). 

Houstonia tenuis Small, FL S.E. US. 1109, 1338. 1903. TyPE: U.S.A. TEXAS. San Saba Co.: 
San Saba, Oct 1850 (?), Thurber 67 (HOLOTYPE: NY)). 

Hedyotis nigricans var. austrotexana B.L. Turner, Phytologia 79:15. 1995. TyPE: U.S.A. 
TEXAS: Karnes Co: roadside 2 mi EF of El Tejano Cafe, dry sandy clay soil, 22 June 
1952, Joe C_ Johnson 833 (HOLOTYPE: LL). 

Hedyotis nigricans var. papillacea B.L. Turner, Phytologia 79:15. 1995. TYPE: U.S.A. NEW 
MEXICO. Otero Co. northern McKittrick Canyon at first crossing of Texas-New 
Mexico boundary, gravels and boulders of stream bottom, 8 Oct 1973, Thomas F. 
Patterson 508 (HOLOTYPE: LL). 


Leaves 0.3-8 mm wide, filiform, linear, narrowly lanceolate, or narrowly ob- 
long, and more than 4 times longer than wide; corollas (2-5-)3-8 mm long; cap- 
sules 1.5-4.5 mm long, 1-2.7 mm wide, 1.3-2 times longer than wide. 

Chromosome numbers.—Lewis (1959) reported 2n=18 for var. nigricans from 
Nacogdoches Co., TX. Lewis (1962) listed n=9 for forma salina (San Patricio Co., 
TX), n=9 and 10 for var. nigricans (Chaves Co., NM and Brewster Co., TX) and 
n=10 for var. rigidiuscula (Brewster Co., TX). Lewis in Terrell et al.1986) counted 
n=10 and 2n=20 for var. pulvinata (St. Johns Co., FL). These varieties and the 
forma are here sunk under var. nigricans. 

Common name.—diamond flowers. 

Phenology.—Flowering usually April to December in Mexico; in U.S.A. usu- 
ally April to October or all year in Florida. 

Distribution —United States. Southern Michigan (Branch Co.) and Ohio 
south to Florida, west to lowa, Nebraska, eastern Colorado,and New Mexico; Mexico: 
Sonora, Sie salute Nuevo Leon, a San Luis Potosi, and Hidalgo. 


Selected ined: MEXICO. Ct I de, 3 km E of Rancho 
E| Niuicielage 29° 52'N, 104° 50' W,M.C Johnston et al. 11293 (CAS, TEX). Coahuila: 8 km S of El Tule, 
24 km N of Castillon, Stewart 447 (CAS, GH, MO). Nuevo Leon: 7 mi W of Iturbide along hwy 60, 
McGregor et al. 32 (KANU, NY, SMU, US). 

UNITED STATES. ALABAMA. Dallas Co.: roadbank 9 mi W of Selma, Webster & Wilbur 3508 
cs MICH, NCSC). ARKANSAS. Clark Co. chee Demaree 15643 (FLAS, NY, TENN). Garland 
Co.: Hot Springs, Harvey 12824 (IA, KANU, MICH). COLORADO. Yuma Co.: 0.75 mi $ of Laird, McGregor 
32365 Teun FLORIDA. Franklin Co.: 6 mi NE of iene Point, Kral 2800 (FLAS, FSU). GEORGIA. 
Decatur Co.: along Flint River 1 mi N of Chattahoochee, Florida, Thorne 5931 (GA, IA, MICH, MT). 
ILLINOIS. Jersey Co.: Pere Marquette State Park, Evers 25019 (ILLS). INDIANA. Harrison Co.: 1 1/2 
mi NE of Davidson, Deam 37238 (IND). IOWA. Fremont Co.: Hamburg, Shimek s.n., 31 Aug 1898, (IA). 
KANSAS. Montgomery Co.: 3 mi S of Independence, Horr £256 (ASU, CM, FLAS, ISC, KANU, MT, 


604 BRIT.ORG/SIDA 19(3) 


NCSC, NO). KENTUCKY. Wayne Co.: Beaver Creek SW of Monticello, Smith & Hodgdon 3907 (GH). 
LOUISIANA. Caddo Parish: 3 mi S of La.526 at Forbing, Thomas & Dorris 96086 (KANU, WNC). MICHI- 
GAN. Branch Co.: loc. not given, Mrs. JM. Mulligan s.n., Jun 1889 (US). MISSOURI. Franklin Co.: 3 mi 
S of Grays Summit, Steyermark 607 (MO). NEBRASKA. Cass Co.: Weeping Water, Sheldon s.n., 17 Aug 
1898 pay MEXICO. Lincoln Co.: 15 mi E of El Capitan, 8500 ft, Hitchcock et al. 4239 (ISC, NO). 
OHIO. Ottawa Co.: Rattlesnake Island, Moseley s.n., 29 Jul 1895 (CM, F, MICH, MO). OKLAHOMA. 
McClain mi E of Blanchard, (PI. Exs. Grayanae 686), Demaree s.n., 30 Jun 1936 (DUKE, GA, A 7 
KANU, MICH, MT, NCSC, NO, PAC, TRT, WIS, WVA). TEXAS. Kerr Co.: Kerrville, Heller 16617 (ARIZ,IA, MICH). 


The preceding citation of specimens lists one specimen from most states of the 
United States where the typical variety occurs. Only a few Mexican specimens 
were cited because of lack of detailed notes on morphology. Turner (1995a) pro- 
vided a map of the Texas distribution. 

Previous to 1940 the name Houstonia angustifolia Michx. was usually ap- 
plied to the species later known as Hedyotis nigricans. Fernald (1940, plate 625) 
presented an illustration of the type of Gentiana nigricans Lamarck from the 
Paris herbarium (shown here as Fig. +). Fernald’s plate shows two plants of the 
type collection along with a plant from Georgia collected by Boykin near the 
type region of Houstonia angustifolia. The three plants are very similar, so much 
so that Fernald commented “That they are the same no one is likely to ques- 
tion.” The plants of the type collection are tall, rather strict, linear-leaved, and 
generally similar to other plants from the southeastern United States. 

Stenarid nigricans isa polymorphic species. Plants from Adams County in 
southern Ohio are short and decumbent; plants from the southeastern U.S.A. 
often are tall and have very narrow leaves; plants from Kansas and Nebraska 
are often very stout and large, with thick, broad, woody bases and heavy tap- 
roots. Plants may also differ conspicuously from one population to another in 
size of corollas. This is only a small example of the variation, much of which 
seems to occur at random. | have not studied S. nigricans in detail, and leave it 
to future workers to study the species in depth. 

Certain varieties are here placed in synonymy. Variety scabra S. Watson is 
only a hairier-than-average extreme connected with the typical plants by a 
continuum. Variety rigidiuscula A. Gray is a confused and ambiguous name 
whose protologue seems to refer to more or less typical plants of S. nigricans: 
the specimens cited as syntypes came from Texas and Florida and are rather 
diverse. Turner (1995a) came to a similar conclusion about this variety. 

Houstonia salina and H. tenuis were discussed previously by Terrell (1986). 
| concluded that they did not merit varietal status. Turner (1995a) agreed with 
this conclusion. 

Terrell (1986) compared var. pulvinata with var. nigricans,and maintained 
the former as a variety. Further consideration, however, leads me to conclude 
that its differences overlap so strongly with those of var. nigricans that it should 


be in synonymy. 
Turner’s (1995a) var. austrotexana and var. papillacea appear to differ from 


TERRELL, TAXONOMY OF STENARIA 605 


var. nigricans in rather minor ways, considering the great amount of variation 
in S. nigricans, e.g., var. dustrotexana: “Resembling var. nigricans but the more 
mature calyces markedly papillose throughout.” Variety papillaced is also pap- 
illose and appears to be delimited by the Texas border. examined specimens of 
both varieties lent tome from TEX and specimens available in BRIT, but Lcould 
not accept their being sufficiently distinct to recognize as varieties (Terrell 
1996b). Turner (1997) again upheld these varieties. Stenaria nigricans is an ex- 
ceedingly variable entity, and I find it somewhat premature to recognize vari- 
eties on minor vestiture characters in Texas without studying the species in 
Oklahoma and throughout its range. 


3b. Stenaria nigricans var. breviflora Terrell, var. nov. Type: MEXICO. NUEVO LEON: 
Sierra Madre near Monterrey, C.G. Pringle 13878 (HOLOTYPE: US! ISOTYPES: CAS! F! 
GH! MEXU! MICH! MO! MSC! SMU! VT). 

Folia saepe 0.7-2(-3.2) mm lata, linearia vel filiformia; corollae 2-4(-4.5) mm longae; capsulae 1.8- 

3(-3.8) mm longae. 1.3-2.2 mm latae, 1-1.5(—2) plo longiores quam latiores. 

Leaves usually 0.7-2(-3.2) mm wide, linear or filiform; corollas 2-4(-4.5) mm 

long; capsules usually 1.8-3(-3.8) mm long, 1.3-2.2 mm wide, 1-1.5(-2) times 

longer than wide, often glabrous. 

Distribution.—Mexico: Nuevo Leon, vicinity of Monterrey, and in adjacent 
Coahuila. I have 30 records of this variety from 15 herbaria. The habitats are 
variously: limestone (travertine) and talc-like soil on gravelly hill; rocky lime- 
stone valley; sandy loam; dry pine-oak forest; crevices of limestone rock; gyp- 
sum plug (of Portrero Chico); bottom of arroyo. 

Selected representative specimens examined: MEXICO. Coahuila: Ojo Caliente, 33 mi SW of 
Monterrey, Warnock & Barkley 14744M (ENCB, F, GH, MO, NY, TEX). Nuevo Leon: 5 km S of Sabinas 
Hidalgo on road to Monterrey, Frye & Frye 2408 (GH, NY, US); 16 mi SW of Villa Santa Catarina, Hernandez 
Cet al. 16M516 (TEX); Diente Canyon, Monterrey, Muller & Muller 111 (F,GH, MEXU, TEX); toll road up 
Sierra Anahuac, 4.5 mi SE of Monterrey, Oliver 202, (SMU, TEX, US); 25 km NW of Monterrey, ane 

Chico, Powell & Turner 2335 (TEX); between Linares and Galeana, 2500 ft, Sharp 45653 (GH, TENN, US). 


The main distinguishing character of this variety is the consistently short co- 
rollas. Also, the leaves are short and linear or filiform. 


3c. Stenaria nigricans var. floridana (Standl.) Terrell, comb. nov. Houstonia 
floridana StandL, N. Amer. FI. 32(1):36. 1918. Hedyotis purpurea var. floridana 
(Standl.) Fosb., Castanea 19:36. 1954. Houstonia nigricans var. floridana (Standl.) 
Terrell, Phytologia 59:79. 1985. Hedyotis nigricans var. Ue ale (Standl.) 
Wunderlin, Sida 11:400. 1986. Type: U.S.A. FLORIDA. Dade Cocoanut Grove, 
eee Bay, Jul 1895, A.H. Curtiss 5484 (HOLOTYPE: US! IsoTYPEs: FLAS! ISC! NY! 
US! VT). 


Stems usually 1-3.5 dm tall, sprawling, spreading, or erect, wiry and diffusely 
branched. Leaves filiform or linear, 0.2-1.2 mm wide. Corollas 3-5(-6) mm long, 
glabrate to densely pubescent within. Mature capsules subglobose, 1.2-2.5 mm 
long, 1.2-2.2 mm wide, 1-1.3 times longer than wide. Chromosome numbers: 


606 BRIT.ORG/SIDA 19(3 


Lewis (1962) reported n=9 for this variety (listed as var. filifolia) from Dade Co., 
Florida. Lewis in Terrell et al. 1986 reported n=9 and 2n=18 for another collec- 
tion from Dade Co., Florida. 

Distribution.—Southern Florida: over limestone at a number of locations 
in Dade County, and at Big Pine Key in Monroe County; Bahamas: Grand Bahama 
Island, Great Abaco Island. 


Selected representative cimens examined: BAHAMAS. Grand Bahama Island: Intersection of 
Settlers Way and Balao Buse, Freeport, Correll 440470 (NY); Buckingham-Leicester Co. line, Govern- 
ment Road, Lewis 718? (FSU, FTG, MO, NY). Great Abaco Island: halfway between Marsh Harbour 
and ae Harbour Airport, Wunderlin et al. 8564 (USF). 
TED STATES. FLORIDA. Dade Co.: Long Pine Key, Brass 32997 (USF); jct. Krome Ave., Moody 

Drive, a mi N of Homestead, $36, 1565S, R38E, Burch et al. 285 (FLAS); Pine Island, 25 mi S of Miami, E of 
Naranja and Homestead, Elder 289 (DUKE); 5 mi S of Florida City on US1,C.& J Janish 418 (CAS):6 mi 
E of Royal Palm Park, pi 7584 (ARIZ, CAS, CM, MICH, MT, NO, US); Sykes Hammock, everglade keys, 

Small, Mosier, & Small 6755 (CAS, DUKE, FLAS, MICH, US, WVA). Monroe Co.: Big Pine Key, Brumbach 
9392 (FSU); N of upper 2 i road, Big Pine Key, Killip 40896 (NO);4 mi N of US1, Big Pine Key, McDaniel 
5813 (FSU). 


Terrell (1986) discussed var. floridana and var filifolia in detail, and concluded 
that the latter variety was a synonym of var. nigricans, as its description re- 
ferred to plants unlike var. floridana. 
Variety floridand is distinguished by having subglobose capsules. Because 
of these capsules Fosberg named it as Hedyotis purpurea |=Houstonia purpurea] 
var. floridana, but it has non-crateriform seeds like those of var. nigricans and 
does not belong in Houstonia. 


3d. Stenaria nigricans val sypsophila (B.L. Turner) Terrell, comb. nov. Hedyotis 
nigh ICans Var. gypsophila B.L. Turner, Phytologia 79:15. 1995, TYPE: MEXICO. NUEVO 
LEON, Santa Rita, 2370 m, sparse pine woods, gypsum hillsides, 11 Jun 1981, Hinton 
et al. 18278 (HOLOTYPE: TEX). 


This new variety was described by Turner as follows: “Resem bling var. nigricans 
but the plants low and much-branched [rom the base, the leaves elliptic-ovate 
and mostly 2.5-3.5(-4) times as long as wide (vs. linear-lanceolate to linear- 
oblanceolate, mostly +-20 times as long as wide) and the mature calyces usually 
markedly hispid with broad-based hairs (vs. glabrous or merely minutely setose).” 

My description incorporates items from Turner’s description. 

Stems woody at base, 0.4-2.5 dm tall. Leaves 4-17 mm long, 1-4(-6) mm 
wide, 2.5-3.5(-4) times longer than wide, elliptic or narrowly elliptic. Corollas 
usually 4-6 mm long. Capsules usually 2.5-3.2 mm long, 1.5-2.2 mm wide. Seeds 
0.45-0.75 mm long, 0.3-0.6 mm wide. 

Chromosome number—Lewis (1962) listed n=9 under the name H. nigricans var 
angulata (=S. rupicola), but the specimen was actually S. nigricans var. gypsophila. 

Distribution —Mexico: Northern and southeastern Chihuahua and northern 
Coahuila south southeast to Nuevo Leon, northeast Zacatecas, southwest 


TERRELL, TAXONOMY OF STENARIA 607 


Tamaulipas, and San Luis Potosi. It has been much collected from the Saltillo 
and Galeana areas. The altitudes range from ca. 1500 to 3195 m. United States: 
Texas: Culberson, Hudspeth, and Jeff Davis counties. The specimens resemble 
Mexican collections, and this extends var. gypsophila northward into the United 
States. The range of habitats of the Mexican specimens includes the following: 
pine and oak forests; pinyon-juniper; pine-douglas fir; steep limestone slope; 
east-facing ravine; rocky slope; bank of dry arroyo; steep igneous slope; and 
gypsum flats in pine-pinyon 

The most conspicuous character is the elliptic leaves on rather small plants. 
As noted (Terrell 1996b) I was aware of this taxon prior to its description by 
Turner. Like Turner, I can not find any other characteristics significantly dif- 
ferent from var. nigricans (hairy mature calyces occur in var. nigricans). | con- 
sider the taxon as misnamed, as I recorded only one collection out of 49 Mexi- 
can collections from 19 herbaria that mentioned a gypsum substrate. It may be 
noted that the type specimen came from a gypsum habitat. 

Plants superficially somewhat resemble those of some variants of 
Houstonia longifolia, which occurs in the eastern and central U.S.A. (Terrell 
1996a). Variety gypsophila may also be confused with the Mexican species 
Houstonia wrightii. The latter species may occasionally occur at higher eleva- 
tions near var. gypsophila, and a very few collections have had both species on 
one sheet. The following comparison may provide a means of distinguishing 
them: capsules of var. gypsophila are somewhat compressed, longer than wide, 
and eventually dehisce widely into four similar segments; seeds are ellipsoid 
with the hilum more or less centered on the ventral side and punctiform; buds 
often have hairy apices. Houstonia wrightii capsules are subglobose, not or only 
slightly compressed, wider than long; seeds have a linear hilar ridge in a ven- 
tral depression; bud apices are glabrous (Terrell 1996a). 


Selected representative specimens examined: MEXICO. Chihuahua: Sierra de la Parra, across Rio 
Grande from Sierra Vieja, 1450-2158 m, 30° 00-02' N, 104° 52-53'W,M.C_ Johnston et al. 11314B (TEX, 
US). Coahuila: 22 km ESE of La Cuesta del Plomo on Muzquiz-Boquillas hwy near intersection of 
hwy from V. Acuna, 1000 m, 28° 38' N, 102° 18' W, Chiang et al. 7550R (CAS, TEX, US): about 35 mi E of 
Saltillo, 5—6 mi E of Los Lirios, 2300-2400 m, McVaugh 12338 (MICH); mpio. Villa Ocampo, Canyon de 
la Vaca, Sierra Santa Fe del Pino, Passini & Robert 5193 (ENCB); Sierra Guadelupe, S of La Cuchilla, 
2300-2600 m, Pennell 17388 (F, PH, US). Nuevo Leon: Canon del Voladero, Galeana, 2200 m, Hinton 
18744 (ASU, US); Cerro Potosi, near microwave tower, 9000 ft, McGregor et al. 351 (TEX, US); mpio. 
Berrumbadero, Cafion de las Capulines above San Enrique Hacienda, San Jose de Raices, Mueller 
2413 (F,GH, MICH, MO). San Luis Potosi: 21.6 mi SE of San Luis Potosi, hwy 86 to Rio Verde, 7500 ft, 
Oliver 190 (MO, SMU). Tamaulipas: 3 mi N of Miquihuana, Stanford et al. ae (DS, US), Zacatecas: 
Sierra del Astillero, 24°34' N, ae 04' W, M.C_ Johnston et al. 11557 (CAS, TEX, 

UNITED eae TEXAS. Culberson Co.: south fork of Mckittrick ae Guadeloupe Mts., 
Correll & Hanson 22 (FSU Victoria Canyon, E margin of Sierra Diablo, 1330-1700 m, 31° 20— 
21' 30" N, 104° 53-55' W, M.CJohnston et al. 10695 (TEX). Hudspeth Co.: Victoria Canyon, 1 mi from 
McAdoo Ranch, Diablo Mts., Muller 8219 (MICH). Jeff Davis Co.: Davis Mts., Madera Canyon, Webster 
4501 (MICH). 


608 BRIT.ORG/SIDA 19(3 


— rupicola (Greenman) Terrell, comb. nov. (Fig. 5).Houstonia rupicola 
reenman, Proc. Amer. Acad. Arts 32:286. 1897, non Hedyotis rupicola Sonder in 
ae S Sonder, FI. Cap. 3:12. 1865. Hedyotis stenophylla Torr. & A. Gray var. 
parviflora A. Gray, Pl. Wright. 81. 1852, non Hedyotis parviflora Walpers, Ann. 
Bot. Syst. 2:772. 1852, nec pa parviflora Holzinger ex Greenman, Proc. Am 
Acad. Arts 32:284. 1897. Hedyotis 1 Fosb. in Shinners, Field & Lab. 17166, 
nom. nov. for var. par aes 1949. He dyotis nigricans var. angulata (Fosb.) WH. 
Lewis, Amer. J. Bot. 49:8605. 1962, nom. superfl. Hedyotis nigricans var. parviflora 
(A. Gray) WH. Lewis, Ann. Missouri Bot. Gard. 55:32. 1968. TyPF: U.S.A TEXAS: Ex- 
pedition from W. Texas to El Paso, N.M., May-Oct 1849, C. Wright 238 (LECTOTYPE 
designated by Lewis (1968): GH}; ISOTYPES: GH! NY! PH! US). 
The protologue states “Crevices of rocks on the San Pedro River; July.” The lectotype 
has atypical plants; the isotype at US has plants that are more typical. The San 
Pedro River is now the Devils River and is near Del Rio, Texas. 


— 


Small perennial herbs with woody taproots becoming | cm thick. Stems 2-15(- 
22) cm tall, sometimes many, quadrangulate, stiffly erect or ascending from 
thick (to lcm), woody, branched base, glabrous imes shiny), scabrous, or 
pubescent on angles. Leaves 2-8(-12) mm long, 0.8-2(-2.5) mm wide, usually 
3-6 times longer than wide, numerous, sessile, narrowly lanceolate to ovate- 
lanceolate or elliptic, thick, leathery, rigidly ascending or spreading, often shiny, 
cuspidate, glabrate, margin scabrid or ciliolate, midrib conspicuous beneath. 
Stipules 0.5-3 mm long, usually with linear or awl-shaped cauda, margins en- 
tire or toothed or ciliolate, teeth sometimes branched. Flowers heterostylous, in 
terminal, few-flowered cymes, pedicels toca. 3 mm long. Hypanthium glabrous 
to pubescent or ciliate; calyx lobes 0.7-2.3 mm long, less than 0.5 mm wide, 
ovate-lanceolate or lanceolate, acute to subcaudate, rigidly erect, with thick 
dorsal rib, slightly to somewhat surpassing the mature capsules. Corollas 3.5-6 
mm long, funnelform, white, buds obovate, often with apical hairs; tubes 2-3.8 
mm long, ca. 0.8-1 mm wide at throat, puberulent or pubescent within; lobes l- 
2.5mm long, 0.7-1 mm wide, lanceolate or ovate, densely pubescent on inner 
faces, sometimes with hairs on apex. Pin flowers (predominating in herbarium 
specimens) with stigma lobes 0.3-0.8 mm long, oblong, stigmas exserted 1-2 
mim beyond corolla throat, styles minutely puberulent, anthers located just be- 
low corolla sinuses. Thrum flowers with anthers 0.8-1.2 mm long, oblong or 
appearing subglobose, straight or curved, on filaments 0.5-1 mm, exserted to 1 
mm beyond throat, stigmas located near midpoint of corolla tube. Mature cap- 
sules 1-2 mm long and wide, slightly longer than wide or subequal, 1/2-3/4 
inferior, subglobose or slightly compressed, glabrous to pubescent. Seeds sev- 
eral-14 per capsule, 0.3-0.7 mm long, 0.25-0.5 mm wide, black or dark brown, 
shiny, ellipsoid, slightly to somewhat compressed, sometimes twisted or angu- 
lar, ventral face with punctiform hilum, testa finely reticulate. Chromosome 
number unknown. Lewis (1962) listed n=9 for Oliver 197 (MO, SMU, TEX) from 
Nuevo Leon as H. nigricans var. angulata (= S. rupicola), however, this collection 


TERRELL, TAXONOMY OF STENARIA 609 


Fic. 5. Stenaria rupicola. CR. Orcutt 753 (US), Sanderson, Texas, 26 Jun 1924. 


is not S. rupicola but S. nigricans var. gypsophila. Stenaria rupicola occurs no 
farther east than Coahuila. 

Phenology.—Flowering May-Novem ber. 

Distribution.—Rock crevices, talus slopes, gravelly soil, often over lime- 
stone, 5300-2000 m (1500-6000 ft). United States: Southwestern Texas (Brewster, 
Crockett, Pecos, Presidio, Terrell, and Val Verde counties); also Culberson (atypi- 
cal); Mexico: northern Coahuila in the Muzquiz area and northwest toward 
Boquillas del Carmen. 


Selected representative specimens examined: MEXICO. Coahuila: 1 mi N of “La Laguna,” Canyon 
del Mulato, Serranias del Burro, about 65 mi NW of Sabinas, 3500 ft, Gould 10584 (ENCB, MICH); S 
Paila (Valley Seco), G. Cepeda, Hinton et al. 16540 (GH, NY, US); 140.3 km NW of Muzquiz on Hwy 53 
towards Boquilla del Carmen, Peterson & Annable 10595 (US); camp below Pichache de Centinela 
about 15 mi, 4600 ft, Warnock 11538 (MICH, SMU). 

UNITED STATES. TEXAS. Brewster Co.: 7 mi E of headquarters, Black a Wildlife Refuge, 
Maravillas Canyon, D.S.& H.B.Correll 35355 (LL). Crockett Co.: Salviastrum Mesa, 32 airline mi NW o 
Ozona, Cory 44377 (TEX). Pecos Co.: 40 mi S of Ft. Stockton on Sanderson Road, ; finckl pera 
main canyon on NE side of —. Madera, 25 mi S of Ft. Stockton, 1300 m, McVaugh ate (MICH). 
Presidio Co.: Capote Canyon, 9 mi NE of Candelaria, Ohlendorf 4717 (LL). Terrell Co.: 1 mi Eo 
Sanderson, 2885 ft, Warnock I 1885 (LL ). 


— 


610 BRIT.ORG/SIDA 19(3) 


This species has been treated by me in the past as Hedyotis angulata, however, 
under Stenaria the earlier epithet, rupicola, may be used. It is quite distinct in 
leaf characters. Some collections are somewhat atypical, possibly due to intro- 
gression from S. nigricans. The species intergrades with S. nigricans and possi- 
bly with S. mullerae. 


5. Stenaria umbratilis (B.L. Robinson) Terrell, comb. nov. Houstonia umbratilis 
B.L. Robinson, Proc. Amer. Acad. Arts 45:401. 1910. Hedyotis umbratilis (B.L. Rob- 
inson) WH. Lewis, Rhodora 63:222. 1961. TYPE: MEXICO. NUEVO LEON: cliffs of 
mountains near Monterrey, 25 Apr 1906, C.G. Pringle 13877 (HOLOTYPE: GH!: 
ISOTYPES: CAS! MEXU-2! MICH! MO! MSC! SMU! US! VT). 

Perennial herb. Stems prostrate, creeping, slender, glabrous or pubescent at 

nodes, rooting at some nodes. Leaves 2-5.5 mm long, 1-3 mm wide, subsessile or 

with petioles to ca. 1.5mm long, l-nerved, ovate or broadly elliptic, glabrous or 
pubescent above, glabrous and slightly paler below, acute or obtuse. Stipules 
minute, glabrous or sparsely pubescent, margins with one or more teeth or 
glands. Inflorescence with flowers solitary, axillary or terminal, on more or less 
straight pedicels to 3 mm long (var. brevipedicellata) or on slender deflected 
pedicels 3.5 to ca. 16 mm long, subtended by one or two bracts or leaves, flowers 

apparently homostylous. Hypanthium glabrous; calyx lobes 0.8-1.5(-2) mm 

long, 0.5-0.7 mm wide, erect, ovate-lanceolate, acutish or obtuse. Corollas 4.7-9 

mm long, funnelform, white; tube 3-5 mm long, 0.5-1 mm wide at base, 1-3 mm 

wide at throat; lobes 1.5-4.2 mm long, 1.0-2.2 mm wide, ovate, spreading, pu- 

berulent or glabrate within. Anthers 0.6-1 mm long, 0.2-0.3 mm wide, narrowly 
oblong or oblong, filaments 0.5-2 mm long, somewhat exserted. Stigma lobes 
ca. 0.5 mm long, included in corolla tube. Mature capsules 1-2 mm long and 
wide, |/2-3/4 inferior, subglobose, thin-walled, dehiscing widely loculicidally. 

Seeds 0.5-0.8 mm long, 0.3-0.6 mm wide, black, shiny, somewhat compressed, 

ovoid or ellipsoid, hilum centric, punctiform, testa finely reticulate. Chromo- 

some number unknown. 

The creeping habit of this species differs markedly from that of the other 
species of Stenaria; however, the ellipsoid seeds led me to place it with the other 
species of this genus. The typical variety has long pedicels that are deflected at 
an angle, whereas var. brevipedicellata consistently has short, more or less 
straight pedicels. The presently known distribution indicates separate ranges. 
The seeds of the two varieties are similar, 


— 


KEY TO VARIETIES OF 5. UMBRATILIS 


1. Stems often densely leafy because of the very short internodes; leaves somewhat 
leathery, revolute; flowers on more or less straight pedicels ca. 1-3 mm long. 
Sb. var. brevipedicellata 
1. Stems not densely le long (3.5-16 mm), 
deflected pedicels. 5a. var. umbratilis 


Ve } | eh fl 


TERRELL, TAXONOMY OF STENARIA 611 


5a. Stenaria umbratilis var. umbratilis (Fig. 6). 
Leaves thin; flowers on long, deflected pedicels. 

Phenology.—Flowering April, May. 

Distribution—Mexico: Nuevo Leon (near Monterrey), Veracruz (mpios. of 
Atzalan and Jalacingo). Rare and disjunct. 
Additional specimens examined: MEXICO. Veracruz: orilla de arroyo, 800 m, Guatemimolo, mpio 
de Atzalan, 13 Apr 1970, Ventura A. 902 (US); orilla de arroyo, 1265 m, Agua Puente, mpio.de Jalacingo, 
17 May 1972, F Ventura A. 5380 (US). 


— 
—= 


These collections have terminal or pseudoterminal flowers on rather long 
pedicels. In April, 1972, |searched unsuccessfully for this species in Diente Can- 
yon, a few miles south of Monterrey. This canyon may have been the type local- 
ity. Seepage areas in this canyon are said to be drier than in Pringle’s time. 


5b. Stenaria umbratilis var. brevipedicellata Terrell, var. nov. (Fig. 7). Type: 
MEXICO. COAHUILA: steep-sided limestone mountains, calcareous gravel, Canon 
de la Gavia south of Rancho de la Gavia, 1250-2200 m, 26° 18'30"-20'N, 1OV 15'-18' 
W, 2-3 Aug 1973, M.C. Johnston, TL. Wendt, F. Chiang, D. Riskind 12032B (HOLO- 
TYPE: US! ISOTYPE: LL!). 


Pedicelli 1-3 mm longi. 


Leaves somewhat leathery, thickish; capsules on pedicels 1-3 mm long. 
Distribution.—Mexico: Coahuila. 


Additional collections: MEXICO. Coahuila. Mina El Aguirreno, N side of Sierra de la sige crevice 
plant, very steep slopes of limestone sierra, calcareous gravel, 1700-2200 m, 26° 05' 30" 26° 06'N, 
101° 36'W,5 July 1973, M.C Johnston, T.L. Wendt, and F. Chiang C.11681G (LL);Mpio.Villa pi ea 
El Rincén, on SW margin of Serranias del Burro (part of Sierra del Carmen), 1400-2100 m, 28° 40' N, 
102° 15'W,ca.80 km SE of Big Bend National Park, Texas, 28 May 1993, Sandra Aguilar Ruiz 320 with 
Diana L. Doan-Crider (TEX). 


This variety has axillary flowers on short pedicels, compared with the long 
deflected pedicels of var. umbratilis 


ACKNOWLEDGMENTS 


Ithank the following for assistance with the illustrations: William Wergin, Chris 
Pooley, Paul Peterson, Harold Robinson, and Marjorie Knowles, lam grateful to 
Joseph Kirkbride, Harold Robinson, and John Wiersema, for providing valu- 
able reviews of the manuscript. Christina Kirkbride helpfully contributed the 
resumen. | also thank curators of the herbaria cited for loans of specimens. 


REFERENCES 


Fernato, M.L. 1940. Soermatophytes of eastern North America. Rhodora 42:299, Plate 625. 

Lawrence, G.H.M. 1951. Taxonomy of vascular plants. New York: The Macmillan Co. 

Lewis, W.H. 1959. Chromosomes of east Texas Hedyotis (Rubiaceae). Southw. Naturalist 3: 
204-207. 


BRIT.ORG/SIDA 19(3) 


i Stenaria umbratilis (B.L. Rob.) Terrell var. ambratitis 
i i Sidayined 
Ror Proc. Amer. Acad. 45:401,1910 
es E.E. Terrell Noy. 2000 Smithsonian Institution 
ars 
Hedy bratilis (B.L. Robinson) W.H. Lewi f SUN 
(Houstonia umpbratilis B.L, Robinson) { (at ‘S-\ 
P= 
Det. EE, Terrell 1990 Ke ws) 
OF ae 
WAL ERES 
nh UNITED STATES NATIONAL MUSEUM 
Howtos uMbratilis Robinson 
ISOTYPE ana aaa SR Tras 
C, G. PRINGLE, 
1973 PLANTA. MEXICAN A. 
as ABO) = —staTE oF NUEVO LEON 4 
Jin r 
tedyotis ts (Robi 2 6 4 me, fp 
Hedyotis umbratilis (Robins.) Lewis 3 i { His f nak ted 
“ a i 
Det. a caws, 1s 64 7 ny 2 
Florbarium | | 2 
Stephen F, Austin Siute College | | me 
00137482 — 


Fic. 6. Stenaria umbratilis var. umbratilis. lsotype, C.G. Pringle 13877 (US 462427). 


TERRELL, TAXONOMY OF STENARIA 


Hedyotis umbratilis (B.L, Robi W.H. Lewis 
(Houstonia umbratilis BLL, Robinson) 
Det. E.E. Terrell 
Unbrersty of Marytand, Cotlepe Park, MD 20742 — Maxey 


Stenaria umbratilis (B.L. Rob.) Terrell brevipedi cell 
Sida, ined. 
Holotype: M.C. Johnston et al. 12032B (US) 


ELE. Terrell Nov. 2000 Smithsonian Institution 


UNITED STATES 
2898060 


waviowat weneariom DIMI 


| 
00610697 


Houstonia Urmbyets Lig Pebinson 


det EE, Tewell, YRS 


No. REC. 

INF. 
—— 
[ROMARE CIENTIFICO 
[pais 


INF, AMBIENTAL, 


—-Sheep-sided limestone mounta 
VELO 


s 
calcareous gravel 
ave lech,, Cercocarpus, Frax ak S_cem 
FORMA BIOLOGICA TAMANO | 


SOCIADA 
Quercus Sp., Dasylirion, 
ABUNDANCIA 
| AN. PERENNE OTROS DATOS 
+ ae C3 
FRUTO 
NOMBRE Loc, 


usaos UT ae amuse meena T 


“ol-M.C,Johnston,?.L.Wendt, F.Chiang, & D. Riskind ~~ no. (LO32LB 


Fic.7.5 i bratili' brevinedicellata Holotype, M.C. Johnston et al. 120328. 


614 BRIT.ORG/SIDA 19(3) 


Lewis, W.H. 1962. Phylogenetic study of Hedyotis (Rubiaceae) in North America. Amer. J. 
Bot. 49: 855-865. 

Lewis, W.H. 1968. Notes on Hedyotis (Rubiaceae) in North America. Ann. Missouri Bot. Gard. 
55:31-33. 

Rogssrecut, E. 1989. A remarkable new Chazaliella (African Psychotrieae), exemplifying the 
taxonomic value of pyrene characters in the Rubiaceae. Bull. Mus. Nat. Hist. Nat., Paris, 
Ser.4, 11,sect.B, Adansonia 4:341-349. 

TerRELL, E.E. 1986. Taxonomic and nomenclatural notes on Houstonia nigricans (Rubiaceae). 
Sida 11:471-481. 

Terrell, E.E. 1991. Overview and annotated list of North American species of Hedyotis, 
Houstonia, Oldenlandia (Rubiaceae) and related genera. Phytologia 71:21 2-243 

Terrett, E.E. 1996a. Revision of Houstonia (Rubiaceae). Syst. Bot. Monogr. 48:1-118. 

TerRELL, E.E. 1996b. Taxonomic notes on Texan and Mexican species of Hedyotis and 
Houstonia (Rubiaceae). Phytologia 81:108-114. 

TerRELL, E.E., W.H. Lewis, H. Rosinson, and J.W. Nowicke. 1986. Phylogenetic implications of 
diverse seed types, chromosome numbers, and pollen morphology in Houstonia 
(Rubiaceae). Amer. J. Bot. 73:103-115 

Turner, B.L. 1995a. Taxonomic overview of Hedyotis nigricans 
lied taxa. Phytologia 79:12-21. 

Turner, B.L. 1995b. Hedyotis pooleana (Rubiaceae), a new species from the Dead Horse 
Mountains, Trans-Pecos, Texas. Phytologia 79:93-96. 

Turner, B.L. 1997. Rebuttal to Terrell’s taxonomic notes of Turner's treatment of Texan and 
Mexican Hedyotis. Phytologia 82:82-85. 


— 


Rubiaceae) and closely al- 


NOTES ON VARIATION IN PSEUDOGNAPHALIUM 
OBTUSIFOLIUM (ASTERACEAE: GNAPHALIEAE) 
Guy L.Nesom 
Botanical Research Institute of Texas 
09 Pecan Street 
Fort Worth, TX 76102-4060, U.S.A. 

ABSTRACT 


l ‘ l Re VALE Rey ioe - . “eg | : Le 
VALIAVDIC OVC! 


ing plac auietis of stems sae upper lea rae iiee’ but the variation is difficu eo- 
graphically. Variety praecox, described from localities on the Atlantic and par acrce as on He 
basis of early-season flowering and an elongate capitulescence, can only be arbitrarily separated 
from the species and does not deserve formal ee asco Variety micradenium, which has been 
treated at varietal rank within both Gnaphalium obtusifolium and the closely related Gnaphalium 


helleri, is distinct in morphology from both and eusnenty reproductively isolated. It is given equiva- 
lent taxonomic rank as Pseudognaphalium micradenium (We eatherby) Nesom, comb. et stat. nov. 


RESUMEN 


D =| Teds 4 ay ft Nala eAda 


i variable a lo largo de su distribucion 
et glandulosidad d llos y haces foliares, aia) i cae ena css 


geograficamente. La variedad praecox, rere de localidades del Atlanhicg y anuras 

Golfo en base a su floracion temprana y una capitulescencia alargada, solo puede ser separada de la 
especie arbitrariamente y no tiene reconocimiento bees La variedad micradenium, que ha sido 
tratada a nivel varietal tanto en Gnaphali , Gnaphalium helleri, es distinta 
BCH CIC Ate de amibas y al parecer aislada Hee aes Se le da el rango taxonomico 
micradenium (Weatherby) N comb. et stat. nov. 


if 


Before its transfer to Pseudognaphalium, as P (Gnaphalium) obtusifolium (L.) 
Hilliard & Burtt, Gnaphalium obtusifolium has, at times, been treated with up 
to five varieties: var. obtusifolium, var. praecox, var. helleri, var. micradenium, 
and var. saxicola (e.g., Fernald 1950; Cronquist 1980; Gleason & Cronquist 1991). 
Aspects of variation affecting the taxonomy of the first four of these are con- 
sidered in the present paper. The status of the morphologically reduced and 
apparently narrowly endemic Gnaphalium saxicola Fassett is currently under 
study by D.S. Feller at the University of Wisconsin. The current study is based 
on examination of more than 750 specimens of P obtusifolium sensu lato from 
BRIT, GH, NCU, TEX-LL, and US. 


1. Variation in vestiture in typical Pseudognaphalium obtusifolium 

The type, distribution, and density of glandular hairs on the stems and leaves 
is significant in the identification of many species of Pseudognaphalium. 
Vestiture is variable within P. obtusifolium and published descriptions have been 
inconsistent. 


SIDA 19(3): 615 — 619. 2001 


616 BRIT.ORG/SIDA 19(3) 


Blake (1918, p. 72) noted that “If the wool of a specimen of [typical] G. 
btusifolium be removed, stipitate glands precisely similar in shape and posi- 
tion to those of the variety [var. helleri] are found.” Fernald described the stems 
and branches as “closely white-lanate” (1936, p. 232) and “covered with a close 
white felt-like pubescence” (1950, p. 1464) without mentioning glands. 
Cronquist (1980, p. 177) observed that the stems are “thinly white-woolly, com- 
monly becoming subglabrous (or even a little glandular) near the base;” in the 
key to species he noted that the stems are “scarcely glandular except sometimes 
near the base.” Observations here are similar to Cronquist’s—glands are uncom- 
mon under the closely lanate stem vestiture, and when glands do occur, they 
are near the base of the plant. Minute, sessile cauline glands rarely may be 
present beneath the cauline tomentum. Plants with glandular stems do not 
show other features to suggest that presence of cauline glands reflects hybrid 
influence of other species. 

Fernald (1936, p. 232) observed that the leaves of Gnaphalium obtusifolium 
“are commonly, though not always, glandular or glandular-papillate above ...” 
Cronquist (1980, p. 177) described the leaves as “from glabrous to slightly aa 
dular or slightly woolly above.” Observations here confirm the variability of 
vestiture on upper leaf surfaces. Glandular upper surfaces are found on plants 
from Illinois, Minnesota, Maryland, Mississippi, Oklahoma, Arkansas, and Texas 
(and probably other states). The glands have short, filiform stipes more like those 
of [var] micradenium than [var] helleri(see key below), but the stipes may be so 
short that the glands are nearly sessile. The persistent, thickened bases of the 
uniseriate, filiform hairs of the tomentum may be gland-like in appearance but 
they are different from the biseriate, glandular hairs. 

The high ratio of pistillate florets to bisexual florets in Pseudognapha 
obtusifolium and the small amount of pollen produced by the bisexual florets 
suggest that self-pollination may be the prevalent mode of fruit production (e.g, 
Noyes 2000); if so, formation of local morphological races would be expected. 
This seems to be the case in central Texas, at the very southwestern corner of 
the range of the species, where plants from counties on the Edwards Plateau 
and slightly eastward have stipitate-glandular leaves, while plants elsewhere 
in the state have leaves eglandular and completely glabrous to sparsely arach- 
noid on the upper surface. Similarly glandular leaves are produced in other parts 
of the range, however, and it would be inconsistent to recognize this Texas en- 
clave with formal taxonomy. Texas plants of P obtusifolium, including those with 
glandular and eglandular leaves, produce the lowest numbers of bisexual flow- 
ers per capitula of any sampled over the whole species range. 


— 


ium 


2. Status of Gnaphalium obtusifolium var. praecox 
Gnaphalium obtusifolium var. praecox Fernald (= Pseudognaphalium obtusi- 
folium var. praecox (Fernald) Kartesz) was described from coastal plain localities 


NESOM 617 


in Virginia, South Carolina, Georgia, and Alabama on the basis of early-season 
flowering and an atypically elongate capitulescence (Fernald 1936). Its presence 
was later noted in New Jersey (Fernald 1950), Maryland (Brown & Brown 1984), 
and North Carolina (Kartesz 1999). In other regional treatments, Ahles (1968) 
treated var. praecox as a synonym of typical Gnaphalium obtusifolium, while 
Cronquist (1980) and Gleason and Cronquist (1991) did not even mention it. 
The taxon as an accepted entity apparently has persisted only ina few floristic 
accounts, e.g., Massey (1961), Brown and Brown (1984), Kartesz (1999). 

Plants of var. praecox cited in the protologue by Fernald (1936) flowered 17 
July, 23 July, 5 August, and 11 August. Other specimens (GH, US) collected by 
Fernald in southwestern Virginia and identified by him as var. praecox were 
flowering in August and September. Based on records from more than 600 col- 
lections of P. obtusifolium at BRIT, GH, NCU, TEX-LL, and US, plants at anthesis 
in the southeastern United States have been collected primarily from the first 
week of August through mid-October, with a few early bloomers in the last two 
weeks of July anda few late bloomers in the first two-thirds of November. Flow- 
ering times in Canada and more northern parts of the U.S. apparently begin 
slightly later and end slightly earlier, as most collections have been made from 
mid-August through September and early October. These observations extend 
the beginning of normal flowering of the species in the southeastern U.S.,com- 
pared to the range of flowering times observed by Fernald in 1936 on the basis 
of GH collections. While the collections of var. praecox cited by Fernald are 
seasonally early, they are not outside the range of normal flowering for typical 
plants of the species and it does not seem unusual that on the Atlantic and Gulf 
coastal plains, with warmer temperatures and generally earlier phenologies, 
flowering in P.obtusifolium should begin earlier than elsewhere in the range. 

The capitulescence of Pseudognaphalium obtusifolium normally is dis- 
tinctly corymboid (flat-topped), this configuration usually attained before the 
capitula reach full anthesis. In plants described by Fernald as var. praecox, flo- 
riferous branches developing from relatively lower nodes and not elongating to 
the full capitulescence height produce a capitulescence that is “elongate-cylin- 
dric to thyrsoid.” In P obtusifolium, a cylindric capitulescence in full anthesis is 
rare, and most of the plants identified by Fernald as var. praecox did not pro- 
duce the distinctly cylindric capitulescence shown in the type specimens (see 
protologue photograph). It is not uncommon to encounter a pumauly 
corymboid capitul at anthesis with lower branches in vari t 
earlier development or with lower branches at anthesis considerably anertey 
than the upper branches. Nor do such variants appear to be strongly correlated 
with early flowering. 

Fernald (1936, p. 233) noted that “search for technical characters [to distin- 
guish var. praecox]in flowers and achenes has failed to reveal them,” and this is 


618 BRIT.ORG/SIDA 19(3) 


confirmed here. Plants segregated by Fernald as var. praecox tend to flower in 
the earlier part of the phenological range of the species and their capitulescenc 
differs in a minor and overlapping way from plants of typical P. obtusifolium, 
but no other differences separate these taxa. This evidence indicates that var. 
praecox has no status as an evolutionary entity and its taxonomic recognition 
is not justified. 


3. Status of Gnaphalium obtusifolium var. micradenium 

Var. micradenium was originally described at varietal rank by Weatherby (1923) 
as “apparently the northern and more inland representative of [Gnaphalium 
obtusifolium] var. helleri (Britton) Blake, which seems to be confined to the 
coastal plain.” It was accepted at varietal rank by Fernald (1950, as G. 
obtusifolium var. micradenium), Mahler (1975, as G. helleri var. micradenium), 
and other floristic accounts. Ahles (1968), however, placed var. micradeniumas 
asynonym of G. obtusifolium (vs.G. helleri), while Cronquist (1980) and Gleason 
and Cronquist (1991, implicitly) treated var. micradenium as a synonym of G. 
helleri. 

Weatherby’s (1923) initial contrast of var. micradenium and var. helleri, 
emphasizing glandularity and leaf morphology, remains relatively accurate. 
Difference in gland morphology and the number of florets per capitulum fur- 
ther sharpen the distinction. Although var. micradenium and var. helleri have 
largely allopatric ranges (generally as mapped by Mahler 1975), a region of sym- 
patry exists in Georgia, South Carolina, North Carolina, and southeastern Virginia. 
Intermediates have not been encountered, suggesting that they are reproductively 
isolated. The two taxa were intermixed and collected together in Northampton 
Co, Virginia—var. micradenium (Fernald and Long 5770,GH), var. helleri (Fernald 
and Long 5550 and 5551,GH)—as noted by the label data on the collections. Evi- 
dence at hand indicates that each is reasonably treated as a distinct species, as 
reflected in the new combination and summarized in the morphological con- 
trasts in the key below. 


Pseudognaphalium micradenium (Weatherby) Nesom, comb. et stat. nov. 
snaphalium obtusifolium var. micradenium Weatherby, Rhodora 25:22. 1923. 
Gnaphalium helleri var. micradenium (Weatherby) Mahler, Sida 6:32. 1975. 
Pseudognaphalium helleri var. micradenium (Weatherby) Kartesz, Synth. N. Amer. 
FL. (ed. 1). 1999. Type: UNITED STATES. MASSACHUSETTS. Barnstable Co. dry, sandy 
openings among scrub oak, Barnstable, 7 Oct 1917, M.L. Fernald 15870 (HOLOTYPE: 
GH!) 


— 


Ss) 


. Stems persistently white-tomentose-felted, sometimes lightly so, usually \dular 
uncommonly glandular near the base; pistillate florets 38-96, eae floiets re 
(-11); plants only slightly if at all fragrant. seudognaphalium ene 
1. Stems quickly losing most tomentum, greenish, persistently and densely stipitate- 
glandular; pistillate florets 47- peote! florets 7- as plants distinctly fragrant. 
2. Stems glandular-puberulent,t -0.2 mm high, relatively even 
in height on any portion of the stem, wit ith a Flitorn stalk of even width and 


NESOM 619 


narrower than the gland width; pistillate florets 47-78, bisexual florets (7—-)11- 
20; leaves linear to linear-lanceolate or oe 1.5-5.5 cm long, 1.5- 
10 mm wide, 6-10 times longer than wide. eudognaphalium micradenium 
. Stems glandular-villous, the stipitate glands (0.1-)0.3- 1 mm high, prominently 
variable in height on any portion of the stem, with a stalk broadened toward the 
base and ca equal the gland width; pistillate florets 83-107, bisexual a 9- 
15; leaves mostly oblong-lanceolate, 25-7 cm long, 4-20 mm wide, 4-8 
longer than wide. Seat itin helleri 


N 


ACKNOWLEDGMENTS 


lam grateful to the staffs of BRIT, GH, NCU, TEX-LL, and US for their help dur- 
ing study at those herbaria and to Walter Holmes and Alexander Krings for 
comments on the manuscript. 


REFERENCES 


Antes, H.E. 1968. Asteraceae. In: Radford, A.E.,H.E.Ahles and C.R. Bell. Manual of the vascular 
flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill. Pp. 1009-1139. 

Biake, S.F. 1918. Notes on the Clayton herbarium. Rhodora 20:21-28, 48-54, 65-73. 

Brown, M.L. and R.G. Brown. 1984. Herbaceous plants of Maryland. Book Center, Univ. of 
Maryland, College Park. 

Cronauist, A. 1980.Vascular flora of the southeastern United States. Vol. 1.Asteraceae. Univ. 
of North Carolina Press, Chapel Hill. 

Fernatp, M.L. 1936. Varieties of Gnaphalium obtusifolium. Rhodora 38:231-233. 

Fernavo, M.L. 1950. Gray's manual of botany (ed. 8). American Book Co., New York, NY. 

Gieason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada (ed. 2). The New York Botanical Garden, Bronx, NY. 

Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the 
vascular flora of the United States, Canada, and Greenland. First Edition. In: J.T. Kartesz 
and C.A. Meacham. Synthesis of the North American flora, Version 1.0.North Carolina 
Botanical Garden, Chapel Hill, NC. 

Manter, W.F. 1975. Typification and distribution of the varieties of halium helleri Britton 
(Compositae-|Inuleae). Sida 6:30-32. 

Massey, A.B. 1961. Virginia flora. Virginia Agric. Exp. Stat. Tech. Bull. 155, Blacksburg. 

Noyes, R.D. 2000. Biogeographical and evolutionary insights on Erigeron and allies 
(Asteraceae) from ITS sequence data. Plant Syst. Evol. 220:93-114. 

WeatHersy, C.A. 1923.Some critical plants of Atlantic North America. Rhodora 25:1 7-23. 


620 BRIT.ORG/SIDA 19(3) 


Book REVIEW 


DaPHNE GAIL FAuTIN, DouGLas J. FutuyMa, and Frances C. JAMES (eds). 2000. Annual 
Review of Ecology and Systematics. Volume 31, 2000. (ISBN 0-8243-1431-X, 
hbk; ISSN 0066-4162). Annual Reviews Inc., 4139 El Camino Way, P.O. Box 
10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.AnnualReviews.org, 
800-523-8635, 650-493-4400, 650-424-0910 fax). $140.00, 630 pp, 6" x 9", 

Readers of Sida will find many subjects of interest in this Annual Review of the ‘mutually support- 

ive’ disciplines of Ecology and Systematics. Volume 31 for the year 2000 has 22 articles followed by a 

Subject Index, Cumulative Index of Contributing Authors, and Cumulative Index of Chapter Titles 


Contents.—Preface: A Millennial View of Ecology and Systematics, and ARES at Age 30 
1, General Principles 
1, The Kinship Theory of Genomic Imprintin 
2. Cenozoic Mammalian Herbivores from the Americas: Reconstructing Ancient Diets and Terrestrial 


Communities 
3. Conservation Issues in New Zealand 
4. The Evolution of Predator-Prey Interactions: Theory and Evidence 
5. The Ecology and Physiology of Viviparous and Recalcitrant Seeds 
6. Inbreeding Depression in Conservation Biology 
7. African Cichlid Fishes: Model Systems for Evolutionary Biology 
8. Shrub Invasions of North American Semiarid Grasslands 
9. The Grasses: A Case Study in Macroevolution 
10. The Ecology of Tropical Asian Rivers and Streams in Relation to Biodiversity Conservation 


i 


11. Harvester Ants (Pogonomyrmex spp.): Their Community and Ecosystem Influences 
12. Origins, Evolution, and Diversification of Zooplankton 

13. Evolutionary Physiology 
14. Mechanisms of Maint { Species Diversity 


15. Temporal Variation in Fitness Components and Population Dynamics of Large Herbivores 

16. Impacts of Airborne Pollutants on Soil Fauna 

17. Ecological Resilience - in Theory and Application 

18. Quasi-Replication and the Contract of Error: Lessons from Sex Ratios, Heritabilities and Fluctu- 
ating Asymmetry 

19. Invasion of Coastal Marine Communities in North America: Apparent Patterns, Processes, and 


Biases 
20. Diversification of Rainforest Faunas: An Integrated Molecular Approach 


21. The Evolutionary Ecology of Tolerance to Consumer Damage 
—Guy L. Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, 
U.S.A, 


SIDA 19(3): 620. 2001 


BIOLOGICAL STATUS OF ARGYTHAMNIA LAEVIS 
(EUPHORBIACEAE) 


B.L. Turner 


Section of Integrative Bio 
University of Texas 
Austin, 1X, 78713, U.S.A. 


ses 


ogy 


Argythamnia laevis (A. Gray) Muell., a taxon of Trans-Pecos Texas and closely ad- 
jacent southeastern New Mexico, was first described in 1859 as Aphora laevis A. 
Gray ex Torrey. It was distinguished from its closest congener, Argythamnia humilis 
(Engelm. & A. Gray) Muell., by its glabrous condition. Other than its striking 
glabrosity, including reproductive organs, A. laevis is seemingly identical to A. 
humilis. Shinners (1956) reduced A. laevis to varietal rank under the latter with 
the observation: “Rather rare in the Trans-Pecos (specimens seen from Jeff Davis 
and Reeves counties); var. humilis is common and widespread on prairies of 
central and western Texas.” The treatment of Shinners has been followed by 
most subsequent workers (eg., Johnston @ Warnock 1962; Correll & Johnston 1970). 
I became interested in the biological status of A. laevis (= Ditaxis laevis[A. 
Gray ex Torrey] Heller) in my preparation of a taxonomic account of Ditaxis for 
Trans-Pecos Texas. Johnston and Warnock (1962) provided a systematic account 
of the varieties concerned. In this they mapped the two as essentially sympatric 
but not intergrading or co-occurring in a given population, this suggesting either 
specific status for A. laevis, or perhaps mere recognition of the latter as a form. 
Discovery of the two taxa within a single population should prove helpful in 
resolving this issue. To this end | began to look intensively at any given popula- 
tion of A. humilis in hopes of finding forms referable to A. laevis. Among five or 
more populations from the trans-Pecos and peripheral areas, only two such 
populations were found, as indicated below and shown in Figure 1. 


XAS. Andrews Co.: northeast shoreline and along roadside of Shafter Lake, 12 May 2000, B. L.& 

Matt Turner 20-263A (TEX). Gaines Co.: 3.8 mi S of Seminole along Farm Rd 181, 12 May 2000, B.L.& 
Matt Turner 20-246 (TEX). 
In the two mixed populations, pubescent forms (var. humilis) were clearly much 
more common than the glabrous forms (var. laevis). Further, I never encoun- 
tered pure populations of the glabrous form. Because of this ] conclude that “var. 
laevis” is but a sporadically occurring form of A. humilis, undeserving of vari- 
etal rank as this is conceived by Turner and Nesom (2000) and perhaps others. 
Better proof might be obtained through sowing field-gathered seeds of “var. 
laevis” so as to show that both pubescent and glabrous forms ptt arise from 
the seedlings concerned, the glabrous condition apparently due to the expr 


SIDA 19(3): 621 - 622. 2001 


BRIT.ORG/SIDA 19(3) 


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of only one or a few genes, but the field observations provided here seemed 
sufficient to establish that likelihood. 

Because of the nomenclatural history and striking appearance of the taxon 
concerned | deem it appropriate to reduce Argythamnia laevis to the category 
of forma, as follows: 

Ditaxis humilis forma laevis (A. Gray ex Torrey) B.L. Turner, forma nova. BASIONYM: 
Aphora laevis A. Gray ex Torrey, Bot. Mexican Bound Surv. 196. 1859. 
REFERENCES 
Corrett, D.S.and M.C. JoHnston. 1970. Manual of the vascular plants of Texas. Univ. of Texas 


Press, Austin. 
JoHnston, M.C. and B.H. Warnock. 1962.The four kinds of Argythamnia in far western Texas. 


SouthW. Naturalist 7:154-162. 
SHINNERS, L.H. 1956. Botanical notes. Field & Lab. 24:38. 
TURNER, B.L.and G.L. Nesom. 2000. Use of variety and subspecies and new varietal combina- 


tions for Styrax platanifolius (Styracaceae). Sida 19:257-262. 


VALIDATION OF THE NAME OROBANCHE LUDOVICIANA 
SUBSP. MULTIFLORA (OROBANCHACEAE) 


Heather L.White and Walter C. Holmes 


Department of Biology 
aylor University 
Waco, TX 76798-7388, U.S.A. 


The combination Orobanche ludoviciana Nutt. subsp. multiflora Collins first 
appeared in Collins (1973), a doctoral dissertation, and was never published. 
Nonetheless, the name has gained some measure of acceptance by botanists 
nationally (Kartesz & Meacham 1999) and in Texas (Jones et al. 1997; Diggs et 
al. 1999). Thus, because the combination has not been validly published, the 
following is proposed. 
Orobanche ludoviciana Nutt. subsp. multiflora (Nutt.) Collins ex H.L. White & 
CT. es, Stat. nov. BASIONYM: Orobanche multiflora Nutt., Proc. Acad. Phila- 
delphia 4:22. 1848. Orobanche ludoviciana Nutt. var. multiflora (Nutt.) Beck, Bibl. 
Bot. 4, 19:81. 1890. Myzorrhiza multiflora (Nutt.) Rydb., Bull. Torrey Bot. Club. 
33:151. 1906. TyPE: U.S.A. NEW MEXICO. Sandy ground along the Rio Grande, 1845, 
Gambel s.n. (Not located at BM, GH, or PH). Since no duplicates of the type have 
been located, we select a neotype herewith. TYPE: U.S.A. TEXAS. Jim Wells Co. 20.5 
mi S of George West, off U.S. [Hwy] 281, near county boundary line, 19 Apr 1944, 
Lundell & Lundell 12809 (NEOTYPE, here designated: LL). 


Phelypaea erianthera Engelm. in A. Gray, Proc. Amer. Acad. Arts 7:372. 1867, as 

helipaea.” TYPE: COLORADO: Huerlano Valley, Parry 147 (GH, MO). 

Orobanche multiflora Nutt. var. arenosa (Suksdf.) Munz, Bull Torrey Bot. Club 57:623. 
1931. TyPE: WASHINGTON. Klickitat Co. Bingin, Suksdorf 2781 (C, G). 

Orobanche multiflora Nutt. var. pringlei Munz, Bull. Torrey Bot. Club 57.623. 1931. 
TYPE: MEXICO. CHIHUAHUA: Chichuahua, 3 Jun 1885, Pringle 31(G). 


me 


The description of Orobanche multiflora was published twice, originally in the 
Proceedings of the Academy of Natural Sciences of Philadelphia and later in 
the Journal of the Academy of Natural Sciences of Philadelphia, ser. 2, 1179. 
1848. Reveal and Spevak (1976) give the date of publication of the Proceedings 
as between 21 March and 4 April, 1848, while publication date of the Journal is 
given as between 1 August and 8 August 1848. Orobanche multiflora is best 
recognized as a subspecies of O. ludoviciana because of the excessive intergra- 
dation of the two taxa. The following brief discussion is largely taken from Col- 
lins (1973), which we have confirmed By eho of herbarium specimens 
from NY and TEX. They are distingui le as subspecies except in the area of 
sympatry in Texas and Oklahoma where the two subspecies appear to inter- 
breed. Corollas of subsp. multiflora are mostly 20-30 mm long, while those of 
subsp. ludoviciana vary from 15-20 mm long. The corolla lobes of subsp. mul- 


SIDA 19(3): 623 - 624, 2001 


624 BRIT.ORG/SIDA 19(3) 


tiflora tend to be broader and either obtuse or acute. Those of subsp. ludoviciana 
are narrower and either obtuse or acute. The anthers of subsp. multiflora are 
woolly and usually inserted and the style is persistent on the capsule while 
anthers of subsp. ludoviciana are glabrous and frequently exserted and the style 
is not persistent on the capsule. Hosts for the two subspecies are also variable; 
subsp. multiflora parasitizes local Compositae except for Artemisia, whereas 
subsp. ludoviciana parasitizes principally Artemisia and Grindelia. 

We wish to thank the curator and staff at TEX for the loan of Orobanche 
specimens. The hospitality and information provided by NY during a visit by 
the senior author is much appreciated. John Pruski of US provided literature 
and also reviewed a preliminary version of the manuscript, while Dan Nicolson, 
also of US, gave advice on the citation of synonymy. Robert Baldridge of the 
Department of Biology, Baylor University, also reviewed and commented on 
portions of the manuscript. 


REFERENCES 


Couuns, T.S. 1973. Systematics of Orobanche section Myzorrhiza (Orobanchaceae) with 
emphasis on Orobanche ludoviciana. Unpub. Dissertation, University of Wisconsin-Mil- 


waukee. 

Dices, G.M., B.L. Liescoms, and R.J. O'Kennon. 1999. Shinners & Mahler's illustrated flora of 
north central Texas. Botanical Research Institute of Texas, Fort Worth. 

Jones, S.D.,J.K. Wiprr, and PM, Montcomery. 1997. Vascular plants of Texas: a comprehensive 
checklist including synonymy, bibliography, and index. University of Texas Press, Aus- 
tin. 

Kartesz,J.T.and C.A. Meacham. 1999, Synthesis of the North American flora (Ver. 1.0).North 
Carolina Botanical Garden, University of North Carolina, Chapel Hill. 

Reveal, J.L. and V.S. Spevak. 1967. Publication dates and current names of 144 names pro- 
posed in two 1848 Thomas Nuttall articles. Taxon 16:407-414. 


AN ANOMALOUS POPULATION OF ASTER 
(ASTERACEAE: ASTEREAE) SENSU LATO IN MICHIGAN 


Guy L.Nesom 
Botanical Research Institute of Texas 
509 Pecan Street 
Fort Worth, TX 76102-4060, U.S.A. 
ABSTRACT 


Plants from a single population on the Keweenaw Peninsula in Houghton County, Michigan, have 
been reported as hybrids between Doellingeria (Aster) umbellata and Oclemena (Aster) nemoralis, 
w features of putative intermediacy ella these two species are equnoes In contrast, 


pores ae features of their roots, leaves florets, and with 


e of Oclemena—in the interpretation here miorpHelogical eudente foe Ene eenoHic< COnbADUr 
tion oon ee very limited, and tl likely arose from 
They are suggested to represent either an sa aa ee of O. xblakei G recurring and persistent 
pybne between O. nemoralis and O.(A i) or a relict ee differentiated earlier 


5 - 4 


history of the genus a separate species. Current 


a > 


the y 
evidence is insufficient to nc sn ape a nek Liypouhesis 7 origin. 


RESUMEN 


a Peninsula de Keweenaw en el Condado de Houghton, Michigan, 
is, pero las 


Plantas de una sola popcen en 
se han ciladore co hibrid Doellingeria oe umbellata y Oclemena Cag) nemora 


<1 


rr 
= ay 


ontraste 


i os 
las caracteristicas morfoldgicas de sus raices, hojas, sspiileceene 4 ideation y cipselas hones 


con los de Oclemena—en la interpretacion que hacemos aqui, las evidencias morfologicas de una 
contribucion de Doe tinea ia es muy limitada, y - plantas sae susie probabler emente surgieron 


e Oclemena.S 


y persistente date O. nemorals y O. (Aster) ac sinieiaea) o una pop ycen relicta diferenciada 
tempranamente en la Blons ev anion a del genet y reconocida a especie 


L 


independiente. | i una nalapeleee unica aE suorigen. 


£ [ 


The recent report (Gerdes 1998) of a population of hybrids between Doellingeria 
(Aster) umbellata (P. Mill.) Nees and Oclemena (Aster) nemoralis (Ait.) Greene 
from the Keweenaw Peninsula of Lake Superior in northwestern Michigan is 
remarkable. Such an intergeneric hybrid would provide corroborative evidence 
that these two genera (Doellingeria Nees and Oclemena Greene) segregated from 
Aster are closely related. Naturally occurring intergeneric hybrids in the 
Astereae have previously been reported only between closely related genera— 
in some cases these hybrids may be better interpreted as between congeneric 
species (Nesom 1994b). 

A hypothesis of close relationship between Doellingeria and Oclemena 
(Nesom 1994a) is based on morphological similarities. In contrast, molecular 
evidence (Semple et al. 1996; Xiang & Semple 1996) suggests that Oclemena may 


SIDA 19(3): 625 — 632. 2001 


= 


626 BRIT.ORG/SIDA 19(3) 


be closely related to Ionactis Greene. A hypothesis of close relationship between 
Doellingeria and Oclemena also is suggested in the disparity of interpretation 
regarding the position of Aster reticulatus Ell, a southeastern USA endemic 
placed by Semple et al. (1991, 1996) in Doellingeria but by Nesom (1993, 1994a, 
2000) in Oclemena. Hybridization between species of Doellingeria and 
Oclemena has not been previously reported, although D. umbellata and O. 
nemoralis are sympatric over most of the range of the latter and both species 


occur in wetland habitats. 

Gerdes noted that the putatively hybrid population was distinctive in the 
field and appeared to be intermediate between nearby populations of 
Doellingeria umbellata and Oclemena nemoralis. While the putative hybrids 
clearly do not fit the typical morphology of either species, the only indication 
of intermediacy in the published observations is in the statement (p. 18) that 
the “overall height and leaf size and shape are intermediate between the as- 
sumed parents.” In the illustration and description, and in the voucher speci- 
mens, other possible indications of intermediacy between D. umbellata and O. 
nemoralis are equivocal. 


Plants studied 

The population described by Gerdes (Gerdes 2210) includes about 20 plants in 
a tamarack swamp situated between populations of Doellingeria umbellata 
(‘common 60 m north of the hybrid site ina mixed wetland complex”—Gerdes 
2211) and Oclemena nemoralis (ca. “60 m southeast of the hybrid site in a 
swale”—Gerdes 2209). Two plants of the putative hybrids and one plant each of 
D. umbellata and O. nemoralis from the Houghton County site were available 
for study (MICH), as well as many other specimens from taxa of Oclemena and 
Doellingeria (BRIT, GH, NCU). The two plants of 2210 are virtually identical in 
micromorphological detail, perhaps representing a single clone, and Gerdes did 
not note the occurrence of significant variation among the putative hybrids. 
The plant of O. nemoralis is morphologically typical of the species—this col- 
lection represents the westernmost known population of the species (see Voss 
1996; Gerdes 1998). The collection of D. umbellata may be arbitrarily identified 
as var. umbellata; hairier plants of the species (D. umbellata var. pubens (A. Gray) 
Britt.) also occur on the Keweenaw Peninsula (as mapped by Semple et al. 1991). 


Houghton County specimens examined: 

1) Doellingeria umbellata (P. Mill.) Nees-MICHIGAN. Houghton Co. Grand 
Traverse Bay ridge and swale complex, ca. 7 mi ESE of Lake Linden, travel E of 
Rice Lake ca. 0.6 mi and walk N of gravel road into swale, edge of sandy road 
and Sphagnum/ericaceous swale; Aster nemoralis also in swale; rays creamy 
white; 18 Sep 1997, L.B. Gerdes 2211 (MICH). 


NESOM AOTCR IN MIRAIOAN 627 


2) Oclemena nemoralis (Ait.) Greene—MICHIGAN. Houghton Co. Grand Traverse 
Bay ridge and swale complex, ca. 7 mi ESE of Lake Linden, and E of Rice Lake. 
Swale, primarily a Sphagnum/ericaceous bog (poor fen); plants scattered and 
numerous with Carex michauxiana, Carex oligosperma, Andromeda glaucophylla, 
etc; rays dark pink; 18 Sep 1997, L.B. Gerdes 2209 (MICH). 


3) The putative hybrid—MICHIGAN. Houghton Co.: Grand Traverse Bay ridge 
and swale complex, ca. 7 mi ESE of Lake Linden and E of Rice Lake. Tamarack 
swamp adjacent to a Sphagnum/ericaceous swale; ca. 20 plants with Aronia 
prunifolia, Chamaedaphne calyculata, Alnus rugosa, Calamagrostis canadensis, 
and Carex oligosperma, Aster nemoralisand Aster umbellatus nearby; rays light 
pink; 18 Sep 1997, L.B. Gerdes 2210 (MICH—2 sheets). 


Evidence for the Doellingeria genome 

Consideration of the illustration and published description of Gerdes 2210 and 
examination of the voucher specimens indicate that features of putative inter- 
mediacy between Oclemena nemoralis and Doellingeria umbellata are rela- 
tively few. 

Height.—Plants of 2210 are 6-7 dm tall, generally between the range of 
height for O. nemoralis (2-7 dm) and D. umbellata (3-20 dm) but also at the top 
of the range for O. nemoralis as well as O. acuminata. 

Leaves.—Leaves of 2210 are narrowly elliptic, 4.5-5.5cm long and 7-10 mm 
wide, with entire, barely revolute margins. Leaves of D. umbellata are elliptic to 
narrowly elliptic, mostly 6-12 cm long, 10-25 mm wide, and flat-margined; 
those of O.nemoralis are very narrowly oblong to linear-lanceolate, mostly 2-5 
cm long, 2-12 mm wide, with entire, strongly revolute margins. Leaves of 2210 
are intermediate in size between D. umbellata and O. nemoralis but they are 
more similar in shape to those of D. umbellata. The abaxial surfaces of 2210 are 
minutely sessile-glandular and could be regarded as intermediate between the 
glandular (larger glands) surfaces of O. nemoralis and eglandular surfaces of 
D.umbellata; the glandularity of 2210, however, also appears to be indistinguish- 
able from that of plants hybrid between O. nemoralis and O. acuminata (see 
below). 

Ray florets.—Rays of 2210 are pink, intermediate between the purple of O. 
nemoralis and white of D. umbellata; pink rays also are characteristic of hybrids 
between O. nemoralis (purple-rayed) and O. acuminata (white-rayed). The 
length of the rays intermediate between O. nemoralis and D. umbellata but 
within the range of variation for O. nemoralis. 

Cypselar vestiture —Cy pselae of 2210 are densely strigose-hispid, more simi- 
lar to the sparsely to densely strigose cypselae of D. umbellata than the typi- 
cally glabrous ones of O. nemoralis (thus this aspect of vestiture is technically 
not intermediate). Otherwise, the densely sessile-glandular cypselar surfaces 


628 BRIT.ORG/SIDA 19(3) 


] 
o 


of 2210 are similar to those of O. nemoralis but not to the surfaces of 
D. umbellata. 

Morphological evidence that Doellingeria umbellata is represented in the 
genome of the plants of Gerdes 2210 is limited, represented by the features of 
equivocal interpretation noted above. In contrast, various features of the roots, 
leaves, capitulescence, florets, and fruits refer Gerdes 2210 to Oclemena. The 
comparisons in Table | summarize morphological distinctions between 
Oclemena nemoralis and Doellingeria umbellata—these also are essentially the 
contrasts that distinguish the two genera (see Nesom 1994a). In each contrast, 
the morphology of 2210 is similar to that of Oclemena. 

If the assessment here of Gerdes 2210 is correct in excluding Doellingeria 
on its close ancestry, two alternative hypotheses would place the evolution- 
ary origin of this anomalous population from within Oclemena. First, it may 
represent a hybrid between O. nemoralis and O. acuminata (Ait.) Greene, or sec- 
ond, it may represent a lineage of Oclemena differentiated early in the evolu- 
tionary history of the genus and now persisting only asa relict at the Hough- 
ton County site. 


Identification as Oclemena <blakei 

The plants of Gerdes 2210 are nearly identical in overall aspect to some indi- 
viduals of Oclemena x blakei (Porter) Nesom, a fertile, recurrent, and persistent 
hybrid between O. nemoralis and O.acuminata—compare Fig. | of Gerdes (1998) 
to Figs. 3. and 11 of Pike (1970). Populations of O. x blakei are scattered through 
the area of sympatry of the parents where ecological conditions allow their 
close contact (Brouillet & Simon 1981). “Extensive colonies of [O.| x blakei are 
often found at the edges of bogs, the shores of ponds, and swampy borders of 
woods, etc., the kinds of areas that are intermediate in wetness between the bo- 
real forest habitat of [O.] acuminata and the open bogs of [O.] nemoralis” (Pike 
1970, p. 401). 

If the plants of Gerdes 2210 are scored on the morphological hybrid index 
developed by Pike (1970; also see Hill & Rogers 1973; Brouillet @ Simon 1981), 
the value is 9 (Table 2), which is within the range characteristic of O. x blakei, 
intermediate between O. nemoralis and O. acuminata. Oclemena xblakei is 
known to produce fertile seeds (Hill & Rogers 1973), but it was not possible to 
make unequivocal observations of fertility for Gerdes 2210. Pollen grains on the 
stigmatic surfaces were regular in size, but the anthers had opened before the 
collection and an estimate of pollen fertility could not certainly exclude pollen 
from other species. None of the cypselae of Gerdes 2210 were completely ma- 
ture when collected; dissected cypselae examined from each plant had produced 
an elongated but otherwise undeveloped embryo. Oclemena xblakei and its 
parental species are known to have a chromosome number of 2n = 18 (Hill & 
Rogers 1970). Scoring of 2210 on this hybrid index does not indicate that it 


NESOM 


Table 1. isla ase ay contrasts between Oclemena 


plants of Gerdes 2 


d Doellingeria umbellata and the 


Oclemena nemoralis 


Gerdes 2210 


Doellingeria umbellate 


Root System — slender, elongate 
rhizomes, without 
a cluster of thick, 
fibrous roots 

Leaves glandular abaxially 

Peduncles long, flexuous 

Ray Corollas pink to purple, 11-18 


Disc Corollas 


Cypselae 


Pappus 


mm long, strongly 
coiling 

narrowly tubular- 
unnelform, slightly 


widened above the tube; 


lobes cut ca. 1/3 of limb, 
erect to spreading 


sessile- -glandular, other- 
e glabrous, fusiform 


slender, ele 
rhizomes, withot 

a cluster orice fibrous 
roots 

glandular abaxially 

long, flexuous 

pink to purple, 10-12 
mm long, strongly 
coiling 

narrowly tubular- 
funnelform, slightly 
widened above the tube; 
lobes cut ca. 1/3 of limb, 
erect to spreading 


sessile-glandular, strigose 


dense cluster of thick fibrous 
oots at the sub-caudex or 
crown, also with eleongate- 
rhizomes 


eglandular 

short, stiff 

white, 5-9 mm long, not 
coiling 


broadly funnelform, abrupt 
widening above the tube; 
lobes cut nearly to base 

of limb, reflexing-coiling 


eglandular, strigose, obovoid 


-hispid, fusiform to 


to nal rowly columnar, Ca. 


1/2-2/3 the phyllary 
length at maturity 

inner and outer series of 
+- even length 


narrowly columnar, Ca. 
1/2—2/3 the phyllary 
length at maturity 

inner and outer series of 
+- even length 


nearly equal the oe 
length at matu 


outer series of short (<1 mm) 


setae 


Taste 2. Morphological hybrid index for Gerdes 2210 


Character State Value 
No. of leaves 35-100 0 
Internode eng (mm) 9-11 ] 
Ratio leaf | length width 5-4 2 
eaf margin—revoluteness -flat 2 
Leaf margin—scabrosity intermat 1 
Leaf margin—toothing entire 0 
No. of bracts per peduncle 2-4 1 
No. of heads per capitulescence 2+ ] 
Ray color pink 0 
Zebra hairs on stem occur | 


Hybrid index value = 9 


630 BRIT.ORG/SIDA 19(3) 


actually isa hybrid between O. nemoralisand O. acuminata but does show that 
its features can be interpreted as intermediate in the same way as those of known 
hybrids can be. 

Two conditions complicate the identification of Gerdes 2210 as Oclemena 
x blakei: (1) O. x blakei previously has been found only within the area of sym- 
patry of the parents, and one of the parental species, O. acuminata, does not 
occur in the Keweenaw region; and (2) plants of Gerdes 2210 have entire leaves 
and hispid cypselae, features not generally characteristic of O. x blakei. 

(1) Oclemena x blakei has been known to occur only much further east— 
the closest known locality to the Keweenaw site is in southeastern Ontario, about 
850 kilometers eastward (maps in Brouillet & Simon 1981, Semple et al. 1966). 
The closest known locality for O. acuminata is the same region of Ontario. 
Oclemena nemoralis grows immediately adjacent to the 2210 population, but 
itis unlikely that O. acuminata has occurred naturally in Michigan since post- 
glaciation revegetation (Brouillet & Simon 1981). Long-distance dispersal would 
the most likely explanation for the far-disjunct occurrence of O. x blakei on the 
Keweenaw Peninsula—at least it isa simpler hypothesis than a postulate of the 
former occurrence and extirpation of O. acuminata far west of its present geo- 
graphic range. “Long-distance dispersal and chance establishment in suitable 
bogs ... are probably responsible for [the] establishment lof O. nemoralis] in the 
eastern Lake Superior area ...and the species could still be expanding its range 
westward...” (Brouillet & Simon 1981, pp. 539-540). The discovery of O. x blakei 
at the westernmost locality of O. nemoralis might also suggest that the former 
is similarly expanding its range. 

In the region of parental sympatry, O. x blakei often occurs with only one 
of the parents or even with neither of the parents in close proximity (Pike 1970), 
apparently persisting through its rhizomatous habit and ranging more widely 
through fertile cypselae. When found with only one of its putative parents, O. 

x blakei is more often associated with O. nemoralis (Pike 1970; Brouillet & Si- 
mon 1981), perhaps because of their greater similarity in habitat. 

(2) The parentage of O. acuminata in the hybrid O. x blakei is usually re- 
flected by at least some degree of foliar toothing. Entire leaves occur rarely in O. 
x blakei, but Fl individuals may sometimes be closer in morphology to the en- 
tire-leaved O. nemoralis than to the toothed-leaved O. acuminata (Pike 1970; 
Hill & Rogers 1973). Evidence for introgression between O. x blakei and its par- 
ents was observed in nature by Pike (1970) and experimentally duplicated by 
Hill and Rogers (1973), but the overall morphology of Gerdes 2210 is more simi- 
lar to an Fl, except for the entire leaves. Leaves of 2210 are elliptic—those of O. 
xblakei vary in shape from oblanceolate (broadest above the middle) to elliptic. 

The cypselae of Gerdes 2210 bear a mixture of sessile glands and numer- 
ous, slender, spreading-ascending, sharp-pointed hairs and are closely similar 
in vestiture to those of Oclemena reticulata (Ell) Nesom, a species of the coastal 


NESOM 631 


plain of Alabama, Florida, Georgia, and South Carolina, far-removed in geography 
and habitat from the other species of the genus. This cypselar vestiture is unlike 
that of O. xblakei from elsewhere in its range or that of its parental species, 
which is glandular but usually otherwise glabrous; cypselae of O.acuminata rarely 
may be sparsely strigose (e.g., Pendleton Co., W. Va., Musselman 3894—NCU), and 
Semple et al. 1996) describe the cypselae of O. nemoralis as “sparsely strigose.” 
The differences in non-glandular cypselar vestiture and the leaf margins are 
morphological differences separating Gerdes 2210 from O. x blakei, but the close 
correspondence in other morphological features suggests that an identification 
as O. x blakei is a reasonable hypothesis for the Keweenaw Peninsula plants. 


Identification as a separate species 

In view of tentative evidence against a hypothesis of hybridity between 
Oclemena nemoralis and O. acuminata (considering the anomalous achene 
vestiture, unusual leaf morphology, and the unlikelihood of the establishment 
of 2210 through long-distance dispersal) for the origin of 2210, a hypothesis 
that this population represents an independent evolutionary branch of 
Oclemena becomes plausible. If this were the true origin, recognition of 2210 at 
specific rank would be appropriate. Current evidence, however, does not con- 
vincingly support or eliminate either of the proposed hypotheses alternative 
to the originally proposed intergeneric hybridization. All three competing hy- 
potheses will be further investigated after an upcoming field season, when it 
will be possible to make observations on pollen and fruit fertility and collect 
fresh material for molecular analysis. 


ACKNOWLEDGMENTS 


[am grateful to the staff of MICH for lending the collections from Houghton 
County, the staffs of GH and NCU for accommodation while working there, to 
Luc Brouillet for his comments on the plants involved and on an early version 
of the manuscript, and to an anonymous journal reviewer. 


REFERENCES 


Brouittet, L. AnD J.-P. Simon. 1981. An ecogeographical analysis of the distribution of Aster 
acuminatus Michx.and A. nemoralis Aiton (Asteraceae: Astereae). Rhodora 83:521—550. 

Geroes, L.B. 1998 [2000]. Aster nemoralis and the apparent hybrid Aster nemoralis xAster 
umbellatus in Houghton County, Michigan. Michigan Bot. 37:1 7-20. 

Hitt, L.M. 1976. Morphological and cytological evidence for introgression in Aster 
acuminatus Michx.in the Southern Appalachians. Castanea 41:148-155. 

Hitt, L.M. and O.M. Rocers. 1970. Chromosome numbers of Aster blakei and A. nemoralis. 
Rhodora 72:437-438. 

Hitt, LM. and O.M. Rocers. 1973.Chemical, cytological and genetic evidence for the hybrid 
origin of Aster blakei (Porter) House. Rhodora 75:1—25. 


ey 


632 BRIT.ORG/SIDA 19(3) 


Nesom, G.L. 1993. Taxonomy of Doellingeria (Asteraceae: Astereae). Phytologia 75:452—462. 

Nesom, G.L. 1994a. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), 
emphasizing the New World species. Phytologia 77:141-297. 

Nesom, G.L. 1994b. Hybridization in the Astereae (Asteraceae). Phytologia 77:298-30/7. 

Nesom, G.L. 2000.Generic conspectus of the tribe Astereae (Asteraceae) in North America, 
Central America, the Antilles, and Hawaii. Sida, Bot. Misc. 20:i-viii, 1-100. 

Pike, R.B. 1970. Evidence for the hybrid status of Aster Blakei (Porter) House. Rhodora 72: 
401-436. 

Sempte, J.C., J.G. CHMIELEWSKI, and C. Leeper. 1991. A multivariate morphometric study and 
revision of Aster subg. Doellingeria sect. Triplopappus (Compositae: Astereae): The Aster 
umbellatus complex. Canad. J. Bot. 69:256-276 

Semete, J.C., S.B. HeAarb, and C:S. XiAnc. 1996. The Asters of Ontario (Compositae: Astereae): 
Diplactis Raf., Oclemena E.L. Greene, Doellingeria Nees and Aster L. (including 
Canadanthus Nesom, Symphyotrichum Nees, and Virgulus Raf.). Rev. ed. Univ. of Water- 
loo Biol. Ser. 38:i-viii, 1-94. 

Voss, E.G. 1996. Michigan Flora. Part Ill, Dicots (Pyrolaceae—Compositae). Cranbrook Insti- 
tute of Science Bull.61 and Univ. of Michigan Herbarium, Ann Arbor. 

XIANG, C. and J.C. Sempte. 1996. Molecular systematic study of Asters.lat.and related genera 
(Asteraceae: Astereae) based on chloroplast DNA restriction site ee and mainly 
North American taxa.In:D.J.N. Hind and H.J.Beentje,eds.Com| Systematics. Proc. 
Interntl. Compositae Conf, Kew, 1994. Vol. 1, pp. 393-423. 


BIBLIOGRAPHICAL NOTES ON THE 
PUBLICATION OF WOOLWARD’S MONOGRAPH 
ON THE GENUS MASDEVALLIA (ORCHIDACEAE) 


Guido J. Braem 


Schlechter Institute 
Naunheimer Str. 17 
D-35633 Lahnau, GERMANY 
and 


Research Associate 


C ahiforniaq Acad, omy of Sciences 


ABSTRACT 
The history of publication of Florence H. Woolward’s monograph on the genus Masdevallia is dis- 
cussed. Publication dates are given for all parts of the work. 


L. ry ee | 


KEY 2 Masdeva 


lia, History of Botany, Botanical Art, Woolward 


RESUMEN 


aa 


Se discute la historia de la publicacion de la monografia de Florence H. Woolward del género 


Masdevallia. Se dan fechas de publicacion de todas las partes del trabajo. 


Miss Florence Helen Woolward (1854-1930) was employed or commissioned 
by the Marquess of Lothian to work and paint in his collection at Newbattle 
Abbey. Contrary to general belief, Miss Woolward was trained neither in Botany 
nor as an artist, as can be deduced from her own words in the introductory chap- 
ter to the book “... although being neither a trained artist nor botanist”. It stands, 
however, without doubt, that Florence Woolward was one of the best botanical 
artists known. The original drawings for the Masdevallia monograph are kept 
at the Natural History Museum in London 

In The Gardeners’ Chronicle for October 25, 1890, the following announce- 
ment was published: 


“The Genus Masdevallia—The first part of the Marquess of Lothian’s Monograph of species growing 


in his collection at Newbattle, is announced as nearly ready. The work will contain hand-coloured 
lithographs and engravings of every available species. The descriptions and plates are by Miss 
Woolward, oe notes by Consul Lehmann. The work will be issued in parts of imperial quarto size, 
at the price of £1 10s. each part, the issue being limited to 250.” 

ON THE DATES OF PUBLICATION OF THE GENUS MASDEVALLIA 


Asso often, the correct dates of publication cannot be obtained from the primary 
source, as the original wrappers merely state the year of publication. Experi- 
ence has shown that such indications are not necessarily correct. Therefore, 
secondary literature must be consulted. Unfortunately, not all authors bother 
to investigate the bibliography in detail, and very often the entire Masdevallia 


SIDA 19(3): 633 — 637. 2001 


634 BRIT.ORG/ SIDA 19(3) 


work of Woolward is erroneously considered to have been published in 1896. 
play et the one literature, however, reveals that the Woolward 
usly issued in 9 parts. Parts one (1) through eight (8) con- 
sisted of 10 ae cack accompanied by the pertinent texts. With part five (5), 
a preliminary index was published (Rendle 1894). Obviously this preliminary 
index would have been discarded at the time the work was completed and bound 
into its final form. Part nine (9) consisted of seven plates, their text, and some 
additional texts such as he title page, the index, and the “List of species now 
excluded from the genus Masdevallia.” 

Stafleu and Cowan (1988) based their publication dates on the notices of 
reception published in the Journal of Botany (J. Bot.) There seem, however, to be 
discrepancies between those data and the information found in The Gardeners’ 
Chronicle, probably resulting from the fact that The Gardeners’ Chronicle was 
published on a weekly basis and is therefore obviously more exact. Both the 
Woolward monograph and The Gardeners’ Chronicle were published in Lon- 
don, and it may be assumed that each installment of the Woolward monograph 
reached the Chronicle without delay. 

Based on the data retrieved from notes of reception and from reviews in 
The Gardeners’ Chronicle, the following dates of publication have been deter- 
mined (Table 1, 2). The full citation as taken from an original copy is: 


The genus Masdevallia. Issued by the Marques of Lothian, K.T., chiefly from plants in his collection at 
Newbattle eee: plates and descriptions by Miss Florence H. Woolward; with additional notes by 
FC. Let Consul in the Republic of Colombia)- With 87 an and 61 woodcuts from 
chirps index ved Map. 


The book is generally referred to as “Woolward, The genus Masdevallia 


Taste 1, Probable publication dates for Woolward, The genus Masdevallia 


Part — Publication dates Data from The Gardeners’ Chronicle Notices in Journal of Botany 

1 1890 (December) Note - se ee aaa 1890 Note: 30 January 1891 
Review: 10 Janua Review: March 1891 

2 1891 (August) ve of Serene ens 189] September 1891 

3 1892 (August) Note of reception: 3 September 1892 November 1892 
Review: 10 September 1892 

4 1893 VJuly) 5 August 1893. Reference in note on August 1893 
leeeien fragrans and Masdevallia 
aristata —-“The recently issued part IV ... 

5 1894 (February) en February 1894 No. 32, 1894 

6 1895 (February) = Review:2 March 1895 ? 

7 1895 (October) Review:9 November 1895 October 1895 

8 1896 (October) Review: 24 October 1896 October 1896 


“We heartily Seneca Miss opine 
on the completion of h ork 


9 1896 (October Review: 24 October ice above) October 1896 


— 


BRAEM, WOOLWARD’S MONOGRAPH OF MASDEVALLIA 


Taste 2. Publication dates of plates arranged alphabetically according to plant names. 


Plant Name Part Publication date 
01. Masdevallia abt Reichenbach fil. 4 July 1893 
02. Masdevallia amabilis Rei ia ] December 1890 
03. Masdevallia aristata Rodrigue 4 July 1893 
04. Masdevallia i Reichenbach fil. 3 August 1892 
05. M Ili Reichenbach fil. 9 October 1896 
06 asdevallia eichenbach fil. 8 October 1896 
07 Hee barlaeana aeichen bach fil. 8 October 1896 
08. Mastenaias bella Reichenbach fil. ] December 1890 
09. Masdev loptera Reichenbach fil. vi October 1895 
10. Masdevallia calura Reichenbach fil. 8 October 1896 
11. Masdevallia ye glossa Reichenbach fil. 8 October 1896 
12. Masdevallia carderi Reichenbach fil. 3 August 1892 
13. Masdevallia caudata sinless 3 August 1892 
14. Masdevallia chestertonl Reichenbach fil. 1 ecember 1890 
15. Meee! We f Reichenbach fil. 5 February 1894 
16. imaera var. backhousiana Veitch 5 February 1894 
17. ee allia chimaera var. roezlii Veitch 5 February 1894 
18. M llia civilis Reichenbach fil 2 August 1891 
19. Masdevallia inea Linden 3 August 1892 
20. Masdevallia coriacea Lindley 3 August 1892 
21. Masdevallia corniculata oo ales fil. 2 August 1891 
22. Masdevallia cucullata Lindle 2 August 1891 
23. Masaevaui cupularis Reichenbach fil. 8 October 1896 
24. Rodrigues 5 February 1894 
25. Masa d valle davisti Reichenbach fil. 3 August 1892 
26. Masdevallia Reichenbach fil. 5 February 1894 
27. camasan ide cues Reichenbach fil. 6 February 1895 
28. M \ ichenbach fil. 1 December 1890 
29. Masdevall id veythacet peleneneden fil. 8 October 1896 
30. M valli Reichenbach fil. 3 August 1892 
31. Masceva allia floribunda neley 4 July 1893 
32. Masdevallia Healer Woolward 6 February 1895 
33. Masdevallia fragrans Woolward 4 July 1893 
34. sdevallia fulvescens Rolfe 9 October 1896 
35. Masdevallia g Reichenbach fil. 7 October 1895 
36. Masdevallia lata Reichenbach fil. 4 July 1893 
37. nee hieroglyphica nplenenoaes fil. 6 February 1895 
38. evallia | Reichenbach fil. 5 February 1894 
39. Hie infracta Lindley 2 August 1891 
40. Masdevallia ionocharis = eitath fil, 4 July 189 
41. Masdevallia laucheana Kraenzlin ex Woolward 8 October 1896 
42 i raeata leontoglossa Reichenbach fil. 2 August 1891 
43 asdevallia macrura Reichenbach fil. ] December 1890 
44 He sdevallia maculata Klotzsch 2 August 1891 
45. Masdevallia marginella Reichenbach fil. 9 October 1896 
46. Masdevallia mel Reichenbach fil. 4 July 1893 


636 BRIT.ORG/SIDA 19(3) 
Plant Name Part Publication date 

47, Masdevallia militaris Reichenbach fil. July 1893 

48. Masdevallia mooreana pauls fil. February 1895 

49, Masdevallia eichenbach fil. February 1895 

50. Masdevallia nidifica peichenback fil. February 1894 

51. Masdevallia nyct Reichenbach fil. October 1896 


Masdevallia o ‘brieniana Rolfe 

ane eee pe eee fil. 
ort. ex Woolward 
Masdevalli Poaien itha Reichenbach fil. 
Masdevallia pachyura Reichenbach fil. 
Masdevallia peristeria Reichenbach fil. 
Masdevallia picturata Reichenbach fil. 
oaeia Senne Reichenbach fil. 
Masdev Reichenbach fil. 
pees ia oer iceps Reichenbach fil. 
Masdevallia pusilla Rolfe 

Masdevallia racemosa Lindley 
Masdevallia radi 
Masdevallia reichenbachiana Endres 
Masdevallia rolfeana Kraenzlin 
NMacdevallia 


- 
gd Lindale 


= 


Reichenbach fil. 


Masdevallia schlimii Linden 
Masdevallia schroederiana eS 
Masdevallia simula Reichenbach fil. 
Masdevallia striatella ee eae a) fil. 
Masdevallia torta Reichenbach fil. 
Masdevallia tovarensis Reichenbach fil. 
Masdevallia triangularis Lindle 
Masdevallia triaristella Reichenbach fil. 
Masdevallia triglochin Reichenbach fil. 
Masdevallia trinema Reichenbach fil. 
Masdevallia troglodytes Morren 
Masdevallia uniflora Ruiz & Pavon 
Masdevallia veitchiana Reichenbach fil. 
Masdevallia velifera Reichenbach fil. 
Masdevallia ventricularis Reichenbach fil. 
Masdevallia vespertilio Reichenbach fil. 
Masdevallia wageneriana Lindle 


= 
— 


L 


Masdevallia xanthina Reichenbach fil. 
Masdevallia yauaperyensis Rodrigues 


al. 

Masdevallia wendlandiana Reichenbach fil. 
/ 
Ii 


MEWDWOANRFUNNONWNHH DH ON HAUHNNYNBKRONWON BND DAANOUDAAA 


July 1893 
October 1895 
August 1891 


December 1890 
August 1891 
October 1896 
December 1890 


February 1895 
July 1893 
February 1894 


REFERENCES 


Anonymous. 1890. The genus Masdevallia. Gard. Chron. ser. 3, 8(209):754. 
Anonymous. 1891. The genus Masdevallia. Gard. Chron. ser. 3, 10(244):252. 


BRAEM, WOOLWARD’S MONOGRAPH OF MASDEVALLIA 637 


Anonymous. 1892. The genus Masdevallia. Gard. Chron. ser. 3, 12(297):279. 

Renote, A.B. 1891. Book notice. Gard. Chron. ser. 3, 9(211):52. 

Renote, A.B. 1892. Book notice. Gard. Chron. ser. 3, 12(298):310. 

Renote, A.B. 1893. Masdevallia fragrans and M.aristata. Gard. Chron. ser. 3, 14(345):162-163. 

Renote, A.B. 1894, Book notice. Gard. Chron. ser. 3, 15(374):240. 

Renote, A.B. 1895. Notices of books. Gard. Chron. ser. 3, 17(427):270-271. 

Renote, A.B. 1895. Book notice. Gard. Chron. ser. 3, 18(463):550. 

Renbte, A.B. 1896. Book notice. Gard. Chron. ser. 3, 20(51 3):490. 

StaFveu, FA.and R.S.Cowan. 1988. Taxonomic literature, second edition, 7:452.Regnum veg. 
116. Bohn, Scheltema & Holkema, Utrecht/Antwerpen, dr. W. Junk b.v., Publishers, The 
Hague/Boston. 


= 


638 BRIT.ORG/SIDA 19(3 
Book REVIEW 
Ken E. Rocers. 2000. The Magnificent Mesquite. (ISBN 0-292-77105-3, hbk.). University 
of Texas Press, PO. Box 7819, Austin, TX 78713-7819, U.S.A. (Orders: http:// 
www.utexas.edu/utpress, 512-471-4032). $24.95, 167 pp, 18 color photos, 
2 line drawings, 5 maps, l2 tables, 51/2" x 9" 


Contents.— 
« Introduction 
* Mesquite: What Is It? 
« Predators of Mesquite 
* Mesquite in Texas and the Southwestern United States 
* The Uses of Mesquite 
* The Mesquite in Verse 
* Worldwide Aspects of Prosopis 
* Texas Ebony-Mesquite’s Astonishing Cousin 


« Mesquite’s Future 

« Appendix |. The Genus Prosopis and Its Species 

* Appendix 2. Sources of Information 

« References and Further Reading 

* Index 
This book summarizes the biology, ecology, uses, and management of mesquite—an overview of in- 
terest to scientists and laymen alike. This broad interest reflects, at least in part, a shift in public 
perception of these plants, which used to be regarded merely as pests but now are finding wide ac- 
ceptance for their utility and aesthetic qualities. A table near the end—“Future Areas of New and 


—notes the fi ollowing uses: fine veneers flooring, ornamentals, desert 


Increased Mesquite Utilization” 
reclamation, feedstock for chemicals and pharmaceuticals, wo 

lastics, biomass for energy feedstock, pods for human and eee consumption, packaged fire- 
wood, and lumber. The prSoiater ranges from a recipe for jelly and instructions on constructing a 
mesquite wood floot predator control, life cycle details, and the physical, 
mechanical, and chemical Sens of the wood. The book is beautifully bound (in handbook size 
e UT Press series) and the contents are nicely organized. Reading is 


into wood 


like “The Pecan Tree” in the 
easy and the whee tee can nbc Hay Teed in detail in a couple of hours. 
eueieice but is skewed toward the technical sIGes in paca a- 
tion, which perhay lerli ious disapy in the mesquite book. The ter esquite 

| | ite of the southwestern USA Prosopisglandutosa 
not impossible to 


his reader 


used tO refer 

as to all species of the genus resp ao Page after page, it is difficult i 
whole genus is being referred to. | surely won't be the only one 

| provide amendments in the next edition of the 


as well 
tell whether a single species or the 
frustrated by this and hope that a. author wil 
‘compleat mesquite. —Guy L. Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, 
TX 76102-40060, U.S.A. 


SIDA 19(3): 638. 2001 


GNAPHALIUM EXILIFOLIUM (ASTERACEAE: 
GNAPHALIEAE) IN 
COLORADO AND SOUTH DAKOTA 


Guy L.Nesom 
Botanical Research Institute of Texas 
509 Pecan Street 
Fort Worth, TX 76102-4060, U.S.A. 


Stricter definition of the genus Gnaphalium L. (e.g., Anderberg 1991) leaves the 
genus in North America north of Mexico with three species: G. exilifolium A. 
Nels., G. palustre Nutt., and G. uliginosum L. The lectotype species of 
Gnaphaliumsensustrictois G. uliginosum—discussion of this choice rather than 
Pseudognaphalium (Gnaphalium) luteo-album (L.) Hilliard and Burtt is given 
in Jeffrey (1979), Hilliard and Burtt (1981), and McNeill et al. 1987). The remain- 
der of the species of Gnaphalium sensu lato in the United States and Canada 
are divided among the segregate genera Pseudognaphalium Kirpichn,, Euchiton 
Cass., Gamochaeta Wedd., and Omalotheca Cass. 

Gnaphalium exilifolium has been treated as conspecific with G. uliginosum 
in various floristic accounts of the western United States (e.g., Great Plains Flora 
Association 1986; Weber & Whittman 1994) but regarded as a separate species 
in others (e.g., Harrington 1954; Welsh et al. 1993; Cronquist 1994). Examina- 
tion of collections of these plants at ARIZ, ASU, BRIT, GH, NCU, NMC, TEX, 
and US have provided perspective on the distinction of G. exilifolium. The three 
species of Gnaphalium sensu stricto that occur in the United States can be iden- 
tified by the contrasts in the key below. 


— 


. Leaves spatulate to Ser eerie, 3-8(-10) mm wide; bracts of capitu- 

lescence oblanceolate to obovate, longest 4-12 mm x 1.5-4 mm, shorter than or 

equalling to slightly surpassing va glomerules; inner phyllaries nee tl 

with blunt apex halium palustre 
. Leaves nest to natrowiy Peer 0. os 3 mm wide; bracts of =e ees 

linear to obla te or obovate, longest 5-25 mm x 0.5-2 mm, distinctly longer 

than the glomerules; inner ie aan with acute a 

2. Leaves oblanceolate, less commonly linear, the alg ]- s cm; capitulescence 

terminal and capitate, rarely with axilllary glomerules below; bracts subtending 

capitulescence linear to oblanceolate or obovate, 1-2 mm wide, the longest 

mostly 5-15 mm. Gnaphalium uliginosum 
. Leaves linear, the largest 0.4-5 cm; capitulescence commonly spike-like, wit 

numerous sessile axillary glomerules; bracts subtending capitulescence linear, 

0.5-1 mm wide, the longest mostly 10-25 mm. Gnaphalium exilifolium 


at 


NR 


The account of the United States, Canada, and Greenland flora by Kartesz (1999) 
records Gnaphalium exilifolium in New Mexico, Arizona, Utah,and Wyoming, 


SIDA 19(3): 639 — 641. 2001 


640 BRIT.ORG/SIDA 19(3) 


and the species also occurs in northern Chihuahua, Mexico (Nesom unpub- 
lished). Records also exist for its occurrence in Colorado and South Dakota—as 
clarified below. Although recent accounts of the flora of Colorado (Weber & 
Whittman 1994, 1996a, 1996b) and South Dakota (Dorn 1977, Van Bruggen 1985) 
have not included G. exilifolium as an accepted species, collections of it (as de- 
fined above) have been made from both states. 


COLORADO. Montrose Co.: Cimarron, 2200 m, 8-10 Sep 1917, Eggleston 14200 
(GH): 

Recognition of the presence G. exilifolium in Colorado results from a more 
focused species concept rather than from its rarity. Gnaphalium uliginosum, 
with which it has been confused, also occurs in Colorado. The distinction between 
the two taxa was correctly made by Harrington (1954, p. 624), although he sup- 
plied the caveat that “all our plants lof G. exilifolium] may actually bea part of G. 
uliginosum L.” Collections at GH of G. exilifolium from Colorado also are from the 
following counties: Clear Creek, Delta, Lake, Larimer, Mineral, Park, and Summit. 


SOUTH DAKOTA. [Lawrence Co.:] Black Hills National Forest, Yellow Creek, 
near Kirk, banks of Ice Pond, 27 Aug 1910, Murdoch 4337 (GH). 

McIntosh (1931) identified G. exilifolium as occurring in the Black Hills of 
South Dakota, although other botanists have apparently identified the same 
plants as G. uliginosum. The latter was specifically noted to occur in Custer 
and Lawren unties, South Dakota (Great Plains Flora Association 1976). The 
cited Murdoch collection unequivocally confirms the occurrence of G. 
exilifolium in South Dakota; the possibility of the existence of G. uliginosum 
there has not been eliminated. Neither species is documented for South Dakota 
in the Forest Service repository at RM. 

The nomenclature details for Gnaphalium exilifolium are as follows. 


Gnaphalium exilifolium A. Nels. [nom. nov. for G. angustifolium|, Bull. Torrey 
Bot. Club 29:406. 1902. Gnaphalium angustifolium A. Nels., Bull. Torrey Bot. Club 
26:357. 1899 (non Lam. 1788). 

Gnaphalium grayii A. Nels. & Macbr. [nom. nov. for G. strictum], Bot. Gaz. 61:46. 1916. 
Gnaphalium strictum A. Gray, Pacif. R.R. Rep. 4:110. 1858 (non Moench 1794; non Roxb. 
1814). 


ACKNOWLEDGMENTS 


| am grateful to the staff of the cited herbaria for loans and accommodation 
during recent visits, to Mark Gabel for comments on the manuscript, and to 
Ron Hartman for checking the RM collections. 


REFERENCES 


AnperberG, A.A. 1991. Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). 
Opera Bot. 104:5-195. 


NESOM DAKOTA 641 


Cronauist, A. 1994. Intermountain flora. Volume Five, Asterales. The New York Botanical 
Garden, Bronx. 

Dorn, R.D. 1977. Flora of the Black Hills. Published by the author, Cheyenne, Wyoming. 

Great PLAINS Flora Association (R.L. McGregor, coordinator; T.M. Barkley, ed.). 1976. Atlas of 
the flora of the Great Plains. lowa State University Press, Ames. 

GreAT PLAINS FLORA ASSOCIATION. 1986. Flora of the Great Plains. Univ. Press of Kansas, Lawrence. 

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

Hiviarb, O.M. and B.L. Burt. 1981. Some generic concepts in Compositae-—Gnaphaliinae. 
Bot. J. Linn. Soc. 82:181-232. 

Jerrrey, C. 1979, Note on the lectotypification of the names Cacalia L., Matricaria L.and 
Gnaphalium L.Taxon 28:349-351. 

Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the 
vascular flora of the United States, Canada, and Greenland. First Ed. In: J.T. Kartesz and 
C.A. Meacham. 1999. Synthesis of the North American flora, Ver. 1.0. North Carolina 
Botanical Garden, Chapel Hill. 

McIntosh, A.C. 1931.A botanical survey of the Black Hills of South Dakota. Black Hills Engi- 
neer 19:159-276 [reprinted Black Hills Engineer 28:3-74. 1949]. 

McNeil, J., EA. Ooett, L.L. Consaur, and D.S. Katz. 1987. American code and later 
lectotypifications of Linnaean generic names dating from 1753:A case study of dis- 
crepancies Taxon 36:350-401. 

VAN Bruacen, T. 1985. The vascular plants of South Dakota (ed. 2). lowa State University 
Press, Ames, lowa. 

Weeer, W.A. and R.C.Wuittman. 1994. Catalog of the Colorado flora: A biodiversity baseline. 
University Press of Colorado, Niwot. 

Weeer, W.A. and R.C. WuitTmaN. 1996a. Colorado flora: Western slope. University Press of 
Colorado, Niwot. 

Weeer, W.A. and R.C.WuHiTTmaN. 1996b. Colorado flora: Eastern slope. University Press of Colo- 
rado, Boulder. 

Wetsh, S.L., N.D. Atwoon, S.Gooprich, and L.C. Hicains (eds.). 1993.A Utah flora (ed. 2, revised). 
Print Services, Brigham Young University, Provo, Utah. 


BRIT.ORG/SIDA 19(3) 


Book REVIEW 


KEITH RUSHFORTH. 1999. A photographic guide to the trees of Britain and Europe. 
Photographs assembled by FLPA (Frank Lane Picture Agency). (ISBN 0 00 
2220013 9, pbk.). Harper Collins Publishers, 77-85 Fulham Palace Road, 
London Wo6 8JB, U.K. (Orders: www.trafalgarsquarebooks.com, www. 
fireandwater.com, 800-423-4525). £16.99 British pounds, as listed on the 
cover; $14.99 on website listing. [= ca. $24.00 US dollars, or ca. $21.00], 825 
+ color photos, 3 3/4" x 71/2" 

“The coverage includes the whole of Europe where plants make trees. This ranges from the Mediter- 

ranean north to beyond the Arctic circle, and from the Urals to the Atlantic coast and islands. ... The 


coverage includes pot native ane erognees (or exotic) trees.” “Nearly 1,200 tree species (plus a 
book, with over 825 illustrated and 750 described 


number of pre 
in detail.” The author notes that “specialist collections such as those of major arboreta or tree collec- 
tions ... will contain a number of trees not featured here,” but a — number of hybrids and selec- 
tions are described under the “Varieties” heading in the commenta 

relatively short, up-front and to-the-point a tie cae a aieser classification and 
nomenclature and gives an overview of aan are defined. A workable “Key to the Colour Plates” 
leave ate and buds alternate, pp. 320-39; leaves digitate 


gives a range of illustrated choices (e.g 
and buds opposite, pp. 310-11) to lead one into ‘the section of photos where a tree in question might 
be found. There also is a similar ‘sort mechanism (using color photos) to fruit types. The pictures are 
small but excellent, witha B00 mix of habit and leaf, flower, and bark details: peace in the Peon 


section are arranged by | I category, but the 
arranged alphabetically by pene aa species. Larger genera are intro uced a) an informal key that 
t f commentary: De- 


separates species into groups. Each species has some or all of t 
scription, Range, Habitat, Varieties, Synonyms, Similar species. Comicn ean variation in chromo- 


some number are a consistent seca As might be expected in a treatment of this scope, some of the 
cultivati appearance, etc., appear to be more first-hand than others. 


} + 
“his is an excellent eee one of great utility for English tree-lovers, and because so 
many of these species are likely to be found in cooler climes of North America, Yanks also will ap- 
preciate it. tis a relatively small-sized book (9.5 x 19 em; advertised as “The only photographic field 

guide you can use in the wild”) but thick with 1336 pages! The pages are thin but seem to be durable; 

same for the binding, and it comes in a plastic cover. The advertised price seems remarkably low, 
apparently designed to get copies into wide distribution. It is one of a Collins Wildlife Trust series 
that includes guides to birds, butterflies, insects, wildflowers, and mushrooms of the same region. 

—Guy L. Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, 


ae) 


SIDA 19(3): 642. 2001 


DOCUMENTED CHROMOSOME NUMBERS 200L 1. 


CHROMOSOME NUMBER OF 
LUPINUS HAVARDII (FABACEAE) 
B.L. Turner A.M. Powell 
Plant Resources Center Department of Biology 
University of Texas Sul Ross State University 
Austin, TX, 78713, U.S.A. Alpine, TX, 79832, USA. 


Chromosome numbers for the American taxa of the large genus Lupinus are 
mostly diploid (2n=24) or tetraploid (2n=48). However, two species of biennial 
(or winter annuals) native to central Texas and closely adjacent Mexico, L. 
subcarnosus Hook. and L. texensis Hook., were found by Turner (1957) to be uni- 
formly diploid with 2n=36. Subsequently, Turner (1994) noted that the closely 
related winter annual, L. havardii S. Wats. of Trans-Pecos, Texas and closely ad- 
jacent Mexico, appeared to belong to this complex, but chromosome counts for 
the species were unknown. To remedy this, bud material for L. havardii was 
obtained by the senior author in the early spring of 2001, these subsequently 
counted by the junior author. 

Meiotic counts were obtained from natural populations of L. havardii us- 
ing the methods of Turner (1957). Voucher specimens are on file at SRSC and 
TEX, these obtained at the following localities: 


Presidio Co.: 2.1 road mi N of Shafter along Hwy 169, 23 Feb 2001, Turner 21-2. 

Presidio Co.: 7 road mi E of Presidio along Hwy 170, 23 Feb 2001, Turner 21-4. 

Presidio Co.: 14 road mi E of Presidio along Hwy 170, 23 Feb 2001, Turner 21-7. 

All counts were determined to be 2n=36 (18 bivalents), except for collections 
21-2 and 21-7, both of which showed circa counts of n=18 bivalents. Turner 
(1994) noted that the chromosome count of L. havardii was “likely to be 2n=18 
pairs since the taxon L. havardii seems closely related to L. texanus,” Which is 
verified by the present paper. 


REFERENCES 


Turner, B.L. 1957. The chromosomal and distributional relationships of Lupinus texensis 
and L. subcarnosus (Leguminosae). Madono 14:13-16. 

Turner, B.L. 1994. Species of Lupinus (Fabaceae) occurring in northeastern Mexico (Nuevo 
Leon and closely adjacent states). Phytologia 76:290-302. 


SIDA 19(3): 643. 2001 


644 BRIT.ORG/SIDA 19(3) 
Book REVIEW 
Gi. Netson. 2000. The Ferns of Florida. (ISBN 1-56164-193-6, hbk; 1-56164-197-9, 
pbk). Pineapple Press, P.O. Box 3899, Sarasota, FL 34230, U.S.A. (Orders: 
www pineapplepress.com). $27.95 hbk, $19.95 pbk, 256 pp, 204 color photos, 
ox? 


A treatment of Florida ferns is useful and welcome—among U.S. states and territories, only Hawaii 
and Puerto Rico have greater numbers of fern species. ee records 164 spontaneously occurring 
axa (153 species, 9 Me brids) within the state's pore | 123 of the taxa are considered native. 42 taxa 


are endangered; 6 are threatened; 3 ar ly exploited. 
The Introduction provides “a brief history of fern study in Florida, a note about the importance 


] | 
nomenciature and 


of fern conservation, a eee of fern classification, a treatment of botanice 
descriptive vocabulary, t f the fern life cycle, a brief discussion fern hybridization, 
and a glossary.” At the end of he book are these: 

* Appendix |. Checklist of Florida Pteridophytes Included in the = rent Volume. 
ix 2. Florida Pteridophytes According to Wunderlin (199 
orida’s Regulated vee ade 


ey 


« Appendi 
. Appendix 3. Pter idophyte s Included on F 
* Appendix 4. Where to Find Ferns in Florida. 


* Bibliography and Inde 
The fern taxa are arrangec ed alphabetically by family, genus, and species, as claimed most ap- 
la for a field guide. For each aac nee is a short technical description, detailed notes on 
habitat, distribution i the gasmall map of the state, divided into four regions), 
early Florida collections, unsuccessful searches for rare species, presumed extirpation, features for 
field recognition and distinction from similar species, hybrids, nativity, spread of non-natives, deri- 
vation of the epithet, references to pertinent literature, and formal categories of rarity from listings 
re. A selection of keys provides distinction for the families 


by the Florida Department of Agricultu 
treated, genera of Dryopteridaceae, genera of Poly podiaceae, genera of Pteridaceae, species of Asple- 
nium, species of Thelypteridaceae, and there is a tabular comparison of species of Marsilea. 

The 240. color photos (all taken by the author) are clustered together in the center of the book, 
ck and white photos are helpful. Pointed comments on habitat, 
ulness as a field guide. The 


—_ 


and scattered | lrawings and bla 
istribution, and aglincred features also support the book’s usef 
author's field experience and first-hand knowledge of the ferns are clear features of the discussions. 
Dimensions to be wished for in the Nelson volume would be keys for all genera with more than 
a single species — this would not be out of place for a field guide, especially as the keys convey the 
author's personal knowledge. Also, ease of reference would be gained (regained) by uniting the illus- 
trations with the species commentaries, although it obviously decreases the cost of production to 
cluster the numerous color illustrations. 
Nelson’s treatment, in company of the volume reviewed next 
contemporary step in the direction set by lar nce 1931) and Fe er ish orida (Long 
re now accounted for, the main 


(Wunderli id Hansen 2000), is 


the 
and Lakela 1976). If we can assume that the native ferns of the 
ne next undee ace will pel the discov ery OF apenas non-native species 


developments in Florida over t 
and the documentation of t 
—Guy L. Nesom, Botanical eee Institute of oe xas, 509 Pecan Street, Fort Worth 


, LX 76102-4060, 


SIDA 19(3): 644. 2001 


THE VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 
Mac H. Alford! 


Department of Botany 
ae see 
0338 


Durham, ee 0338, U.S.A. 
mha&@cornell.edu 


ABSTRACT 


A survey to document the vascular plants of Amite County, Mississippi, was conducted from 1997 to 
2000. Occupying 732 square miles, the county lies along an edaphic and physiographic transition 
zone from loess in the west to sandy loam and gravel in the east. Species composition changes with 
the gradient, and the county serves as a westernmost limit to several sues plants (eg., 
Gelsemium rankinii Small, Illicium eae Ellis). Amite County also harbors a residual ele- 
ment of north-south Pleistocene migrations in the loess hills (e.g., Adiantum se L., Pachysan- 
dra procumbens Michx., Cynoglossum virginianum L.). A total of 923 species were recorded, with 
new records for the state (Solidago auriculata Shuttlew. ex ne Clinopodium gracile (Benth.) 
Kuntze, aur ae psittacina Lehm., Ipomoea indica (Burm.f.) Merr., Photinia eis (Desf.) 

Kalkman) and a confirmation of recently published records of Physalis carpet Riddell and 
Dryopteris ae (Kunze) Small in Mississippi. 


RESUME 
Un arpentage pour documenter les ue culai lu Comté d’Amite, Mississippi, a été conduit 
de 1997 4 2000. Occupant 1874 kil carrés, le comté s’étend le long d’une zone de transition 
édaphique et phy caer du loess 4 l'ouest au terreau sablonneux et au gravier 4 l’est. La com- 
position d’espéces se modifie selon le gradient du terrain, et = comté sert de limite occidentale a 
plusieurs plantes du sud- -est (e.g., Gelsen mium anki Small, I floridanum J.Ellis). Le Comté 
d’Amite contient a un élé 1éi 5 {-sud parmi les collines 


de loess (e.g. Fens ties L., Pachysandra on um bes ns Michx., Cynoglossum virginianum L.). 
Un total de oe nes peces est jee coun avec de ou es uote pour Vetat sere 


Ipomoea indica (Burm.f.) Merr., Poti a serratifoli id eS ) tees et la Sogn des i 
mentations récemment publiées de ell et Dryopteris ludoviciana (Kunze) 
Small en Mississippi. 


INTRODUCTION 


Relative to most of the eastern United States, the vascular flora of Mississippi is 
poorly known (Duncan 1953; Pullen 1966; Bryson & Carter 1992; Bryson et al. 
1996; Sorrie & Leonard 1999). This problem was remedied to some extent by 
the Flora of Mississippi project funded by the National Science Foundation and 
executed by Samuel B. Jones, Jr. (University of Southern Mississippi, later GA), 
Thomas M. Pullen (MISS), and Ray Watson (MISSA). Despite a number of vital 


'Present address: L.H. Bailey Hortorium, Cornell University, 462 Mann Library, Ithaca, New York 14853 U.S.A, 
mha8@cornell.edu 


SIDA 19(3): 645 -699. 2001 


646 BRIT.ORG/SIDA 19(3) 


publications (e.g., Evans 1978; Jones 197 4a, 1974b, 1975, 1976; Pullen 1966; Pullen 
et al. 1968; Temple & Pullen 1968), however, the project was never completed 
and has only recently been reinvigorated (e.g., Bryson & Carter 1992). 

One of the major components of a statewide flora is information assembled 
from smaller-scale floras and the herbarium specimens they yield. Most of 
Mississippi's vascular floristic work has been concentrated in the northeast and 
southeast, near the major universities and in the longleaf pine belt, respectively. 
In fact, of all Mississippi vascular floras, only one (McCook 1982) is a survey in 
the southwestern part of the state. The adjacent area to the south in the Florida 
parishes of Louisiana can claim a much better record, but surprisingly the first 
flora of that region was not completed until 1972 (Allen 1972). 

The area chosen for this floristic study is Amite County, Mississippi. This 
choice is significant, considering the following reasons. First, Amite County is 
one of the counties that borders Louisiana in the southwest and as such is likely 
to produce new records merely by its periphery for the state of Mississippi. Sec- 
ond, Amite County occupies an interesting edaphic and physiographic loca- 
tion, serving as the transition zone from heavy loessal deposits in the west to 
the Citronelle-derived sandy loam and gravel of the east. The vegetation is in- 
fluenced by this gradient, and one can expect to find a variety of species, from 
those of mixed mesophytic areas in loessal deposits to those of coastal plain 
piney woods in the southeastern part of the county. Third, rare plants and plant 
communities can be observed and reported to the Mississippi Natural Heritage 
Program as a foundation for conservation work in the area. 


= 


THE PHYSICAL ENVIRONMENT 


Amite County, Mississippi, is located in southwestern Mississippi (Fig. 1). Amite 
County borders East Feliciana Parish, St. Helena Parish, and Tangipahoa Parish, 
Louisiana, on the south at 31.00°N; Wilkinson County, Mississippi, on the west 
at approximately 91.06°W; Pike County, Mississippi, on the east at approximately 
90.55°W, and Franklin County and Lincoln County, Mississippi, on the north at 
approximately 31.35°N. The county is generally rectangular but is irregular in 
the northwestern corner where the boundary follows a historical path of Fos- 
ter Creek and of the Homochitto River. 

Amite County is about 30 miles east to west by about 24 miles north to 
south, encompassing 732 square miles (1874 km? or 466,560 acres) (Milbrandt 
1976). The population in 1990 was 13,328 people (Clark 1997), with the only 
incorporated towns being Liberty, Gloster, Centreville, and Crosby. 

Climate.—Weather data supplied by the National Climatic Data Center 
(NOAA) for 1962-1997 are summarized in Table 1. Amite County is mild and 
humid with major weather influence from the Gulf of Mexico. The tempera- 
ture ranges from an average maximum of 9L.6’F (33.1'C) in July to an average 
minimum of 33.4’F (0.8'C) in January. The frost-free growing season lasts on 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 647 


Taste 1.Climatic data for Amite County, Mississippi. Based on data from the National Climatic Data 
Center, National Oceanic and Atmospheric Administration, for Liberty Station, 1962-1997. 


TEMPERATURE (°F) 
MONTH mean mean mean mean # of mean # of rainfall 
maximum minimum days with days with (inches) 
max = 90° min <32° 
January 45.2 56.9 33.4 0 15.1 5.85 
February 48.9 61.5 36.3 0 11.4 5.44 
March 57.0 69.9 44.] 0 49 6.42 
April 64.7 776 51.8 0.2 0.5 515 
May 715 84.0 58.9 3.9 0 5.17 
June 779 90.1 65.6 17.0 0 5.13 
July 80.3 91.6 68.9 23.2 0 5.29 
79,7 91.5 67.8 22.6 ¢) 448 
September 75.4 87.6 63.2 11.9 0 460 
October 64.9 79.3 50.4 2.0 0.5 3.06 
November 56.2 69.3 43.0 0 6.3 467 
December 48.5 60.7 36.3 0 13.9 6.21 
ANNUAL 64.2 76.7 51.6 81.3 51.6 61.47 


average from March 22 until November 3, or about 227 days. For the years avail- 
able, the temperature never dropped below 3'F (-16°C), and the average number 
of days per year with even a minimum temperature of 32’F (0'C) is 52. Average 
annual rainfall is 61.5 inches (154 cm), with the driest months in autumn and 
the wettest in winter and early spring. See Milbrandt (1976) for additional cli- 
mate information. 

Disturbance.—Forestry, agriculture, and fire are the major causes of veg- 
etation disturbance. Although many fires are set as part of forestry manage- 
ment practices, numerous fires are presumably natural. For the years 1959-1998, 
an average of 2078 acres of land burned each year in Amite County witha range 
of 402 to 6137 acres per year (Mississippi Forestry Commission, pers. comm.). 
Tornadoes are also a major natural cause of disturbance. From 1984 to 2000, 
there were 12 tornadoes, damaging an average area of 1144 acres each (Missis- 
sippi Forestry Commission, pers. comm.). More data on the effects of forestry 
and agriculture can be found under the section entitled Present Vegetation Types 
and Land Use. 

Topography.—Amite County lies within the generalized gulf coastal plain 
physiographic province (Fenneman 1938) and consists of rolling hills cut by 
several shallow valleys. The northwestern corner of the county deviates from 
this pattern somewhat by its dissection, and probably belongs to Fenneman’s 
(1938) loess hills physiographic belt. Elevation in the county ranges from 136 
feet above sea level along the Homochitto River to 500 feet in the northeastern 


648 BRIT.ORG/SIDA 19(3) 


part of the county. Large bluffs and ravines are only locally common, concen- 
trated in the Homochitto River basin and along the forks of the Amite River. 
Amite County is mapped on the USGS topographic maps Auburn, Berwick, 
Bewelcome, Busy Corner, Centreville, Crosby, Gillsburg, Gloster, Homochitto, 
Lake Tangipahoa, Liberty, Peoria, Smithdale, Street, and Terrys Creek 7.5' series. 

Numerous rivers and streams traverse the county (Fig. 1). The heavily dis- 
sected area in the northwestern part of the county belongs to the Homochitto 
River drainage, which flows into the Mississippi River. Small parts of the Buf- 
falo River drainage, which also flow into the Mississippi River, drain the county 
just west of Gloster. Most of the rest of the county lies within the Amite River 
drainage, which includes the West Fork Amite River, East Fork Amite River, 
Comite River, and Beaver Creek. This is the first drainage east of the Mississippi 
River that does not flow into the Mississippi River; it empties into Lake Maurepas 
of Louisiana. The Tickfaw River drains a part of the southeastern corner of the 
county, and the Tangipahoa River drains a very small part of the northeastern 
corner of the county. 

Geology. Most of Amite County rests upon reddish sedimentary deposits 
of sand, silt, clay, and gravel called the Upland Complex of the Citronelle For- 
mation (Bicker 1969; Spearing 1995). These sediments were probably deposited 
in the Pliocene or Pleistocene (~2 to 17 million years ago) as a broad alluvial 
fan of many streams (Doering 1935, 1956; Spearing 1995). Mineral composition 
(Spearing 1995), along with freshwater mussel (Stern 1976) and stonefly (Alford 
1998) distributions, suggest an Appalachian origin to these sediments 

The very northwestern corner of the county rests upon Miocene deposits 
that are, like much of the Citronelle deposits, obscured by a layer of loess. The 
Miocene deposits consist of green and bluish-green clay, sandy clay, gray silt- 
stone, and sand and are locally fossiliferous (Bicker 1969). Thought to be wind- 
blown deposits from glacial till (krinitzsky & Turnbull 1967), the loess—un- 
like most coastal plain soils—is rich in calcium and magnesium, has higher pH, 
and is able to retain a larger percentage of water (Caplenor 1968; Krinitzsky & 
Turnbull 1967). Loess deposits that are less than 8 ft (2.5 m) deep, however, like 
those in Amite County, are often leached of their calcium (Krinitzsky & 
Turnbull 1967). Areas with significant loessal soil are easily recognized from 
the adjacent coastal plain because they do not support the longleaf pine veg- 
etation so characteristic of much of the eastern gulf coastal plain. 

Larger deposits of loess are farther west in neighboring Wilkinson County, 
Mississippi, where they reach 100 feet deep (Krinitzsky @ Turnbull 1967). From 
this narrow band called the “Tunica Hills,” a gradually thinner layer of loess is 
deposited eastward, resulting in a gradient from thick loess rolling hills in the 
west to a thin layer of loess in the east called brown loam (Milbrandt 1976; 
Milbrandt & Hale 1968; Leggett et al. 1968). Although the loess bluffs and thick 
loess (>2.5 m deep) region has been recognized as a distinct physiographic belt 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 649 


mf 31 20° 


Ayo) Ald 


Amite County ; A J aX : 
East Feliciana Parish UISIA i Tangipahoa 
ari 


Leek se 


Fic. 1. Map of Amite County, Mississippi. Major highway pink, rivers in blue, incory ( ) in gray 
and the Homochitto National Forest in green. 


in the coastal plain province (Fenneman 1938; Holmes & Foster 1908; Lowe 1913), 
the thin loess region has generally been lumped into the “longleaf pine belt” or 
“southern pine hills” belt. 

Soils. —As just discussed under Geology, an important component of the soils 
of Amite County is loess (brown loam). The loess is thickest in the west and 
thins to an almost undetectable layer in the southeast (Fig. 2). 

According to Milbrandt (1976), there are seven major soil associations in 
Amite County (Fig. 2). Two associations, Gillsburg-Ariel-Peoria and Collings-Bude, 
are floodplain soils that are nearly level and poorly drained. The major upland 
associations, Providence-Bude and Providence-Ruston, consist primarily of small 
slopes with moderately well-drained soils based on loam (loess or loess-derived). 
One other major upland soil association is found along margins of major river 
dissection. This Saffell-Smithdale-Providence association, which is hilly and 
often consists of much gravel and silt, is of particular botanical interest, as will 
be noted later. In the southeastern corner of the county, the soil is less influenced 
by loess, tends to be better drained, and often has a reddish (instead of brown) 
color. It is named the Ora-Smithdale-Providence association. In the northwest- 
erncorner, the soil is influenced by loess but is highly dissected and often hasa 
clayey subsoil. This clayey subsoil frequently outcrops where loess has been 
eroded at high hills in the Homochitto River basin. These areas are where one is 
most likely to find longleaf pine and associated species in the region. 


THE BIOLOGICAL ENVIRONMEN 
Presettlement Vegetation and Early History—In 1720, only ae 700 European set- 
tlers lived in Mississippi, mostly confined to the areas around present-day 


650 BRIT.ORG/SIDA 19(3) 


Fi, 2. Soil f Amite C Mississippi (adapted Milbrandt, 1976). The Gillsburg-Ariel-Peoria association (flood 
plain) is in light green, Collings- -Bude (flood plain) i in dark green, Ora-Smithdale-Providence (upland coastal plain) in 
yellow, Providence-Bude in orange Biel Ruston oN Mapes in blue, Saffell-Smithdale-Providence (often 
gravelly) in red, and Smithdale 


ane J 


Natchez and Vicksburg (Gillis 1963). A massacre by the Natchez Indians in 1729 
eliminated most Europeans from the state, and settlement did not begin again 
until 1763 when Great Britain acquired the land from France. Settlement in- 
creased quickly during the Revolutionary War, when the land was under Span- 
ish control. The Spanish census of the region in 1792 showed 4690 people liv- 
ing in Mississippi, most still confined to areas near the Mississippi River (Gillis 
1963). However, in areas close to Amite County, there were already 112 families 
along Buffalo Creek and 136 families along the Homochitto River (Gillis 1963). 

Amite County was officially separated from Wilkinson County in 1809 
while Mississippi was still a territory (Clark 1997). The population continued 
to grow at an astonishing rate: the census of 1820 showed 6859 people in Amite 
County, 2833 of whom were slaves (Darby & Dwight 1836, U.S. Census 1820). In 
only four years (1816-1820), the population of neighboring Pike County grew 
from 330 families [1500? people] (Gillis 1963) to 5402 people (Darby & Dwight 
1836; U.S. Census 1820), also attesting to the rapid influx of people into the area. 
Furthermore, by 1840, 9511 people lived in Amite County (USS. Census 1840). 
Additional history of the county and surrounding area can be found in Casey 
and Otken (1948), Casey et al. (1950), and Casey (1957), 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 651 


Because Mississippi—like most of the United States outside of the original 
thirteen colonies and Texas—was surveyed ina standardized rectangular format 
(cf. Pattison 1957), the presettlement (or at least early) vegetation can be inferred 
from surveying records. “Marked trees” or “bearing trees’ at each quarter-section 
can be evaluated statistically for bias and used as a statistical sample of the 
woody vegetation. Many areas were surveyed long before settlement, but other 
areas—like that in question here—were settled at approximately the same time 
as the surveys. Amite County was surveyed in 1847 and 1848. Because the Land 
Ordinance of 1785 gave priority to previous surveys in these regions (Pattison 
1957), any areas already settled and surveyed by the time of the rectangular 
survey are readily noted ona political map asit regularly shaped sections. There- 
fore, one can safely assume that most early settlement (and disturbance of veg- 
etation) in Amite County was along major rivers and streams, as the irregular 
so-called “Spanish Land Grants” occur in these areas. 

A statistical analysis of the land survey records for determining presettlement 
vegetation has shown much promise (Bourdo 1956). However, the surveying of 
Amite County was accomplished by three different people with varying degrees 
of botanical expertise and note-taking, and the data are not statistically useful. As 
a coarse-grained tool, though, the data are interesting. For instance, In the south- 
western part of the county (Township | North, Range 2 East), 37% of corners 
were described as “oak, gum, beech” forest, 35% included “pine” in the descrip- 
tion, and only 7% were described as “timber all gone.” Most vegetation descrip- 
tions were permutations of oak (Quercus spp.), gum (Liquidambar styraciflua 
and/or Nyssa sylvatica and N. biflora), beech (Fagus grandifolia), pine (Pinus 
spp.), holly (Ilex opaca), bay (probably Magnolia gra ndiflora or M. virginiana), 
and hickory (Carya spp.). In the eastern part of the county (Township | North, 
Range 4 East), however, the survey repeats again and again “poor pine land,” 
probably of Pinus palustris, with only a few references to oak and hickory. 

A neighboring area has been studied in this fashion (Delcourt & Delcourt 
1974). Delcourt and Delcourt (1974) show that much of West Feliciana Parish, 
Louisiana, was dominated by a presettlement forest of Fagus grandifolia, Mag- 
nolia grandiflora, and Ilex opaca. However, West Feliciana Parish is a special 
case, because it almost entirely falls into the loess hills belt of the coastal plain 
province (Fenneman 1938). And, not surprising, in the northeast corner of West 
Feliciana Parish, not far from the border of Amite County, Mississippi, Delcourt 
and Delcourt (1974) record a forest composition change to Quercus alba, Pinus 
sp. or spp.,and Fagus grandifolia. Whether or not this presettlement forest was 
or was like the climax forest of the area is beyond the scope of this paper but 
has been much discussed (Blaisdell et al. 1974; Delcourt & Delcourt 1974, 1977, 
Hodgkins 1958; Kurz 1944: Monk 1965, 1968; Nesom & Treiber 1977; Pessin 1933; 
Quarterman & Keever 1962; Wells 1942). 

The best available information on the presettlement vegetation of Amite 


652 BRIT.ORG/SIDA 19(3) 


County is probably from notes of botanically-trained individuals who traveled 
through the region before heavy settlement. Darby and Dwight (1836), while 
describing neighboring Wilkinson County as “one of the most productive cot- 
ton districts in the U.S.” describe Amite County with “some good land... along 
and near the streams, [although] the great body of the county is... covered with 
fine timber [presumably Pinus palustris]” Darby (1817) describes the vegeta- 
tion of the thick loess, which would include Wilkinson County and perhaps 
some of Amite County, as “thickly timbered” and lists 33 species—all hard- 
woods—as the primary timber, including seven species of oak and four of 
hickory. Darby (1817) also lists some prominent vines and understory shrubs, 
most notably Vitis spp. and “brakes of the arundo gigantea (great cane)” 
[Arundinaria gigantea]. As one moves east, Darby (1817) describes the land in 
Amite County becoming of three qualities: “alluvion near the streams, that Spe 
cies of slopes called Hammock, and the open pine hills.” The eastern half of 
Amite County and eastward then become forests of longleaf pine with an ad- 
mixture of Quercus falcata, Q. alba, and Liquidambar styraciflua (Darby 1817). 
Darby’s three categories correspond directly with the present vegetation types 
given later: bottomland hardwood forest, ravines / upland hardwood and mixed 
forest, and longleaf pine / loblolly-shortleaf pine forests for alluvion, hammock, 
and open pine hills, respectively. 

Visits to the area by William Bartram in 1787 also confirm this general pattern 
(Harper 1958). Although Bartram did not travel through Amite County, he gives 
a personal account of the loess hills vegetation, noting that pine was “viewed 
here as a curiosity” and that eleven species of hardwoods were the “magnifi- 
cent” trees of the forest here, while farther east pines were the dominant trees. 

For the years 1807-1809, Cuming describes western Wilkinson County as 
“hilly, ... [with] the soil rich, though thinly inhabited”. other areas in Wilkinson 
County he describes as “comparatively well cultivated” (Thwaites 1904). He also 
writes of a trip through forest just south of the Mississippi border in Louisiana 
“abounding with that beautiful and majestick [sic] evergreen, the magnolia or 
American laurel” (Thwaites 1904), affirming the interpretation of land survey 
records by Delcourt and Delcourt (1974). 

Studies this century (Holmes & Foster 1908, Lowe 1913) continued to record 
a vegetation pattern similar to that of Darby (1817): hardwood in extreme west- 
ern Wilkinson County, followed by a mixed pine-hardwood forest, followed by 
longleaf pine forest from eastern Amite County eastward. Holmes and Foster 
(1908) report that Wilkinson County was 63% cleared, Amite County was 25% 
cleared, and Pike County (still including Walthall County) was 30% cleared. 
These figures do not imply that the uncleared land was primary forest, as the 
authors also speak of large tracts of secondary woods (Holmes & Foster 1908). 
Secondary woods, Holmes and Foster (1908) assert, became dominated by Pinus 
taeda rather than Pinus palustris. 


—— 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 653 


Recent Botanical Effort—Amite County has been visited a number of times this 
century by botanists. The earliest collections from the county were by Andrew 
Allison (1903-1907) and Thomas Bailey (1915-1916), which were subsequently 
noted in E.N. Lowe’s “Plants of Mississippi: A List of Flowering Plants and Ferns” 
(1921). Despite its antiquated nomenclature and several inaccuracies, Lowe's 
compilation was the only statewide checklist of plants for Mississippi until the 
publication of Kartesz (1999). A. J. Eames visited the county in 1942, but only 
one specimen has been seen (Phoradendron leucarpum, Eames s.n. [BH!)). 

The 1960s and 1970s showed the greatest amount of floristic work in Amite 
County. Louis Temple (MISS and later Mississippi College), Clair Brown (LSU), 
and especially Samuel B. Jones, Jr. (University of Southern Mississippi and later 
GA) and Jones's accompanying wife, students, and friends made numerous col- 
lections, turning up rarities for the area such as Chromolaena ivifolia and Hy- 
drangea arborescens. Jones and associates collected over 250 specimens from 
the county. Collections since that time have mostly been limited to small student 
collections. One exception is the late John Allen Smith, a local school teacher 
who was contracted by the Mississippi Natural Heritage Program to find rare 
plants in the Homochitto National Forest. He located several populations of 
Antennaria solitaria, Mikania cordifolia, Pachysandra procumbens, and Ste wartia 
malacodendron in the county. 

Present Vegetation Iypes and Land Use.—The vegetation of Amite County 
isnow mostly (if not exclusively) secondary. This is not surprising considering 
the early settlement coupled with an economy based primarily upon timber, 
beef cattle, and dairy cattle. Despite this somber note, one may be surprised 
that 78% of the county is forested and 22% is open land (Mississippi Forestry 
Commission 1998), an irony primarily due to the resurgence of the timber in- 
dustry through tree farming. 

Today, dairy production is restricted mostly to the southeastern corner of 
the county, an area once dominated by longleaf pine. Beef farms remain scat- 
tered throughout the county. Dairy and beef cattle production are the two major 
reasons for open land in the county, with smaller areas attributed to crops such 
as corn, soybeans, and ryegrass. Timber harvesting, too, continues throughout 
the county but in this age of tree farming leaves open areas only for short periods 
between harvest and replanting operations. In the year 1998, the merchantable 
timber (growing stock) in Amite County was estimated to be 309.0 million cubic 
feet of pine and 206.1 million cubic feet of hardwood (Mississippi Forestry Com- 
mission 1998). Only three counties in Mississippi have a greater volume of mer- 
chantable pine timber. During that same year, 68,314 MBF (thousand board feet, 
Doyle Rule) of pine were harvested, 5033 MBF of hardwood were harvested, 
103,946 cords of pine pulpwood were harvested, and 68,689 cords of hardwood 
pulpwood were harvested (Mississippi Forestry Commission 1998). 

Forestry management practices differ throughout the county. The north- 


654 BRIT.ORG/SIDA 19(3) 


western corner of the county, for instance, is part of the Homochitto National 
Forest, and as such enjoys relatively longer timber rotation times, mandatory 
streamside protection zones, several older-growth areas, and even longleaf pine 
management and replacement. The rest of the county is owned privately, and 
timber production and management is determined more by economics and by 
soil properties. In the western part of the county, where the loess deposits are 
relatively thick, harvested areas require much attention (e.g., burning, discing, 
chemical treatments) because of rapid post-harvest non-timber growth CE. 
Alford, Mississippi Forestry Commission, pers. comm.). Open areas are quickly 
exploited by trees suchas Liquidambar styracifluaand Acer rubrum var. rubrum 
and other plants such as Rubus spp., Lactuca spp.,and Eupatorium spp. The east- 
ern part of the county, where the soil is less loessal, is not quickly exploited by 
vigorous undergrowth, and thus timber practices like seed-tree regeneration 
are more common CE. Alford, Mississippi Forestry Commission, pers. comm.). 

Pine forests comprise 45% of forested lands in the county. Mixed forest (19%) 
and hardwood forest (36%) also make up a sizable portion of forested land in 
the county (Mississippi Forestry Commission 1998). Of the pine forests, the pri- 
mary type is loblolly or loblolly-shortleaf, and most hardwood forests belong 
to bottomland forests, with fewer acres in uplands and ravines, swamps, and 
other areas. 

For purposes of explaining general vegetation patterns and to give the reader 
a background template upon which to interpret the vascular plant checklist, 
the vegetation of Amite County is divided into twelve categories. Naturally, this 
is a simplification and overlooks variation in a great continuum. A short de- 
scription and common species of each type will be given (in alphabetical order, 
so as not to imply a rigorous statistical measurement of abundance). Species 
primarily restricted to certain types are noted, as are rare species. In general, these 
vegetation types correspond to those of Allen (1972), modified by the author 
and treatments by Braun (1950), the United States Forest Service (1995), Chris- 
tensen (2000), Clewell (1985), and Delcourt and Delcourt (2000). 


1. Longleaf Pine Forest (Fig. 3).—Longleaf pine forest is uncommon in the 
county, occurring only in scattered localities in the southeast and in the north- 
western corner on high ridges of the Homochitto River basin. Typically, these 
forests are upland, open, and regularly burned. The dominant tree species, of 
course, is Pinus palustris, which is usually accompanied to some extent by P. 
echinata or P. taeda. Depending on the burning regime, other associates may 
include Callicarpa americana, Carya tomentosa, Cornus florida, Crataegus spp., 
Diospyros virginiana, Ilex vomitoria, Malus angustifolia, Morella cerifera, Quercus/ 
falcata, Q. marilandica, Q. stellata, Rhus copallina, Vaccinium arboreum, and V. 
stamineum. The understory is dominated by composites, grasses, and legumes, 
primarily Andropogon virginicus, Aristida spp., Desmodium spp., Lespedeza 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 655 


Fic. 3. photograpis of majer Negerablon types in Amite oe Lo Upper: longleaf pine forest. Middle: 


Inhlally.ct Fagus grandifolia Arundinaria 
g g 


ODIony 


gigantea, and the predominance of hardwoods. 


656 BRIT.ORG/SIDA 19(3) 


repens, Schizachyrium scoparium, Symphyotrichum dumosum, S. patens, and 
Tephrosia spicata. Some species in the county are found primarily in these for- 
ests, suchas Aletrisaurea, Asclepias viridiflora, A. viridis, Drosera brevifolia, lonactis 
linariifolius, Liatris elegans, L. pycnostachya, Phlox pilosa, Polygala incarnata, 
Pycnanthemum albescens, Quercus incana, Rhynchosia_ reniformis, 
Symphyotrichum adnatum, and Tragia smallii. An example of this vegetation 
type is on private property about six miles south-southeast of Liberty, T2N R4E 
Sec 28 NW/4. An example from the Homochitto National Forest is at the inter- 
section of West Homochitto Road and Royal Chapel Road, T4N R2E Sec 19 E/2. 
All longleaf pine forest ison upland sites, and thus, there are no flatwoods (sensu 
Clewell 1985) in the county. 

2. Loblolly-Shortleaf Pine Forest (Fig. 3).—This is the most abundant veg- 
etation type in the county. Under this category are included old field pine succes- 
sion, mature upland non-longleaf pine forest, and pine plantation. These forests 
are dominated by Pinus taeda. If burning is frequent or the soil is gravelly or 
sandy, Pinus echinata becomesa clear co-dominant species. Understory or sup- 
pressed species include Acer rubrum var. rubrum, Diospyros virginiana, Ilex 
vomitoria, Liquidambar styraciflua, Quercus falcata, Prunus serotina, Rhus 
copallina, and Vaccinium arboreum. The ground layer is typically species-poor 
in unburned or plantation pine, inhabited by Callicarpa americana, Clitoria 
mariana, Desmodium spp, Dichondra carolinensis, Lonicera japonica, Lygodium 
japonicum, Mimosa microphylla, Oxalis spp., Panicum spp., Rubus spp., Toxico- 
dendron radicans, and Vitis rotundifolia. Some pine plantations may consist 
only Pinus taeda and Lygodium japonicum. With burning arrives Hypox 
hirsuta, Lespedeza spp.,and Tephrosia virginiana. Mature, open loblolly- anes 
pine forest is home to Andropogon ternarius, A. virginicus, Aristida spp., Ascl- 
epias tuberosa, Cirsium carolinianum, Coreopsis spp., Eragrostis spp. Gentiana 
villosa, Helianthemum carolinianum, Hypericum drummondii, Ipomoea 
pandurata, ee Oxalisditlenti , Polygala mariana, P nana, Rhus 
glabra, Rosa carolina, Ruelli is, abundant Schizachyrium scoparium, 
Sabatia spp., Scleria: Spp., Scutellaria integrifol ia, Seymeria cassioides, Solidago 
odora, Stylosanthes biflora, Vernonia texana, and Viola pedata. Wet ditches andl 
old roads through these forests further add species such as Carex glaucescens, 
Cyperus spp., Hypericum mutilum, Ludwigia spp., Rhynchospora inexpansa,and 
Xyris laxifolia var. iridifolia. Rarer species in these forests include Galactia 
erecta, Gaylussacia dumosa, Ludwigia hirtella, Oenothera linifolia, Orbexilum 
pedunculatum, Rhynchosia tomentosa, Scutellaria incana, and Trichostema 
setaceum. An example of this type of vegetation is at the northern corner of the 
intersection of Mary Wall Bridge Road and River Road, TIN R5E Sec 19 W/2. 

3. Upland Hardwood and Mixed Forest.—This vegetation type is also known 
as oak-pine or oak-pine-hickory forest. This type is quite similar to loblolly- 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 657 


shortleaf pine vegetation, only that the dominant and co-dominant hardwoods 
more conspicuously join the pines, especially Carya pallida, C. tomentosa, 
Cornusflorida, Fraxinusamericana, Hamamelis virginiana, Ilex decidua, L longipes, 
Ostrya virginiana, Prunus serotina, Quercus alba, Q. falcata, Q. hemisphaerica, 
Q. nigra, Q. velutina, Ulmus alata, Vaccinium elliottii,and Viburnum scabrellum. 
The herbaceous layer tends to be less developed, and species like Chasmanthium 
sessiliflorum, Matelea gonocarpos, Mitchella repens, Osea acrostichoides, 
and Smilax pumila join the list. Uncommon s} found in this vegetation type 
are Chamaelirium luteum, Lilium michauxii, Malaxis unifolia, Polygonatum 
biflorum, Smilax pulverulenta, and Tipularia discolor. An example of this type 
of vegetation is found near the pipeline crossing on Dickey Mills Road, T3N 
RGESe6sh5/2, 

4, Ravines.—These areas are also known as hammocks and bluffs. The typical 
ravine is rich in both woody and herbaceous species. Common trees and large 
shrubs include Carpinuscaroliniana, Carya spp. Cornusflorida, Fagusgrandifolia, 
Frangula caroliniana, Hamamelis virginiana, Hydrangea quercifolia, Ilex opaca, 
Liriodendron tulipifera, Magnolia acuminata, Ostrya virginiana, Oxydendrum 
arboreum, Quercus alba, Q. michauxii, Q. pagoda, Q. velutina, and Viburnum 
rufidulum. Common herbaceous species include Carex spp., Chasmanthium 
sessiliflorum, Melica mutica, Oxalis violacea, Podophyllum peltatum, Phlox 
divaricata, Sanicula smallii, Spigelia marilandica, and Viola walteri. Less com- 
mon species that are found primarily in this type of vegetation are Antennaria 
solitaria, Aristolochia serpentaria, Carya cordiformis, Dioscorea quaternata, Ilex 
ambigua, Matelea carolinensis, Phegopteris hexagonoptera, Schisandra glabra, 
Solidago auriculata, Spiranthes ovalis, Stewartia malacodendron, and Uvularia 
perfoliata. Ravines are often the areas preserved by private landowners, and 
whether or not these rarer species are restricted to these areas because of their 
vegetation-edaphic properties or as a result of less disturbance is unknown. 

Ravines fall into two major subtypes: A. Homochitto Ravines (Fig. 3).—These 
ravines are located in the Homochitto National Forest and surrounding areas. 
They generally have a thick loess aye! over Miocene a and the hills are 
highly dissected. These ravines have th of Acer barbatum, 
Magnolia macrophylla, Quercus shumardii, and Symplocos tinctoria (as a mid- 
to upper-slope tall shrub) and much more abundant Magnolia acuminata and 
Pinus taeda. In the understory, ferns reach their peak with large areas domi- 
nated by Phegopteris hexagonoptera, Polystichum acrostichoides, and Thelypteris 
kunthii. Also added are Antennaria solitaria, Botrychium virginianum, abundant 
Carex flaccosperma, Cynoglossum virginianum, Laportea canadensis, Mikania 
cordifolia, Piptochaetium avenaceum, Smilax pumila, Urtica chamaedryoides, 
and Woodsia obtusa. Because of the dissection of land in parts of the Homochitto 
National Forest, there can of ten be a continuum between this type, upland hard- 


658 BRIT.ORG/SIDA 19(3) 


wood and mixed forest, and pine forest, depending on the size of the ridges. An 
example of this type of vegetation can be found in the Homochitto National 
Forest off New Hope Road in T3N R2E Sec 12. 

B. Gravel or Saffell outcrops.—Gravel outcrops occur primarily as ridges 
along the margins of the Amite River and associated large drainages. The soil 
here is very gravelly and sandy, and species like Carya glabra, Frangula 
caroliniana, Styrax grandifolius, and Tilia americana var. caroliniana become 
more common. One adds many species seldom found elsewhere in the county, 
including Agrimonia microcarpa, A. rostellata, Cocculus carolinus, Desmodium 
glutinosum, D. nudiflorum, D. rotundifolium, Fleischmannia incarnata, Litho- 
spermum tuberosum, Phryma leptostachya, Physalis heterophylla, Potentilla sim- 
plex, Quercus coccinea, Silene stellata, Smallanthus uvedalius, Symphyotrichum 
drummondii var. texanum, and Tragia cordata. Pleopeltis polypodioides var. 
michauxiana can be found here growing both epiphytically and on outcrops 
of conglomerate rock and sandstone. Pachysandra procumbens is also found here 
in one locality. An example of this vegetation type can be found at the Ethel S. 
Vance Natural Area just west of Liberty on MS Hwy 24, T2N R4E Sec 6 SE/4. 

5. Bottomland Hardwood Forest (Fig. 4).—Bottomland forest, too, is very 
rich in both woody and herbaceous species. There is much variability in this 
type, and one ood: is provided for the most conspicuous outlier. There is 
also some intergradation with spring-seeps and swamps, ponds and lakes, and 
streamsides, each of which is considered a separate category. Typically, bottom- 
land hardwood forest is dominated by Asimina triloba, Carpinus caroliniana, 
Fagus grandifolia, Halesia diptera, Ilex opaca, Pigainann oa: styraciflua, Lirio- 
dendrontulipifera, Magnolia grandiflora, Q uercus laut Tene 9. niga O.michauxii, 
Q. pagoda, and Symplocostinctoria. Li | ,Decumaria 
barbara, and Smilax spp. Herbaceous erie can commonly be non-exis- 
tent in deeper forest, but at edges, small openings, small drains, and woods roads 
appear Carex intumescens, C. joorii, C. lurida, Chasmanthium laxum, Cuphea 
carthagenensis, Elephantopus carolinianus, Gratiola spp., Hexastylis arifolia, 
Hydrocotyle spp., Hymenocalliscaroliniana, Impatiens capensis, Leersia virginica, 
Lycopus virginicus, Macrothelypteris torresiana, Packeraglabella, Polygonum spp. 
Ranunculus abortivus, R. pusillus, Rhynchospora caduca, R.glomerata, Samolus 
valerandi ssp. parviflorus, Saururuscernuus, Selaginella apoda, Thelypteris kunthii, 
Verbesina walteri,and Viola primulifolia. Uncommon species found primarily 
in bottomlands are Circaea lutetiana ssp. canadensis, Claytonia virginica, 
Listera australis, Solidago discoidea,and Trillium foetidissimum. In the Homochitto 
drainage, Acer rubrum var. drummondii, Geum canadense, Lindera benzoin, 
and Ulmus americana join the list. Bottomland forest is commonly invaded by 
exotics, especially Ligustrum sinense and Lygodium japonicum. An example of 
the generalized type can be found at the Ethel S. Vance Natural Area just west 
of Liberty on MS 24 at the West Fork Amite River, T2N R4E Sec 6 S/2. 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 659 


Fic. 4. Rhotodiatte of Har vegetation types in Ants eo pees Dis ll Upper: bottomnland hardwood forest. 


Middle: swamp, here dominated by Nyssa biflora and Cep d arm pond, wit 
Hydrocotyle and Saururus evident. 


660 BRIT.ORG/SIDA 19(3) 


There is one major distinctive subtype of bottomland forest. Magnolia- 
Beech-Spruce Pine-Ilicium Thickets.—The locals call this type of vegetation 
“stink-bush thickets.” These are forests of Fagus grandifolia, Magnolia grandi- 
flora, and Pinus glabra with a dense understory of Illicium floridanum. Often 
there are no herbaceous plants, and the Illicium floridanum can become im- 
penetrable. Occasional species in these areas are Athyrium filix-femina, Epi- 
dendrum conopseum, Epifagus virginiana, Polystichum acrostichoides, Stewartia 
malacodendron, Symplocos tinctoria, and the rare Trichomanes petersii. An ex- 
ample of this type can be found along the East Fork Amite River just south of 
South Greensburg Road, TIN R4E Sec 27. 

6. Spring-Seeps and Swamps (Fig. 4).—Spring-seeps and swamps fall into 
the broader category of forested wetlands. Woody species typically include 
Cephalanthus occidentalis, Fraxinus pennsylvanica, Ilex opaca, I. verticillata, 
Itea virginica, Magnolia virginiana, Nyssa biflora, Photinia pyrifolia, Quercus 
spp., Sabal minor, Smilax laurifolia, and Viburnum nudum. The herbaceous 
layer consists of Carex albolutescens, C. intumescens, C. leptalea, C. lonchocarpa, 
C.lurida, Gratiola floridana, Hydrocotyle spp., Osmunda cinnamomea, O. regalis 
var. spectabilis, Panicum ¢g gymnocarpon, P rigidulum, Pilea pumila, Polygonum 
spp., Proserpinaca palustris, Spare icanum, Triadenum walteri, Viola 
primulifolia, and Woodwardia dreolata. Uncommon species found primarily 
in these areas include Arnoglossum plantagineum, Dryopteris ludoviciana, 
Leucothoe racemosa, Melanthium virginicum, Platanthera flava, Smilax walteri, 
Spiranthescernua, Stachys tenuifolia,and Woodwardia virginica. In occasional 
ox-bows, Taxodium distichum will form monotypic stands. Asclepias perennis, 
Carex decomposita, and Leersia lenticularis are uncommon species that occur 
in these areas. A typical swamp can be found at the Ethel S. Vance Natural Area 
just west of Liberty on MS 24 downhill from the above-mentioned Saffell out- 
crop, T2N R4E Sec 6 S/2. Several typical spring-seeps can be found on school 
lands near Royal Chapel, T4N R2E Sec 18 NE/4. A mature baldcypress swamp 
can be found in the Homochitto National Forest, Brushy Creek at Robertson 
Road, T4N R2E Sec 2 SW/4. 

7. Cultivated and Fallow Fields.—Land that is under cultivation or has 
been under cultivation recently (including home gardens) falls into this cat- 
egory. The common species are typical weedy herbs, including Amaranthus 
spinosus, Brassica rapa, Cyperus rotundus, Eclipta prostrata, Geranium 
carolinianum, Jacquemontia tamnifolia, Lolium perenne, Physalis angulata, 
Raphanus raphanistrum, Senna obtusifolia, Sesbhania herbacea, Solanum 
carolinense, Soliva pterosperma, Trifolium repens, Urochloa platyphylla, Vicia 
sdtiva ssp. nigra, and V. tetrasperma. Rarely one finds Melothria pendula, 
Muscari neglectum, or Narcissus jonquilla. An example of this vegetation type 
can be found in the areas surrounding Pumpkin Patch Creek at MS 569N, T4N 
RGE Sec 7 35/2. 


— 


— 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 661 


8. Pastures and Old Successional Fields.—T hese areas are the conspicuous 
autumn flower shows. Common species include Agalinis fasciculata, Ambro- 
sid artemisiifolia, Andropogon spp., Buchnera americana, Chrysopsis mariana, 
Croton capitatus, Eremochloa ophiuroides, Eupatorium spp., Helenium amarum, 
H. flexuosum, Hordeum pusillum, Lactuca spp., Lespedeza spp., Paspalum spp. 
Pityopsis graminifolia, Polypremum procumbens, Ranunculus spp., Rumex spp. 
Sisyrinchium spp., Solidago canadensis, and S. gigantea. In moist pastures, 
Cyperus spp. and Juncus spp. become common. Fencerows and edges of this 
type are home to Albizia julibrissin, Diospyros virginiana, Prunus serotina, and 
Triadica sebifera. An example of this vegetation type can be found along Cut 
Through Road, T3N R5E Sec 28 N/2. 

9. Roadsides.—Roadsides support a diverse array of species, although the 
habitat and species are generally called “ruderal.” Common species include 
Antennaria plantaginifolia, Boltonia diffusa, Campsis radicans, Chamaecrista 
fasciculata, Cicuta maculata (in ditches or wetter sites), Erigeron strigosus, 
Helianthus angustifolius, Ipomoea cordatotriloba, Linum medium, Oenothera 
spp., Panicum spp., Paspalum dilatatum, P urvillei, Plantago aristata, Pueraria 
montana, Pyrrhopappus carolinianus, Robinia pseudo-acacia, Rudbeckia hirta, 
Sorghum halepense, Tradescantia ohiensis, Tridens flavus, Trifolium spp., Ver- 
bena brasiliensis, Vernicia fordii, and Vicia villosa. Uncommon species in this 
vegetation type are Ailanthus altissima (apparently only beginning its spread 
in southern Mississippi), Lysimachia lanceolata, and Penstemon laxiflorus, as 
well as several waifs collected previously but not seen during this study (e.g, 
Crotalaria spectabilis, Geranium dissectum). 

10. Ponds, Lakes, and Beaver Impoundments (Fig. 4).—This vegetation 
type encompasses all areas that are in or on the margin of stationary water, 
primarily ponds, lakes, and beaver impoundments. There is some overlap with 
the vegetation type “Spring-Seeps and Swamps,” and one should reference that 
section as well. Common woody plants include Cephalanthus occidentalis, 
Nyssa biflora, Smilax walteri, and Taxodium distichum. The herbaceous layer 
includes Bidens spp., Brasenia schreberi, Eleocharis obtusa, Eryngium 
prostratum, Juncus repens, abundant Ludwigia spp., Nuphar lutea, Panicum 
hians, Rhexia mariana, Saccharum spp., Sagittaria latifolia, Sparganium 
americanum, Thelypteris palustris var. pubescens, Typha latifolia, and Utricu- 
laria biflora. Uncommon species in this type include Carex louisianica, 
Gelsemium rankinii, Hydrolea uniflora, Potamogeton diversifolius, and P. 
pulcher. An example of this vegetation type (more precisely, an open beaver 
impounded pond) can be found at the Ethel S. Vance Natural Area just west of 
Liberty on MS 24, T2N R4E Sec 7 N/2. 

11. Riversides.—Riversides represent a special case of bottomland hard- 
wood forest. The soil of ten has much more sand and gravel, and the flora is strik- 
ingly distinct. Moving upstream, as the rivers become smaller streams, the veg- 


662 BRIT.ORG/SIDA 19(3) 


etation type, however, begins to grade with bottomland hardwood forest. Riv- 
ersides are typically dominated by the woody plants Acer negundo, Betula ni- 
gra, Platanus occidentalis, and Salix nigra. The herbaceous layer includes Apios 
americana, Carex lupulina, Chasmanthium latifolium, Cleome hassleriana, 
Commelina diffusa, C. virginica, Digitaria ciliaris, Hygrophila lacustris, Justicia 
ovata var. lanceolata, Lobelia cardinalis, Mikania scandens, and Vernonia 
gigantea. Uncommon plants found in this vegetation type are Leersia oryzoides 
and Melochia corchorifolia. Anexample of this vegetation type can be found on 
private property along the West Fork Amite River south of Powell Road, TIN 
RAE See 51. 

12. Cemeteries, Yards, and Ruderal Areas in Towns.—This last category 
incorporates all “ruderal” areas except roadsides, which do share a number of 
elements. Many species of trees are planted and grow natively in yards, so con- 
centration here will focus on herbaceous species. The most common species 
are Cerastium glomeratum, Duchesnea indica, Galium aparine, Glechoma 
hederacea, Houstonia spp., Krigia spp., Lamium amplexicaule, Lepidium 
virginicum, Modiola caroliniana, Nothoscordum bivalve, Nuttallanthus 
canadensis, Oxalis rubra, Paspalum notatum, Poa annua, Salvia lyrata, 
Sporobolus indicus, Stellaria media, Trifolium spp, Triodanis spp., Valerianella 
radiata, Veronica spp.,and Youngia japonica. Uncommon species in these habi- 
tats include Cyrtomium falcatum pees matinalized), ea helix (ex- 
panding only around old home sites), Le} t 
Ophioglossum spp. (cemeteries), Phyllanthus urinaria (sidewalk cracks in town), 
Plantago heterophylla (cemeteries), and Portulaca oleracea (sidewalk cracks in 
town). Anexample of botha yard and ruderal area in town is Old Jackson Road 
at Pecan Street in Liberty, T2N R4E Sec 4. An example of a cemetery is the 
Stewart- Wall Cemetery near the intersection of Mary Wall Bridge Road and 
Mt. Vernon Road, TIN R4E Sec 24. 


Ke s) 


ANNOTATED CHECKLIST 


The following checklist is the compilation of all native, naturalized, or espe- 
cially long-persisting vascular plant species occurring in Amite County, Mis- 
sissippi. Families are arranged in alphabetical order under the divisions of 
Lycopodiophyta, Polypodiophyta, Coniferophyta, Magnoliophyta: Liliopsida 
(monocots), and Magnoliophyta: Magnoliopsida (dicots). Family delimitation 
in the Lycopodiophyta, Polypodiophyta, and Coniferophyta follows the system 
of Kartesz (1999). Delimitation of monocot families follows the system of 
Dahlgren et al. (1985), which primarily means that the Liliaceae sl. are split 
into various smaller families. Delimitation of dicot families generally follows 
the system of Kartesz (1999) with modifications to avoid polyphyletic or un- 
natural taxa, especially regarding the older concepts of Saxifragaceae and 
Loganiaceae. More specifically, Viburnum and Sambucus are considered part 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 663 


of Adoxaceae (Donoghue 1983; Donoghue et al. 1992), Lepurapetalon part of a 
broadly defined Celastraceae (Soltis & Soltis 1997), Itea part of Iteaceae (Soltis 
& Soltis 1997), Penthorum part of Penthoraceae (Haskins & Hayden 1987; Soltis 
& Soltis 1997), Gelsemium part of Gelsemiaceae, and Polypremum part of 
Buddlejaceae, despite much uncertainty regarding its position (Struwe et al. 
1994; Backlund et al. 2000). Labiatae and Verbenaceae are retained and used as 
set forth in Kartesz (1999), although their circumscriptions are likely to change 
in the future. Genera and species are in alphabetical order in each family. Spe- 
cies entry data are arranged in the following manner: 


[@ = new state record] [t = species taxonomy or nomenclature differing from 
Kartesz, 1999]|* = non-native] Species Authority [following Brummitt & Powell 
1992], Collection Number. Frequency. Vegetational associations. Note(s). 


Species nomenclature (scientific names plus authorities), in general, follows 
Kartesz (1999). If there is a departure from his system, a cross and a reference 
citation or note accompanies the entry. The only major departure from his no- 
menclature is my retention of Panicum in the large sense. Because the taxonomy 
of Panicums..is controversial and obviously still rudimentary, | follow the last 
treatment of the genus for Mississippi (Lelong 1986). Only four hybrids are part 
of the list: Eupatorium x pinnatifidum, Gladiolus x gandavensis, Rhexia nashii 
x R. virginica, and Quercus x comptoniae. Each of these iscommon enough that 
explorers of the region should be aware of their presence. If the species is not 
native, then the citation is accompanied by an asterisk. References for this cri- 
terion are Radford et al. (1968), Clewell (1985), Correll & Johnston (1970), 
Weakley (in prep.), and the Mississippi Natural Heritage Program (pers.comm.). 

Specimens have been deposited at DUKE, and duplicates of most species 
have been deposited at IBE. Additional duplicates have been variously distrib- 
uted to BEO, BH, LSU, MICH, MISS, NCU, US, the Mississippi Museum of Natu- 
ral Science (mmns), and the personal herbarium of Charles T. Bryson (ctb) (her- 
barium abbreviations following Holmgren et al. 1990). Collection numbers with 
an A refer to Mac H. Alford (also used when other collectors assisted), with an 
FA refer to E. Earl Alford, Amite County Forester, Mississippi Forestry Com- 
mission, and with a WAGM refer to Robert L. Wilbur (DUKE), Mac H. Alford, 
and Gerry Moore (BKL). 

Frequency, a measure of how common and widespread a species is, will 
follow the pattern outlined in Table 2. For maximal information retrieval, the 
plant association data is clearly, though sometimes verbosely, described. If the 
descrit given are terse, the vegetation types of the previous section are fully 
adequate to describe the habitat(s) of occurrence. 

Any species occurring in Amite County that was not directly planted or 
vegetatively produced in close proximity toa cultivated individual is considered 
a part of the flora. For example, Camellia japonica, though widely reproducing 


664 BRIT.ORG/SIDA 19(3) 


Taste 2. Abundance scale for vascular plants in the checklist of Amite County, Mississippi 


Abundant Frequent, dominant, or Epagaininany) in one or more habitats; widespread. 

Common Relatively pt frequen 
in a subset of habitats. 

Locally Common Fither widely distributed but completely restricted to a certain habitat 
or locally distributed with many individuals. 

Infrequent Uncommon in the county, often small populations and/or restricted to 
certain habitat 

Rare ne or two al populations known from the county. 


vegetatively under the growth of planted individuals, is not found outside a 
close perimeter of planted individuals and was never found in an area where its 
origin was ambiguous; it is not included. On the other hand, Cyrtomium falcatum, 
though remaining in close vicinity to planted (or persisting) individuals, often 
produces spore-derived individuals around old, moist concrete foundations and 
brick walls; it is included. Other species such as Photinia serratifolia and Eu- 
onymus fortunei show few Gf any) signs of reproduction, but persist so long, 
even after abandonment of property, that they are included, asa botanist is likely 
to encounter them with little inherent evidence to suggest a cultivated origin. 

Methods.—Field collections were made from 1997-2000. Because the county 
is divided according to the rectangular system of surveying, one section of each 
township is designated for public schools. These public lands served as evenly 
distributed points for collection. Because they are maintained by the Missis- 
sippi Forestry Commission for various management purposes, a diversity of 
plant communities and edaphic conditions were available. Public land was also 
accessible in the Homochitto National Forest, a significant portion of the land 
area in the northwest corner of the county (Fig. 1). Only one sizeable park oc- 
curs in the county, the Ethel S. Vance Natural Area near Liberty. This park, too, 
was extensively surveyed. Private lands and roadsides were also searched, but 
usually less extensively and in a more random manner, although a combina- 
tion of topographic maps and the soil map were used to pinpoint potential in- 
teresting areas. 

Field identification was undertaken primarily with Clewell (1985), Radford 
et al. (1968), and Correll and Johnston (1970). Upon return to DUKE, monographs 
and other references, especially Allen (1992), Bailey 1949), Chapman (1897), 
FNA (1997), Godfrey (1988), Godfrey and Wooten (1979, 1981), Small (1933), 
Steyermark (1963), and Weakley (in prep.), were consulted, and specimens were 
compared to those in the herbarium for accuracy. 

Previous collections were consulted at GA, IBE/MISSA, LSU, MICH, and 
MISS. Several monographic and floristic studies had Amite County marked as 
occurrences on dot-maps or in specimen citation lists. If these specimens were 
not seen, the reference is given. 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


LYCOPODIOPHYTA 


SELAGINELLACEAE 
Selaginella apoda (L.) Spring, A1945. Infrequent. 
Bottomland forest. 


POLYPODIOPHYTA 


ASPLENIACEAE 

Asplenium platyneuron (L) B.S.P,A1049.Common. 
Saffell oe upland hardwood forest, 
and pine 

BLECHNACEAE 

Woodwardia areolata (L.) T.Moore,A1296. Locally 

n. Spring-seeps, swamps, and 

WO ie areas near impoundments. 

Woodwardia virginica (L.) Sm., A1719. Rare. 
Spring-seep in the Homochitto National 
Forest. 


DENNSTAEDTIACEAE 

Pteridium aquilinum var. pseudocaudatum (Clute) 
A.Heller, A1695. Common. Roadsides and 
pine forest. 


DRYOPTERIDACEAE 
Athyrium filix-femina ssp. asplenoides (Michx.) 
Hultén, A7050. Abundant. Mesic forest and 


fores 
ae um faleatur (L.A) C. Presl, A1784. Rare. 
and shady brick and concrete walls. 
es protrusa (Weath.) Blasdell,A 7086. Rare. 
mland hardwood forest along 
chitto River. 

Dryopteris ludoviciana (Kunze) Small, A2029.Rare. 
S along the East Fork Amite River. 
Onoclea sensibilis L., A1341. Locally common. 
Roadside ditches, mesic forest, and swamp 

gins 


Polystichum acrostichoides (Michx.) Schott, 
W, 921. Abundant. Mesic forest, rich 
ravines, Saffell outcrops, and upland hard- 


wood forest. 

Woodsia obtusa (Spreng.) Torr.ssp.obtusa,A2114 
Rare. Sandy bottomland forest and associ- 
ated loessal hills of the Homochitto River 
basin. 

HYMENOPHYLLACEAE 

Trichomanes petersii A.Gray,A2028. Rare. Bases of 

us grandifolia trunks near springs of the 
East Fork Amite River. 


LYGODIACEAE 

*lygodium japonicum (Thunb. ex Murray) Sw., 
WA&M 71082. Abundant. Roadsides, pine for- 
est, mesic hardwood forest, and swamp 
margins 


OPHIOGLOSSACEAE 
Botrychium biternatum (Savigny) Underw.,A352. 
m Forest 


Botrychium virginianum (L.) Sw., A1007. Infre- 
and mixed forest, primarily in the 
Homochitto River basin. 
Ophioglossum crotalophoroides Walter. Rare. 
Sandy lawns and cemete 
Evans (1978): specimen not s 
Ophi ogi nudicaule | f. Rare ic awns, pasture 
farenced in Fyan (1978); 


ries. Referenced in 


specimen not seen. 


OSMUNDACEAE 

Osmunda cinnamomea L., A524. Infrequent. 
Spring-seeps and moist ravines. 

Osmunda regalis var. spectabilis (Willd.) A. Gray, 
A531. Locally Common. Spring-seeps, 
swamps, and wooded areas near impound 
ments. 


POLYPODIACEAE 
Pleopeltis polypodioides ssp. michauxiana (Weath.) 
Andrews & Windham, A1526. Locally 
common. Epiphyte primarily on Quercus 
stellata, Quercus michauxii, Carya illinoinensis, 
and Juniperus virginiana and infrequently grow- 
ing terrestrially on exposed outcrops of 
Saffell sandstone 
PTERIDACEAE 
Adiantum pedatum L.,A2257.Rare. Saffell outcrop 
along the East Fork Amite River 
THELYPTERIDACEAE 
re castes torresiana (Gaudich.) Ching, 
Al085. Co n. Mesic forest and forest 


oads. 

nea hexagonoptera (Michx.) Fée, A1289. 
Infrequent. Hardwood ravines, primarily in 
the Homochitto River basin. 

Thelypteris hispidula var. versicolor (R.P. St John) 
Lellinger, A1008. Infrequent. Sandy bottom- 
land hardwood forest. 

Thelypteris kunthii (Desv.) C.\V.Morton, 41339. 
Common. Mesic forest and forest roads. 

Thelypteris ie var. pubescens (G. Lawson) 


Fernald, A/342. Infrequent. Swamp margins 
and perennially wet ditches. 
CONIFEROPHYTA 
CUPRESSACEAE 
Juniperus eed L. var. virginiana, A1357. |In- 
requent. Fencerows and old home sites. 
Taxodium dean (L.) LC. Rich., A1347. Infre- 
quent.Ox-bow sloughs, swamps, and flood: 


plain swales. 
PINACEAE 
Pinus echinata Mill.,A1167.Common. Pine forest 
and old fields. 
*Pinus elliottii Engelm. var. elliottii, EA 43. Infre- 
quent. Planted in plantations, especially in 
the 1950s and 1960s, and also in yards and 


towns. 
Pinus glabra Walter,A793.Infrequent.Bottomland 


forest. 

Pinus palustris Mill, A1 192. Infrequent. Pine for- 
est of the eastern half of the county and high 
ridges of the Homochitto River ba 

Pinus taeda L., FA 204. Abundant. aaa every- 
where except cultivated fields and in per- 
manent water. Commonly cultivated in 
plantations. 


MAGNOLIOPHYTA: LILIOPSIDA 
ogelaaadien 


(L.) Salisb.ex Rose, L.C. Temple 
9821 (MISSI) inf eau eat Pine forest and 


roadsides 

t Yucca fect Haw. [Y. filamentosa L. in part], 
WA&M 70997. Infrequent. Pine forest, particu- 
larly in the Homochitto River basin. Species 
taxonomy follows Trelease (1902). 


ALISMAT. 
Sagittaria ae Willd., A1459. Infrequent. 
Swamps, pond margins, and springs. 


IACEAE 

*Allium ampeloprasum L.,A778.Rare, Perennially 
wet roadside ditches in full sun 

Allium canadense L. var. canadense, A817. Com- 

NON, dsides and fields. 

Nothoscordum bivalve (L.) Britton, A264. Abundant. 
Fields. roadsides, yards. and te 

*Nothoscordum gracile (Aiton) ane ee 
Rare. Gravelly ruderal areas in Liberty. 


BRIT.ORG/SIDA 19(3) 


ALSTROEMERIACEAE 
@ *Alstroemeria psittacina Lehm.,A1919.Rare.Yard 
in Liberty 


AMARYLLIDACEAE 
at ects caroliniana (L.) Herbert, EFA 266. In- 
ent. Bottomland hardwood forest. 
ec tae L.,A374.Infrequent. Persisting 
ome sites and naturalizing into row 
crop fields. 


ARACEAE 

Arisaema dracontium (L.) Schott, A1004. Infre- 

uent. Mesic hardwood forest. 

Arisaema triphyllum (L.) Schott [including A. 
guinatum (Buckl.) Schott], A506. Common. 
Mesic forest, ravines, and occasionally pine 
forest. 

Orontium aquaticum L.,A533.|Infrequent. Springs 
and sloug 


ARECACEAE (see PALMAE) 


BROMELIACEAE 

Tillandsia usneoides (L.) L., A1397. Infrequent. 
Epiphyte on various trees and shrubs in 
mesic and lowland forest, and on Quercus 
spp.in upland forest or in yards. 


COMMELINACEAE 

*Commelina diffusa Burm.f., A1362. Infrequent. 
Sandy margins of rivers. 

Commelina virginica L.,A1361. Infrequent. Sandy 

margins of rivers. 

Tradescantia hirsutiflora Bush, A741. Rare. Mixed 
pine-hardwood upland forest and roadside 
ditches in the Homochitto River basin. 

Tradescantia ohiensis Raf., A1889. Locally com- 
mon. Roadside ditches and yards. 

CONVALLARIACEAE 

Polygonatum biflorum (Walter) Elliott, A2065. 
Infrequent. Upland hardwood forest and 
ravines, 


CYPERACEAE 

*Bulbostylis barbata (Rottb.) C.B. Clarke, A23715 
Rare. Cemetery in the southeastern corner 
of the county 

Bulbostylis capillaris (L.) Kunth ex C.B.Clarke,A2325. 

cally common. Sandy pine forest and 

sandbars in the Homochitto River basin. 

Carex abscondita tla A1012. Infrequent. Ma- 
ture mixed fore 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


Carex albicans var.australis (L.H. Bailey) Rettig [=C. 
ee ee oe Sveti Hard- 
ood ra and ce 
ie pie ee ae Infrequent. 
Cypress swamps and spring-seeps 
Carex atlantica L.H.Bailey [including C. atlantica 
ssp. capillacea (L.H. Bailey) Reznicek], A909. 
Common. Spring-seeps and mesic ravines. 
Carex basiantha Steud., A1013. Infrequent. Up- 
land hardwood forest, ravines, and bottom- 
land hardwood forest. 
Carex blanda Dewey, A1998. Infrequent. Ravines 
ane uplene hardwood forest. 
ar. bromoides Schkuhr ex Willd., 
A2005. Rare. senna swamp along the 
st Fork Amite River. 
Carex pan ane Schwein., A103. Rare. Hard- 


Carex cherokeensis Schwein., A634. Infrequent. 
Thick loess and outcrops of clayey subsoil, 
primarily in the Homochitto River basin. 

Carex complanata Torr.& Hook.,A733.Common. 
Sandy bottomland, floodplains, clearcuts, 
and pine forest. 

Carex corrugata Fernald [=C. amphibola var. 
turgida Fernald], WA&M 71147.Rare.OQutcrops 

yey subsoil in longleaf pine forest of 
the Homochitto River sewe 

Carex crebriflora Wiegand, A Common. 

ides, upland and ecu hard- 
wood forest, and ravines. 

Carex debilis Michx., A847. Abundant. Upland 
mixed forest, longleaf pine forest, outcrops 
of clayey subsoil in the Homochitto River 
basin, bottomland and associated mid- 
slopes, crevices of Nyssa biflora bark. 

Carex oa Muhl., A1077. Rare. Cypress 
S p in crevices and bark of Taxodium 


dist ieee : 

Carex digitalis var.asymmetrica Fernald, A867.Com- 
mon. Ravines, upland hardwood forest, bot- 
tomland hardwood forest, and roadside 


ditches. 

Carex festucacea Schkuhr ex Willd., A735. Infre- 
quent. ee outcrops and upland hard- 
wood fo 

Cre faces Dewey, A696. Apu ney Boe 

d forest, outcrops of clayey subsoil in 
. ae River basin, and roadside 
ditches 


667 


Carex frankii Kunth,A1054.Common.Bottomland 
openings, moist ditches in pine forest, up- 
land loess, and oxbow lakes. 

Carex glaucescens Elliott,A1779. Infrequent. Moist 

as associated with pine forest. 

Carex intumescens Rudge,A732. Locally common. 
Swamps, mesic hardwood ravines, and bot- 
tomland oak flats 

Carex joorii LH.Bailey, A1617. Locally common. 
Bottomland hardwood forest and swamps. 

Carex leptalea Wahlenb.,A736. eee common. 
Spring-seeps and associated swamps. 

Carex lonchocarpa Willd., A2032. erecta 
Stream banks and swamps, primarily in the 
southeastern part of the count 

Carex longii Mack., WA&M 71218.Common. Bot- 

mland forest and mesic ravines. 

Carex louisianica L.H.Bailey, A127. Rare. Old bea- 
ver-impounded pond. 

Carex lupulina Muhl.ex Willd.,A 1445. Locally com- 
mon. Riverbanks, swamp margins, and bot- 
tomland openings 

Carex lurida nea A1286. Locally common. 
Spring-seeps, bottomland forest, and 
swamps. 

Carex muehlenbergii Schkuhr ex Willd. var 
muehlenbergii,A1148. Rare. Saffell outcrops. 


mw 
baal 
Om 


Carex rosea Schkuhr ex Willd., A101 1. Infrequent. 
Saffell outcrops and hardwood ravines 
Carex striatula Michx., A1033. Infrequent. Hard- 
wood ravines and bottomland forest. 

Carex texensis (Torr.) LH. Bailey [=C. retroflexa var. 
texensis (Torr.) Fernald],A747.Common.Grav- 
openings, yards, and alongside 

uildin 

mee ee Wahlenb., A1047. Common. 
Bottomland hardwood forest and Saffell 
outcrops. 

pees a Michx., WA&M 71055. Infrequent. 

ell outcrops and pine cutov 

ane compressus L., A241. Rare. padaial in 

Liberty 


a atte : .) Wood, WA&M 70959. Infre- 
t. Pine 
By. aes oe ‘Muhl, A2355. Infrequent. 
Sandy river margins 
*Cyperus esculentus L.,A2332. Infrequent. Sandy 
river margins and cultivated fields. 


668 


Cyperus haspan L., Al020. hale Roadside 
ditches and pond margin 

*Cyperus iria L., WA&M 71065. ieccnen Road- 
side ditches. 

gi polystachyos Rottb., A2359. Infrequent. 

y river margins 

cers pan Steud.,A1027.Common. 

Moist ditc 


Cyperus retrorsus Coon FA 123.Common. Old 
fields, pine forest, yards, and roadsides. 
*Cyperus rotundus L., A556. Locally common. 


s, ditches, and riversides. 
Cyperus virens Michx., WA&M 71174. Common. 
rgins and ditches in pine forest. 
Fleocharis baldwinii (Torr.) Chapm.,A 1929. Infre- 
quent. Pond margins 
Eleocharis aaa a A118/. Rare. Wet 
ditches in pine 
chs os Se A1346.Common. 
wamps,and roadside ditches. 
Eleocharis ace (Michx.) Roem. & Schult, 
sey Infrequent. Ditches in ied pine 
forest of the Homochitto River bas 
ee aes (L.) Roem. eta A2243. 
Infrequent. Bald-cypress swamp of the 
Homochitto River and pond margins. 
mobristylis decipiens Kral, A2422. Rare. Ruderal in 


Cyperus strigosus L.,A1710.Common.Bottomland 
edges, 


Liberty, 

Fimbristylis dichotoma (L.) Vahl,A2401.Infrequent. 
Ditches in sulle forest of the Homochitto 
River bas 

eae ails (L.) Vahl [as commonly 

din the U.S, or F littoralis Gaud.], A1519 
Common. Moist areas in pine forest sit) 
sandy river margins. See Strong & Kral (1999 
for a discussion of the nomenclatural diffi- 
culties associated with this taxon 
yy fOMepIenG Vahl, ue MIRE SH 


Isolepis carinata Deak & Arn. ex Torr. [=Scirpus 
oilolepis (Steud.) Gleason],A734. Locally com- 
.Moist open areas. 
*Kyllinga brevifolia Rottb.,A2420.| 
as in towns. 
*Kyllinga odorata Vahl, Al 783. Infrequent. Yards. 
Kyllinga pumila Michx., A2360. Infrequent. Sandy 
river margins 
Rhynchospora caduca Elliott, A7574. Abundant. 


freq Rud- 


BRIT.ORG/SIDA 19(3) 


Bottomland fields, el ae spring- 
nd roadside ditch 
Hivacnosoer ae ae & Gale, 
SJones et al. 19951 (MISS!). Rare. Swamps 
fara corniculata (Lam.) A. Gray, A2035. 
Infrequent, ies wet ditches and 


mp ma 
ie eat ee (Chapm.) Small, WA&M 
71210. Infrequent. Upland clearcuts in the 
seis part a county. 
hy .) Vahl, WA&M 71226. 
Gomnivan: pele fields and roadside 
ditches 
Riynenestora inexpansa (Michx.) Vahl, WA&M 
71200. Infrequent. Moist oe openings, 
such as old ditches in pine fores 
ilies mixta Britton, A617. en 
argins of dry, upland longleaf or mixed 
oale pine forest. 


ava Ir R 


Loh, yf 

. globular 
Var. recogni ita Gale], A997 samen Roadsides 
ields, pine forest, and outcrops of clayey 
subsoil in the Homachitto River basin. 

Scirpus cyperinus (L.) Kunth, A1317. Common. 
Roadside ditches, pond margins, swamp 
margins, and Beaveh impoundments. 

Scleria ciliata Michx., A444. Infrequent. Longleaf 

pine forest. 

cleria Acne Michx.,A905.Common. Upland 

and mland hardwood forest, ial 

ae layey subsoil, and pine 

cleria pauciflora Muhl. ex se var. paucl ie 

A886. Infrequent.cem 

Scleria tri enka aa Wit ie 71134. Infre- 
quent. Pine fore 


DIOSCOREACEAE 
Dioscorea quaternata Walter ex J.F. Gmel., A1669. 
Infrequent. Mesic hardwood forest and rich 
ravines. 
Dioscorea villosa L., A2107. Infrequent. Saffell 
utcrops 


5G 


ald 


GRAMINEAE 
pen Se (Walter) B.S.P,A7930.Common. 
st, roadsides. and fields. 


Aas sect anal A1596.\nfrequent. 
t roadsides, and Saffell 


outcrops. 
Alopecurus carolinianus Walter, A7446. Infrequent. 
Pond margins and cattle pasture. 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


Andropogon glomeratus var. pumilus Vasey ex 
Dewey, A1691. Common. Clearcuts and old 
fields. 

Andropogon gyrans Ashe var. gyrans, Ray 5471 
(MISSA; specimen not seen). Cited in Camp- 
bell (1983). Now reportedly extirpated from 
MS (Kartesz 1999) 

Andropogon ternarius Michx.var.ternarius,A 1650. 
Infrequent. Pine forest. 

Andropogon virginicus L. var. virginicus, A1627. 
Abundant. Old fields, pine forest, clearcuts, 


and roadsides. 
Aristida dichotoma sake var.dichotoma,A1788. 
frequent. Pine for 
Aristida longespica a var. longespica, A1787. 
Common. Pine forest. 
Aristida oligantha Michx.,A2801.Infrequent. Pine 
forest and roadsides 
sees sane Poir: var. purpurascens, 
undant. Pine forest. 
a hispidus (Thunb.) Makino, A2328. 
andy margins of the Homochitto River. 
Arundinaria ae (Walter) Muhl 
A mmon. Bottomland forest, ravines, 
and iad hardwood and mixed for 
*Avena sativa L., A855. Infrequent. ola falde: 
Axonopuls fissifolius (Raddi) Kuhlm., A2446. Infre- 
quent. Fields and yards. 
*Briza minor L.,EA 24?.Common. Yards, old fields, 
and disturbed areas in town. 
*Bromus catharticus Vahl, A722. Infrequent. Dis- 
Rice: areas, eC arc Uts and roadsides. 
Schrad., A856. Abundant. 
Pastures, fields, and roadsides 
ie teeth latifolium (Michx.) H.0.Yates, 
Al3 cally common. Riversides and road- 
side di a 
Chasmanthium laxum (L.) ie Yates, A1337. In- 
frequent. Forest of alls 
Stora sessi es a H.O. Yates, 
A1680. Abundant. Upland hardwood and 
mixed forest, ravines, and less commonly 


ssp gigantea 


pine ; 
*Cynodon dactylon (L.) Pers., A854. Abundant. 
Pastures, old fields, yards, roadsides, and riv- 
ides 


*Dactylis glomerata L., Al 173. Infrequent. Road- 
sides and disturbed ey areas. 

Danthonia sericea Nutt 67. Common. 
Roadsides 


669 


* Digitaria ciliaris (Retz.) Koeler,A1447.Common. 

Riversides and bottomland edges 
ci meee (Schreb.) Muhl. var. 
mum, A1599. Common. Yards and 


ee ides. 
* Echinochloa colona (L.) Link, A1405. Common. 
se neles pastures, and 1 pine oF res = 


A2335. Rare. Sandbars of the se in 


River. 

*Eleusine indica (L.) Gaertn., A2424. Infrequent. 
Ruderal areas in towns 

Elymus virginicus L., A916. Abundant. Bottomland 
open areas, roadside ditches, and riversides. 

Eragrostis aie: (L.) Nees, A2805. Infrequent. 
Pine forest and roadsides. 

a a ottii SWatson,A1625.Common.Pine 

roadsides, and ditches 

ets hirsuta (Michx.) Rees A2331. Infre- 
quent. Sandy river margins and associated 
pine forest 

see al a ee B.S.P, A2829. Infre- 
quent. Pond marg 

bogs refacto ‘ub Scribn., A2284. Com- 

e forest and rarely old fields 

Pe meee is (Pursh) Steud.,A/ 1546. Abun- 
ee nt. Pine forest, old Tae and roadsides. 

unro) Hack., A1524. 
Common. Yards, fields oe and cem- 
eteries. 

Hordeum pusillum Nutt., A726. Common. Old 
fields and pastures 

Leersia lenticularis 
Baldcypress swamp margins. 

Leersia oryzoides (L.) Sw.,A1794. Infrequent. Riv- 
ersides. 

Leersia virginica Willd., A1727. Common. Moist 
bottomland forest, swamps,and spring-seeps. 

Lolium arundinaceum (Schreb.) Darbysh. [=Festuca 
arundinacea Schreb.], A2013. Infrequent. 
Roadsides and farms, . in the 
northeastern part of the c 

*/olium perenne L., A583. Aceon Fields and 
roadsides. 

Melica mutica Walter, A674. Locally common. 
Saffell outcrops, gravelly forest, and bottom- 
land roadsides. 

bleak: schreberi J.-.Gmel., A2834. Infre- 

t. Roadside ditches and bottomland 


hl be neh 
MIOCTHOC OP HUPOhdes 


A1l307. Rare. 


a 


670 


Oplismenus hirtellus ssp. setarius (Lam.) Mez ex 
E n (=O. setarius (Lam.) Roem. & Schult], 
A1464. Infrequent. Bottomland forest, 
swamps, spring-seeps, and shaded yards. 

Panicum: Taxonomy follows Lelong (1986). Rec- 
ognition of Dichanthelium without 
Phanopyrum will require the transfer of nu- 
merous darn to other genera, or 

ichanthelium with 

ai require transfer of 
Dichanthelium to Phanopyrum (the older 
name) or conservation (cf. Zuloaga et al. 
[1993] and phylogenetic reanalysis [Alford, 
unpubl. data)). 

Panicum aciculare Desv.ex Poir.[=Dichanthelium 
aciculare (Desv. ex Poir.) Gould & C.A.Clark, 
in part], A426. Abundant. Pine forest and 
cemeteries. 


Lelong) [=P lanuginosum Elliott, Dichan- 
thelium acuminatum (Sw.) Gould & CA. Clark, 
in Soe ere Pine forest, clear- 
cuts, and roadsides 
+Panicum acuminatum var. fasciculatum (Torr. 
Lelong [=Dichanthelium acuminatum va 
fasciculatum (Torr.) Freckmann],A 1979. |Infre- 
quent. Clearcuts. 
Panicum acuminatum var. lindheimeri eae 
elong [=Dichanthelium acuminatum 
Ht le (Nash) Gould & C.A.Clark], Wee 
_Infrequent. Clearcuts. 
nee acuminatum var. unciphyllum (Trin.) 
Lelong aunene meridionale (Ashe) 


— 


YS 


= 


ae nn], WA&M oe Common: 
Clearcuts, side forest, sandy t l,and 
ravell 


Panicum anceps eee lincluding P anceps var. 
rhizomatum (Hitchc.& Chase) Fernald],A 1198. 
Common. Roadside ditches and pine forest. 

tPanicum ee Elliott [=D eee 
aciculare (Desv. ex Poir.) Gould & C.A.Clarke, 
in part], A2022. Infrequent. ae pine oan 

+Panicum boscii Poir. [=Dichanthelium boscii 
(Poir.) Gould & C.A.Clarke],A1985.Common. 
Upland hardwood and mixed forest and 
ravines 

+Pani Schult. var.commutatum 
[=Dichanthelium commutatum (Schult. 
Gould, in part], WA&M 71057. Infrequent. Pine 
forest 


—_ 


BRIT.ORG/SIDA 19(3) 


+Panicum dichotomum var. ramulosum (Torr) 
Lelong [=Dichanthelium dichotomum L. var. 
dichotomum, in part], A1670. Common. Bot- 
tomland hardwood forest and low-water 


swamps. 

tPanicum gymnocarpon Elliott [=Phanopyrum 
AG Elliott) Nash], A1822. Locally 

mon.Old river beds and swamp margins. 

o. cum hians Eliot [=Steinchisma hians (El- 
liott) Nash], A206 1. So .Swam 
margins and pond ma 

Panicum laxiflorum tes hy eanietin 
laxiflorum (Lam.) Gould, P. xalapense Kunth], 
A1064. Common. Upland _ forest, bot- 
tomland edge, and cemeterie 

+Panicum polyanthes Schult. [=D pieantheliun 
sphaerocarpon var. isophyllum (Scribn.) 
Gould & C.A. Clark], A969. Infrequent. Mixed 
forest. 

tPanicum ravenelii Scribn. &  Merr. 
[=Dichanthelium ravenelii (Scribn. & Mert.) 
Gould], A450. Rare. Outcrops of clayey sub- 
soil in longleaf pine forest of the Homochitto 
River basin 

Panicum rigidulum Bosc ex Nees, A1598. Locally 
common. Bottomland forest edges, spring- 
seeps, swamp margins, and roadside ditches. 

+Panicum scoparium Lam. [=Dichanthelium 
scoparium (Lam.) Gould], A1600. Common. 


— 


e 
+Panicum sphaerocarpon Elliott [=Dichanthelium 
sphaerocarpon (Elliott) Gould, in part), A872. 
cally common. Pine forest and clearcuts. 
+Panicum strigosum Muhl.ex Elliott var. strigosum 
[=Dichanthelium strigosum (Muhl. ex Elliott) 


Freckmann var. strigosum], — Rare 
Loblolly and shortleaf pine fores 
tPanicum tenue uhl. [= ae 


dichotomum var. tenue (Muhl.) Gould & C.A. 
Clark], A449. Rare. Outcrops of clayey subsoil 
in cen pine forest of the Homochitto 
Rive 

ae verrucosum Muhl., A1597. Common. 
Roadside ditches and pond margins. 

Panicum virgatum L. var. virgatum, A1545. Com- 
mon. Old fields and roadsides 

Paspalum boscianum Fliggé,A1614. Infrequent. 
Roadside ditches. 

*Paspalum dilatatum Poir., A962. Abundant. Pas- 
tures, old fields, roadsides, and yards. 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


Paspalum floridanum Michx.,A1748. Infrequent. 
Pine forest and roadsides 

Paspalum laeve Michx., A1655. Infrequent. Pine 
forest in the southeastern part of the county. 

*Paspalum n m var. saurae Parodi, A939. 

bundant. Pastures, old fields, disturbed ar- 
as in town, and yards. 

Paspalum plicatulum Michx., WA&M 71132. Rare. 
Pine forest of the Homochitto River basin. 

Paspalum setaceum Michx., A957. Infrequent. 
Ene forest oh the Homochitto River basin. 

*P i Steud., A938. Common. Road- 
ade: and old fields 

*Phalaris angusta Nees ex Trin., A860. Rare. Road- 
S| itches 

Phalaris caroliniana Walter, A584. Infrequent. 

oadsides and old fields 

*Phyllostachys aurea Carriére ex Riviere & 
CRiviére, A1224. Infrequent. Bottomlands in 

the southwestern part of the county and 
persisting around old h 

Piptochaetium avenaceum a. ; pare [=Stipa 
avenacea L.],A1897. Rare. Steep ravine of the 
Homochitto River basin. 

*Poa annua L.,A210.Locally common. Yards, dis- 

rbed areas in town, and fields. 

Poa autumnalis Muhl. ex Elliott, A275. Common. 
Bottomland forest, upland hardwood and 
mixed forest, Saffell outcrops, and yards. 

Saccharum baldwinii Spreng. [=Erianthus 
strictus Elliott], A1475. afiequent Bottom- 
| dges and riversides. 

Saccharum brevibarbe var. contortum (Elliott) 


R.D.Webster [=S. contortum (Elliott) Nutt., 
Erianthus contol pa eaainl ex x Elliott], A1648. 

clear-cuts 
ey dees 


Saccharum nose (Walter) Pers. [=Erian- 
thus giganteus (Walter) PBeauv.], A1664. In- 
frequent. Pond margins and ditches in pine 


forest. 

Sacciolepis striata (L.) Nash, A2358. Rare. Sandy 
gin of the West Fork Amite River 
seta scoparium (Michx.) Nash, A1626. 

ndant. Pine forest, clearcuts, and 
alte 
*Setaria glauca (L.) PBeauv., Al404. Common. 
Pastures, old fields, and moist roadsides. 
Setaria parviflora (Poir.) Kerquélen [=S. geniculata 
’Beauv.], A1647. Rare. Xeric roadsides 


671 


Sorghastrum elliottii (C.Mohr) Nash, A2389. Infre- 
nt. Pine forest 
* Sorghum halepense (L.) Pers.,A1117.Common. 


Roadsides. 
Sphenopholis filiformis (Chapm.) Hitchc., A2550. 
are. Cemetery in the eastern half of the 
county. 
sa le obtusata (Michx.) Scribn., A587. In- 
fre 


oadsides, pastures, and disturbed 


n town. 
hea: hee (L.) R.Br, A1732. Common. 
Fields, pastures, yards, and disturbed areas 


in town. 

Sporobolus junceus (PBeauv.) Kunth,A 1645. Infre- 
quent. Pine forest in the eastern part of the 
county. 

* Stenotaphrum secundatum (Walter) Kuntze, EA 
265.Common. Lawns and fields. 

Tridens flavus (L.) Hitche.,A 7484. Common. Road- 
sides and fields. 

Tripsacum dactyloides (L.) L.,A1227. Infrequent. 
Roadside ditches, riverbanks, and pine 
forest 

* Triticum aestivum L., A835. Infrequent. Fields and 
food plots. 


sas ee aie (Munro ex C. Wright) R 

=Brachiaria platyphylla hae ex 
ae Nash], A7215. Infrequent. Culti- 
vated fields 

*Vulpia myuros (L.) C.C.Gmel., A721. Common. 

Disturbed areas and roadsides. 


fl A664. 


Infrequent. Roadsides 
HEMEROCALLIDACEAE 
*Hemeracallis fulva (L.) L., A814. Infrequent. Road- 
sides and old home sites. 
HYACINTHACEAE 
*Muscarineglectum Guss.ex Ten.,A376. Rare. Fal- 
low soybean fields. 


HYDROCHARITACEAE 

Vallisneria americana Michx.,A337. Locally com- 
mon.In channel and margins of both forks 
of the Amite River and its tributaries, occa- 
sionally in spring-fed streams. 


HYPOXIDACEAE 

Hypoxis hirsuta (L.) Coville,A2021.Infrequent. Pine 
forest, primarily those that are occasionally 
burned, in the Homochitto River basin and 


in the southeastern corner of the county, 
rarely along sandy stream margins of the 
Homochitto River basin. 

IRIDACEAE 

*Gladiolus x gandavensis Van Houtte, A689. In- 
frequent. Roadside ditches. 


Gladiolus communis ssp. eel hee (Mill) 
A.PHam., A529. Rare. Old house 
Iris brevi a is Raf., A790. Rare. ai patton 
land hardwood forest along the East Fork 
Amite Riv 
*Iris eee: L.,A537. Infrequent. Perennially 
wet ditches and shallow abandoned ponds. 
Sisyrinchium angustifolium Mill, A977.Common, 
Pine forest, midslope mixed forest, and 
ungrazed pasture land. 
~ inchium exile EP. Bicknell (S. rosulatum E.P. 
nell,in part), A556.Infrequent. Roadsides 
ae pastures. Species taxonomy follows 
Shinners (1962) 
Sisyrinchium mucronatum Michx., A1880. Rare. 
Pine forest in the eastern half of the county. 
Sisyrinchium rosulatum E.P. Bicknell, A637. Com- 
Roadsides and pastures. 


JUNCACEAE 

Juncus eel Michx., A895. Infrequent. Bot- 
d fields and openings. 

eae ea Elliott, A638. Locally common. 
aist areas and roadside ditches. 

Juncus en. Engelm., WA&M 70996.Rare. 


Juncus bufonius L.,A1948. Rare. Bottomland hard- 


ce 
Juncus coriaceus Mack.,A1446.Locally common. 
wamps, roadside ditches, riverbanks 
moist fields, and hardwoocd forest. 
Juncus diffusissimus Buckley, A973. Common. 
Open swampy areas, roadside ditches, and 
moist bottomland fields 


|| 
all 


Swamps, 


Juncus effusus L.,A620.L 
dside ditches, and moist fields. 
ue ee Rostk., WA&M 71161. Infre- 
nt. Moist open 

hee repens Michx., io on ateale Pond 
margins and perennially wet ditc 

Juncus tenuis Willd., A699. Abundant. ere 
ditches, forest edges, old woods roads, 
clearcuts, and swamps. 


BRIT.ORG/SIDA 19(3) 


Juncus validus Coville, A022. Locally common. 
mps and roadside ditches 
Luzula acuminata var. carolinae (S.Watson) 
Fernald, A2522. Rare. Steep gravelly bluff 
overlooking the East Fork Amite River. 
Luzula bulbosa (Wood) Smyth and L. Smyth, 
A297. Infrequent. Roadsides in town 
Luzula echinata (Small) FJ. Herm.,A277.,cCommon. 
affell outcrops, mature hardwood forest, 
bottomland hardwood forest, and yards. 


LEMNACEAE 
lemna ae Welw., A1351. Infrequent. 
ant ditches, small streams, beaver-im- 
poun a and slough 
Lemna va eee eulbet A258] a Slough of 
the East Fork Amite River. 


LILIACEAE 

Chamaelirium luteum (L cm A. a Infrequent. 
Upland hardwood fore 

Lilium michauxii Poir., A2018 eae Upland 
hardwood forest. 

MELANTHIACEAE 

Aletris aurea be ae Infrequent. Longleaf 
pine fores lI tcrops of clayey 
subsoil in ie Homechitte River basin. 

Melanthium virginicum L., A1298. Rare. Spring- 
seep in the Homochitto River basin. 

ee 


nance) 


m WT. Aiton, A1369. Infre- 
quent. oa on Magnolia grandiflora 
along rivers, especially in the East Fork Amite 
River drainage. 

Listera australis Lindl, A423. Rare. Bottomland 
hardwood forest along West Fork Amite River. 

Malaxis unifolia Michx., A845. Rare. Mesic upland 


ardwood forest. 
Platanthera ciliaris (L.) Lindl, A47.Rare. Moist hard- 
wood drains between longleaf pine domi- 
nated ridges of the Homochitto River 


Platanthera flava (L.) Lindl. var. flava, A1328. In- 
frequent. Spring-seeps and low-water 
swamps 

Spiranthes cernua (L.) Rich.,A60. Rare. Spring-seep 
and low-water swamp of the West Fork 


Amite River. 
Spiranthes ovalis Lindl.,A1668. Rare. Mesic hard- 
wood ravine. 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


Spiranthes praecox (Walter) S.Watson, A1941. 
frequent. Pine forest. 
Spiranthes tuberosa Raf.,A1191.Infrequent.Cem- 
eteries and yards 
Spiranthes vernalis Engelm. & A.Gray, A2079. 
Fields, yards, roadsides, and 


cemeteries. 
Tipularia discolor (Pursh) Nutt.,A 1388. Infrequent. 
Upland hardwood forest. 


PALMAE 
Sabal minor (Jacq.) Pers., A1211. Infrequent. 
Spring-seeps, bottomland hardwood forest, 


and swamp margins. 
POACEAE (see GRAMINEAE) 


POTAMOGETONACEAE 
pacer diversifolius Raf,,A2294. hese les 
Tributaries and open spring-fed waters 
hate te River drainage. 
eee pulcher Tuck., A2582. Rare. Open 
ugh of the East Fork Amite River. 


SMILACACEAE 

milax bona-nox L., A889. Common. Upland for- 
est, secondary forest, and Saffell outcrops. 

Smilax glauca Walter, A1920,. Abundant. Pine for- 
est and roadsides 

Smilax laurifolia L., A1839. Locally common. 
Swamps, swamp margins, and bottomland 
hardwood forest. 

Smilax pulverulenta Michx., a Infrequent. 
Upland hardwood fores 

Smilax pumila Walter, Al Locally common. 
Saffell outcrops and unburned longleaf pine 
forest in the Homochitto River basin and 
southeastern corner of cou 

Smilax oe ia L., A2044. naneres Forest of 

alls 
Smilax va eer A1829. Abundant. Forest 
of alls 

Smilax tamnol nee L.,A1162. Abundant. Upland 
forest, secondary forest, bottomland forest, 
and Saffell outcrops. 

Smilax walteri Pursh, A 1840. Infrequent. Swamps 


SPARGANIACEAE 
a americanum Nutt., aedae, iS 
uent.Swa 
water near forest, and perennially- -wet road- 
side ditches. 


amps, marshes, | 


TRILLIACEAE 

Trillium eed J.D. Freeman, A300. Infre- 
quent. Bottomland hardwood forest and 
ae in rich ravines. 


TYPHACEAE 
Typha latifolia L., A881. Infrequent. Roadside 
ditches and pond margins 


UVULARIACEAE 
Uvularia perfoliata L., A1885. Infrequent. Mature 
hardwood forest and steep ravines. 


XYRIDACEAE 

*Xyris jupicai Rich., A1657. Rare. Old farm ponds 

in southeastern corner of county. 

Xyris laxifolia var. iridifolia (Chapm.) Kral [=x. 
iridifolia Chapm.],A1297. Infrequent. Spring- 
seeps, Nyssa swamps, and perennially-wet 
ditches in full sun. 

MAGNOLIOPHYTA: 
MAGNOLIOPSIDA 
ACANTHACEAE 


Hygrophila lacustris (Schltdl. & Cham.) Nees, 
A1109.|nfre Sah Riverbank sand and gravel, 


ae 


often submerged. 
Justicia ae var. Gane (Chapm.) R.W.Long, 
A7 cally common. Riverbank sand and 
eee margins, and swamp margins. 
Ruellia caroliniensis (J.F.Gmel.) Steud. var. 
caroliniensis, A920. Common. Fields, open 
bottomland, and roadsides. 


ACERACEAE 
saat barbatum Michx., A2115. Infrequent. Mature 
wood or mixed a primarily in the 
i a River bas 
eee L,A395. en River-banks and 
andy bottomland forest. 
Acer rubrum L. var. rubrum, A259. Abundant. Forest 
of all sorts 
Acer rubrum L. var. drummondii (Hook. & Arn. ex 
Nutt.) Sarg.,A7076. Infrequent. Swamps and 
bottomland, particularly along the 
ochitto River. 
Acer saccharinum L.,A2025.Rare. Bottomland for- 
st in the southeastern corner of the county. 
ADOXACEA 
+Sambucus canadensis L. [=S. nigra ssp. 
canadensis Bolli], A792. Abundant. Mesic 


roadsides, fencerows, and edges of bottom- 
land forest. Rank is arbitrary here, and | 

e to maintain the historical usage. 
Viburnum nudum L., AS69. Locally common. 


Swamps. 
Viburnum rufidulum Raf. A433. Infrequent. Saffell 


ea scabrellum Te & A. Gray ex Chapm. 

i: ytatum var. scabrellum Torr.& A. Gray], 

 exiea _ Upland forest, ravines, and 

sites outcrops. Species taxonomy follows 
McAtee (1956). 

MARANTHACEAE 
* Alternanthera philoxeroides (Mart.) Griseb., A795. 
Infrequent. Bottomland fields and roadside 


itches. 
*Amaranthus spinosus L.,A1220.|Infrequent. Pas- 
tures and cultivated fields. 


ANACARDIACEAE 

Rhus peace: L.,A1739. Abundant. Pine forest, 
secondary forest, fencerows, and roadsides. 

Rhus es L., A941. ale Pine forest, 

s,and ro 

eee ae 5 Kuntze Al 126.Abun- 
dant.Widespread, but e 
secondary forest and nb rned pine or up- 

and hardwood fores 

Toxicodendron bese Mill. [=7. toxicarium 

Gillis, =f. quercifolium (Michx.) Greene], 

All 185. See Pine forest, | ilyin the 
southeastern part of the count 

Toxicodendron vernix (L.) Kuntze, A1705. Rare. 

p along the East Fork Amite River. 

ANNONACEAE 

Asimina triloba (L.) Dunal, FA 54. Infrequent. Bot- 
tomland hardwood forest and Saffell out- 
crops. 

APIACEAE (see UMBELLIFERAE) 


CYNACEAE 
Trachelospermum difforme (Walter) A.Gray,A717. 
Infrequent. Sandy bottomland forest, upland 
pine forest, and upland forest edges. 
*Vinca major L.,A35/.Infrequent. Roadsides and 
around old homes. 


AQUIFOLIACEAE 
llex ambigua (Michx.) Torr. A1390. Infrequent. 


Mature hardwood forest of Saffell outcrops. 


BRIT.ORG/SIDA 19(3) 


Ilex coriacea (Pursh) Chapm., A1116. Infrequent. 
Swamps in the East Fork Amite River drainage. 
llex decidua Walter, A412. Common. Various for- 
est and forest 
llex longipes Chapm.ex viel A439. mal Mid- 
slope or upland mature for 
llex opaca Aiton, A538. aan vee forest. 
Ilex verticillata (L.) A.Gray,A510.Locally common. 
Swamps, margins of en 
ponds, and bottomland fores 
ex vomitoria Aiton, A437. a Upland for- 
sadsides, and fencerows. 


= 


ARALIACEAE 

Aralia spinosa L.,A1552. Infrequent. Upland pine 
forest and roadsides 

*Hedera helix L., A1230. inheauent Old home 
sites 


ARISTOLOCHIACEAE 
Aristolochia serpentaria L., A2082. Infrequent. 
affell ou oie ravines, bluffs, and mature 
hardwood for 
Hexastylis arifolia 
[ rum arifolium Michx.], A 


aes Small var. arifolia 
1886. Infre- 


quent 


ASCLEPIADACEAE 
Asclepias amplexicaulis Srm., WA&M 70994. Infre- 


v 


quent. Pine forest. 
ae ie en A1938. Rare. Pine 


forest. 

Asclepias perennis Walter, A1006. Rare. Old dry 
oxbow lake with baldcypress in the 

mochitto River drainage. 

Asclepias tuberosa L.,A688.Common.Pine forest 

and roadsides. 

Sh vga L., A591. Common. Forest of 

alls 
was ias etl L., WA&M 70866. Rare. Pine 
n the southeastern aes of the county. 

ie ee Raf., A1281. Infrequent. 
Longleaf pine forest and ridges of the 
Homochitto River basin 

Asclepias viridis Walter, A565. Infrequent.Longleaf 
pine forest and ridges of the Homochitto 


Aue 


ASC 


iver basin. 
Matelea carolinensis (Jacq.) Woodson, A952. In- 
Trequent Rich ravines Saffell outcrops. 
Aa (Walter) Shinners, A972. In- 
cen: Upland Lanne forest and open 
ravines. 


> 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


ASTERACEAE (see COMPOSITAE) 


BALSAMINACEAE 
Impatiens capensis Meerb., A650. Infrequent. 
Floodplains and riverbanks. 


sa hear anaheth 
andina domestica TOAD A 1678. iniequent, 
sear ts | t ngly naturaliz- 


ing to all kinds of forest. 
Podophyllum peltatum L., A338. Infrequent. Mid- 
slope hardwood forest and ravines. 


BETULACEAE 
Betula nigra L., A2506. Locally common. 
Riverbanks and pond margins. 

Carpinus caroliniana Walter ssp. caroliniana, 
WA&M 70905. Abundant. Bottomland forest, 
ravines, and mature forest of various sorts. 

Ostrya virginiana (Mill) K.Koch,A1994. Common 

e forest, bottomland forest, ravines, 
and Saffell outcrops. 


BIGNONIACEA 

Bignonia ae L., A512. Abundant. Forest, 
forest edges, and dense fencerow 

Campsis radicans (L.) Seem. ex cn A852. 
Abundant. Forest edges and fencerows. 

Catalpa bignonioides Walter, A796. Infrequent. 
Old fields, fencerows, and persisting from 
cultivation at homes and ponds. Possibly 
non-native. 


BORAGINACEAE 
Cynoglossum virginianum L. var. virginianum, 
A1001. Rare. Upland mature forest of the 
Homochitto River basin. 
*Heliotropium indicum L.,EA 184.Infrequent. Rud- 
ral areas in towns. 
Lithospermum tuberosum Rugel ex DC. Al1/4. 
Infrequent. Saffell outcrops. 
eae ae ee Engelm.,A492. ee 
semi-openings and yard 
BRASSICACEAE (see CRUCIFERAE) 
BUDDLEJACEAE 
Polypremum procumbens L., A832. Abundant. Pine 
forest, clearcut areas, road-sides, and dis- 
turbed sites 
BUXACEAE 
Pachysandra procumbens Michx., A2574. Infre- 
quent. Mixed upland forest in the Homochitto 


River basin and hardwood forest of Saffell 
outcrops in the Amite River drainage 


CABOMBACEAE 

Brasenia schreberi J.-.Gmel., WA&M 71089. Infre- 
quent.Beaver impounded ponds and old im- 
poundments of various sorts. 


CALLITRICHACEAE 
Callitriche Ses Me Pursh,A2433. ea 
ds beaver-impoundm 
Callitriche te Nutt. var. pepoi ae A255/. 
e.Cleared forest in the northeastern part 
ae county. 


CAMPANULACEAE 

Lobelia appendiculata A.DC., A874. Eine Ort 
upland sae areas, and gravelly bott 
woods ro 

Lobelia ae is haga cardinalis,A70.Common. 
Bottomland openings and riverbanks. 

Lobelia puberula Michx.,A1633. Abundant. Road- 


ThHalla 


sides. 
Triodanis biflora (Ruiz & Pav.) Greene, A365.Com- 
oadsides, ruderal areas in towns, and 


yards. 
Lek Acie .) Nieuwl., A500. Infrequent. 
d semi-open areas and yards. 
Wet marginata (Thunb.) A.DC.,A778. 
Roadsides and gravelly disturbed areas. 


CAPPARACEAE 
*Cleome hassleriana Chodat, WA&M 70900. |Infre- 
ent. Riverbanks and sandbars. 


CAPRIFOLIACEA 
ee ame A478. Abundant. Sec- 
forest, roadsides, and fencerows. 
me sempervirens L. var. sempervirens, A434. 
Infrequent. Forest edges. 


CARYOPHYLLACEAE 

Arenaria lanuginosa (Michx.) Rohrb., A713. Infre- 
quent. Saffell outcrops. 

*Cerastium glomeratum Thuill., A266. Abundant. 
Roadsides, fields, ruderal areas in towns, and 


yards. 

Sagina decumbens (Elliott) Torr. & A.Gray, A369. 
Infrequent. Old fields, yards, and ruderal ar- 
eas ne owns. 

Silene antirrhina L.,A2553. Infrequent. Roadsides 
a ms in the eastern half of the county. 

Silene eae (L.) W.T.Aiton, A17137. Infrequent. 
Saffell outcrops. 


676 


*Stellaria media (L.) Vill. A285. Abundant. Old 
fields, roadsides, ruderal areas in towns, and 
yards 


CELASTRACEAE 

ees americanus L.,A1/02. Infrequent. Up- 
land h fores 

See pete nee Hand.-Mazz.,A2800. 
Rare. Long-persisting after cultivation. 

Lepuropetalon spathulatum Muhl.ex Elliott,A 7872. 
Infrequent. Cemeteries. 


CHENOPODIACEAE 
*Chenopodium ambrosioides |., A2354. Infre- 
quent. Sandy river margins. 


CISTACEAE 
Helianthemum carolinianum (Walter) Michx., 
320. Infrequent. Pine forest and cemeteries. 
Lechea mucronata Raf., A1636. Infrequent. Pine 
forest, clearcuts, and Sia especially in 
the eastern half of the cou 
Lechea tenuifolia Michx.,A2010. a couene Road- 
and pine forest in the eastern half of 
the ne 
CLUSIACEAE (see GUTTIFERAE) 
COMPOSITAE 
Acmella oppositifolia var.repens (W. 
[=Spilan 


ie americana var. repens (Walter) 
1.Infrequent. Riversides and 


| DL 


Jansen 


A.H.Moo 
sandy tae 

Ageratina aromatica (L.) Spach, Al715. Infre- 
quent. Pine forest. 

Ambrosia artemisiifolia L., EA 61. Abundant. Old 
fields, clearcuts, and roadsides 

Ambrosia trifida L., A1708. Common. Roadside 


ditches, 

Antennaria plantaginifolia (L.) Richardson, A382. 
Infrequent. Gravelly prin roadside 
banks, ane Uplane pine fores 

Rydb., A1894. - aa ra- 
vines of the Homochitto River bas 

*Anthemis cotula L., A819. ae Cattle 
farms and old fields. 

Sa sean plantagineum Raf., A2041. Infre- 

mps and spring-se 

Baccharis aa ia L., EA 137. aon Pine 
f 


est. 
Bidens aristosa (Michx.) Britton [including B. 
polylepis S.FBlake], A1586. Common. Road- 
side ditches 


BRIT.ORG/SIDA 19(3) 


Bidens bipinnata L.,A2349. Rare. Roadside 

Bidens discoidea stele & A. Gray) Britton, A2250. 
Infrequent. Swam 

Boltonia astercides (. ; a Al442. eae: 

sides and sandy river margins. 

Boltonia diffusa Elliott,A 1273. ene Pine for- 

des. 

Chromolaena ivifolia (L.) King & H.Robins. [=Eu 
patorium ivifolium L.J, A2808. Infrequent. 
Open roadsides 

Chrysopsis mariana (L,) Elliott, A1635. Abundant. 
Upland pine forest, old fields, and roadsides. 

ie ee casa Nutt.,A7294. Infrequent. Upland 

e forest, particularly in the Homochitto 
er basin. 

aa altissimum (L.) Spreng.,A1673.Infrequent. 
Upland forest edges and gravelly clearcuts. 

Cirsium carolinianum (Walter) Fernald & 
B.G.Schub., A1939. Infrequent. Pine forest, 
especially in the Homochitto River ne and 

n the southeastern of the 
794, Hype Old 


Cirsium horridt nitiaa) Mic 


fields, roadsides, and t edges. 

Cirsium nuttallii OC. A1324. eur Upland 
pine forest and gravelly ar 

*Cirsium ae (Savi) Ten. a Infrequent. 
Clearcuts. 

crs coelestinum (L.) DC. [=Eupatorium 
coelestinum LJ], FA 33. Abundant. Old fields 
a dsides 

Conyza canadensis (L.) Cronquist, A1290. Infre- 
quent. Upland pine forest and roadsides 

tConyza parva Cronquist [=C, Snadlensis » var. 
pusilla (Nutt.) Cronquist], A1728. Common. 
Moist roadsides. Species taxonomy follows 
Cronquist (1980). 

Coreopsis lanceolata L., A441. Common. Road- 
sides and upland forest edge 

Coreopsis pubescens Elliott var. Hneens A806. 
(reaucne Roadsides and upland forest 
edge 


ges 
et, is inctoria Nutt. var. tinctoria, A1019.|In- 
orest, especially of the 

H chitto River basin. 

Coreopsis tripteris L.,A1539. Infrequent. Pine for- 
est, clearcuts, and roadsides. 

Eclipta prostrata (L.) L., A1535. Infrequent. Old 
fields and lawns. 

Elephantopus carolinianus Raeusch., A1454. Lo- 
cally common.Bottomland hardwood forest. 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


Elephantopus tomentosus L., A1329. Common. 
Upland hardwood and mixed forest. 
Erigeron annuus (L.) Pers., A936. Infrequent. 

Clearcuts, old fields, and roadsides 
Erigeron eee L,A228. Comimot: Road- 
upland hardwood forest, and 


as strigosus var. eee (Fisch.& C.A.Mey.) 
A. Gray, A598. Abundant. 


ae ides and es Ids. 
Frigeron tenuis Torr. & A. Gray, A224. Infrequent. 
ooded yards. 


aes album L.,A1283.|Infrequent. Pine for- 
t and upland mixed forest. 
ee eae (Lam.) Small, EA 132. 
t. Old fields, upland forest edges, 
dies roadsides 
Eupatorium ieee L., A2279. Infrequent. 
ine forest 
cee le iatum L.,A1605.Infrequent.Old 
ields, forest margins, and roadsides. 
nee ee x aaa Elliott, FA 107. Abun- 
dant. Clearcuts, roadsides, and pine forest. 
Fupatorium ane ium L. var. rotundifolium, 
A1275.Common. Pine fo 
Fupatorium semiserratum a A1l661.Common. 
Pine forest edges, secondary se edges, 
and old fields 
ae um serotinum Michx.,A1688. Abundant. 
es and clear 
i pas ie ica eee Nesom [=Aster 
hemisphaericus Alexander, Aster paludosus 
ssp. hemi pphesiads ierenieet) ae 
A nee Common and pine 
fore 
eta ia tend (Torr. & A. Gray) Greene, 
0. Common. Pine forest, forest edges, 


ee 


old fields 
*Facelis retusa (Lam.) Sch.Bip., A800. Infrequent. 
Roadsides and old fields 
epee incarnata (Walter) R.M. King & H. 
atorium incarnatum Walter], A 76. 
ee al outcrops. 
Gamochaeta american (Mill.) Wedd. 
[=Gnaphalium pay Mill.], A554. 


Cabrera 
(=Gnaphalium falcatum Lam.], A2556. Infre- 
quent. Fields and roadsides. 


Gamochaeta pensylvanica (Willd.) Cabrera 
[=Gnaphalium pensylvanicum Willd.J,A2393. 
Common. Roadsides, clearcuts, and sandy 
fields. 

Gamochaeta purpurea (L.) Cabrera li 
purpureum L.], A599. Abundant Road: 
sides, clearcuts, yards, and disturbed forest 

argins 

Helenium amarum (Raf.) H. Rock, A744. Abun- 
dant. Old fields, cattle farms, roadsides, and 
disturbed areas in towns. 

Helenium flexuosum Raf., A1107. Common. Old 
fields, upland pine forest, and roadsides. 
Helianthus angustifolius L.,A1603.Abundant. Pine 

orest and roadsides 

Heli _ atrorubens L, FA 109. Rare. Upland 


oadsides 

bee aie divancats L., A1197. Common. Up- 
land pine forest, especially in the Homochitto 
River basin. 
elianthus microcephalus Torr. & A. Gray, EA 35. 
Common. Old fields and roadsides. 

Helianthus resinosus Small, A1408. Infrequent. 
Pine forest and roadsides, especially in the 
eastern half of the county. 

Helianthus strumosus L.,A1266. Infrequent. Pine 
forest and roadsides 

Heliopsis helianthoides var. gracilis (Nutt.) Gandhi & 

_Thomas,A1147. Rare. Saffell outcrops. 
tHeterotheca subaxillaris var. latifolia (Buckley) 
Gandhi & R.D. Thomas, A1397. Infrequent. 
Roadsides and gravelly areas. Infraspecific 
taxonomy follows Gandhi & Thomas (1989). 

Hieracium gronovil L.,A1713. Infrequent. Hard- 
wood ,cemeteries, and pine forest 

onactis linariifolius (L.) Greene [=Aster linariifolius 
LJ,A111. Locally Outcrops of clayey 
subsoil in longleaf pine forest of the 
Homochitto River basin 

lva annua L.,A1746. Rare. Upland pine forest of 
the Homochitto River bas 

Krigia 1. (Raf.) KL. ee A207. Infre- 

t.Fields and yards. 

Krigia ondion .) Nutt.,A370. Infrequent. Yards 
disturbed areas in town, and bottomland 
fie " ; 

Krigia virginica (L.) Willd. A627. Infrequent. Edges 
of pine forest and outcrops of clayey sub- 
soil in the Homochitto River basin 


— 


Lactuca canadensis L., Al1106. Common. Road- 
sides and old fi 
Lactuca floridana (L.) Gaertn., A1394. Common. 
sides and gravelly areas. 
Lactuca graminifolia Michx., A1285. Infrequent. 
Longleaf pine forest of the Homochitto River 
basin 


*leucanthemum vulgare Lam., A851. Rare. 
Roadsides 


Liatris — (Walter) Michx., FA 5. Rare. Out- 
crops of clayey subsoil in longleaf pine for- 
est ee Homochitto River basin. 

Liatris pUuane na Michx., A1264. Locally 

on. Palo pine forest clearcuts, and 
roadsides, especially in the Homochitto River 
basin. 

Liatris squarrosa (L.) Michx., A1249. Locally 
common. Upland pine forest, clearcuts, and 
roadsides, especially in the Homochitto River 


basin. 

Liatris squarrulosa Michx. [=L. earlei (Greene) K. 
Schum.], A1637. Common. Pine forest and 
roadsides, cea in the southeastern part 
ae 


ja (L.A) Willd., A1778. Infrequent. 
Ravine: of ite Homochitto River basin and 


Saft crops. 

Mikania mee (L.) Willd., A71460. Common. 
potomland openings. 

Pa nyma (Wood) W.A. Weber & A. Léve 
[=Senecio anonymus Wood], A590. Locally 
common. Roadsides of the eastern one-third 


of the county. 

Packera glabella (Poir.) C. Jeffrey [=Senecio 
glabellus Poir.], A308. Locally common. Moist 
fields, margins of beaver impoundments, 
streamsides, and bottomland hardwood 
forest. 

Pityopsis graminifolia (Michx.) Nutt., FA 139. 
Abundant. Upland pine forest, old fields, and 
gravelly areas 

Pluchea camphorata (L.) DC. A15178. Infrequent. 
Bottomland hardwood forest and swamp 


argin 
Se a ida (L.) DC. ea Infrequent. Moist 
ditches in pine fores 
ees altissima L., i Infrequent. Ravines 
e Homochitto River basin. 
Pseudognaphalium helleri a Anderb., W. 
Allison 481 (MISS!).R 


= 


BRIT.ORG/SIDA 19(3) 


Pseudognaphalium obtusifolium (L.) Hilliard & B.L. 
Burtt [=Gnaphalium obtusifolium LJ, EA 74. 
Common. Old fields. 

Ss ae a (Walter) DC., A653. 

sides and forest edges. 
ee ia a rta is A600. ee Roadsides, 
old fields, and upland forest edges. 

Smallanthus uvedalius (L.) Mack. ex Small 
[=Polymnia uvedalia (L.) LJ, A698. Infrequent. 
Upland mixed forest, Saffell outcrops, and 
forest edges. 

Solidago auriculata Shuttlew. ex S.F. Blake, 
A16/0. Rare. Mature hardwood ravine near 
center of county. 

= caesia L.,A1606.Common. Mesic hard- 


ood forest. 

ne one L., FA 113. Abundant. Old 
fields, clearcuts, and ro es, 

Solidago a idea Elliott eee discoidea 
(Elliott) Greene], A1 706. Infrequent. Bottom- 
land ie and swamps. 

a gigantea te A1325. Common. Old 


fields. 
So seas hispida Mubl. ex Willd., A1685. Infre- 
nee oe and roadsides. 
Solid iton var.odora,A1274.Common. 
Pine forest and old fields. 
Soli eae ce Mubl. ex Willd.,A2407. Infrequent. 


a rugosa Mill [in ik var. celtidifolia 
(Small) Fernald], A7683. Abundant. Pine for- 
est, a of other forest, roadsides, and 
clea 

as er folia Muhl. ex Willd, A1559. Infre- 

uent. Saffell outcrops 

*Soliva sessilis Ruiz & Pav. [=S. pterosperma (Juss.) 
Less], A857. Infrequent. Old fields and yards. 

* Sonchus asper (L.) Hill, A370.Infrequent. Around 
buildings in towns and on farms 

Symphyotrichum adnatum (Nutt. Nesom [=Aster 
adnatus Nutt.],A7712.Locally common. Out 
crops of clayey subsoil in longleaf pine for- 
est of the Homochitto River basin. 

Symphyotrichum concolor (L.) Nesom [=Aster 
concolor L.J,A2395. Infrequent. Pine forest of 
the Homochitto River basin. 

Symphyotrichum drummondii var. texanum 
Burgess) Nesom [=Aster drummondii var. 
texanus (Burgess) A.G.Jones], A682. Infre- 
quent. Saffell outcrops. 


— 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


Symphyotrichum dumosum (L.) Nesom [=Aster 
dercieus A1744.Abundant.Roadsides and 
pine fore 
Smet lateriflorum (L.) A. Love & D. 
ve [=Aster lateriflorus (L.) Britton], A 1643.In- 
a Open pine forest, clearcuts, road- 
sides, and bottomland forest edg 
Symphyotrichum patens var. gracile Soso 
Nexom [=Aster ok var. gracilis Hook], 
A642. Rare. Outcrops of clayey subsoil in 
ae pine forest of the Homochitto River 
basi 


na 


Smo ichum ae (Aiton) Nesom var. pat- 
ns ns Aiton var. patens], A168/. 
ae ee clearcuts, and pine 


orest. 

Symphyotrichum praealtum (Poir.) Nesom [=Aster 
praealtus Poir.], A106. Infrequent. Old fields 
and pine forest. 

Symphyotrichum undulatum (L.) Nesom [=Aster 
undulatus L.],A1771.Infrequent. Upland pine 
or mixed forest. 

* Taraxacum officinale Weber ex FH.Wigg.,A225. 
Common. Old fields, yards, and disturbed 
areas ns 

Verbesina virginica L.,A1491.\nfrequent. Roadside 
ditches in the southeastern part of the 


county. 
— walteri Shinners, A1449. sees com- 
mland forest and ditche 
ene: ae (Walter) Trel. ssp. ee FA 
2. Common. Mesic openings and forest 


edges. 
“— ° texana (A. Gray) Small,A1256.Common. 
and pine forest and ee 
orc i strumarium L., A1206. infleauent 
ndy stream margins. 
* Youngia japonica (L.) DC.,A273.Infrequent.Yards 
nd disturbed areas in towns. 


CONVOLVULACEAE 
Dichondra carolinensis Michx., EA 208.Common. 

Old fields and yards. 

Ipomoea cordatotri ae Dennst.,A578.Common. 

adsides. 

Ipomoea hederacea Jacq., A2814. Infrequent. 
Roadsides and fields. 

@ */pomoea indica (Burm.f.) Merr., EA 17. Rare. 
Ditches in Liberty. 

Ipomoea lacunosa L.,A1538. Infrequent. Old fields 
and cultivated areas. 


Ipomoea pandurata (L.) G. sea A752. Abundant. 
Roadsides and pine fore 

Ipomoea sree re EA 3  osieal Roadsides 
and 


ion tamnifolia (L.) Griseb., A1104. 
Abundant. Old fields, roadsides, fencerows, 
and forest edges. 

Stylisma humistrata (Walter) Chapm., A1094. |n- 
requent. Pine forest, especially in the 
Homochitto River basin. 


CORNACEAE 
Cornus florida L., A1869. Abundant. Forest of all 


sorts. 

Nyssa biflora Walter, A1348. Locally common. 
Swamps, margins of beaver impoundments, 
and occasionally in typical bottomland 


forest. 

Nyssa sylvatica Marshall, A1396. Common. Up- 
land forest of various sorts, old fields, fence- 
rows, and occasionally bottomland forest. 

CRUCIFERAE 

* Brassica rapa L.,A377.Infrequent.Old fields and 
persisting in gardens. 

Cardamine bulbosa (Schreb. ex Muhl.) B.S.P., EA 
217. \Infrequent. Bottomland hardwood 
forest, moist semi-open forest, and Saffell 


outcrops. 
*Cardamine hirsuta L., A280. Common. Fields, 
nd ruderal areas in tow 
Lesley virginicum L.,A78 eed ale 
roadsides, and ruderal areas in tow 
eee ee lis ee Common oe 


dsides 


dot 


half of the county. 
Rorippa sessiliflora (Nutt.ex Torr.& A. Gray) Hitchc., 
00. ee Riverside gravel in the 
Amite River drain 
* Sisymbrium offici fae q ?) mag) ae re 
quent.R 


of the county. 


CUCURBITACEAE 
ale quinqueloba (Raf.) Shinners, A1487. 


em on of the county. 
*Cucumis melo L.,£A 255.Infrequent. Bottomland 


fields. 
Melothria pendula L., S. Jones et al. 19957 (MISS!). 
Rare. Cultivated fields. 


CUSCUTACEAE 

Cuscuta c Juss.ex Choisy var. pa 
A?4/0. Rare. Parasitic on roadside 

Cuscuta cuspidata Engelm., A1466. ee 
Parasitic on bottomland field herbs or 
riverbank herbs, ae in the southeast- 
ern part of the cou 

Cuscuta pentagona en A973. Common. 
Parasitic on roadside and clearcut-area 
herbs. 


DROSERACEAE 

Drosera brevifolia Pursh, A448. Rare. Outcrops of 
clayey subsoil in long-leaf pine forest of the 
Homochitto River basin. 


EBENACEAE 

Diospyros virginiana L., WA&M 70894. Abundant. 
Pine forest, mixed forest, roadsides, and old 
fields. 


ERICACEAE 

Gaylussacia dumosa (Andrews) Tort.& A. Gray var. 
dumosa,A209/. he Pine forest in the 
Souineasien part of the c 

; 1 latifoli Rare ae in the Buf- 


falo River diinaoe of the western part of the 
county. Apparently native, as referenced by 
a ai aoa (R.Richardson, pers.comm.) 
but suspiciously close to John James 
udu Arboretum in Gloster. If not origi- 
nally extending this far west, the species has 
definitely naturalized along certain streams. 
Leucothoe racemosa (L.) A. Gray, A2039. Infre- 
sheen 7 jeerlte in the south-central and 
astern part of the co 
hoe Pee .) DC, A2417, ane Swamp in 
the southeastern part of the county. 
Oxydendrum arboreum (L.) DC., A754. Common. 
Pine forest and upland mixed or hardwood 
forest. 


Ww 


Rhododendron canescens (Michx.) Sweet, A387. 
Locally common. Bottomland forest, espe- 
cially along ne swamp margins, and 
rarely in pine forest 

Vaccinium arboreum Marshall [=Batodendron 

arboreum (Marshall) Nutt.], 4546. Common. 

Mature pine forest and upland mixed or hard- 

wood forest. 

-cinium elliottii Chapm., A614, Abundant. Pine 

forest and upland mixed forest. 


Va , 


BRIT.ORG/SIDA 19(3) 


Vaccinium fuscatum Aiton, A1039. Infrequent. 
ine and mixed forest. 
Vaccinium stamineum L. [=Polycodium 
stamineum (L.) Greene], A528.Common.Pine 
and upland mixed forest 


EUPHORBIACEAE 

Acalypha gracilens A. Gray, A842. Common. Pine 
forest, gravelly areas, and roadsides 

Acalypha rhomboidea Raf., A1785. Infrequent. 
Ruderal areas in towns and yards 

Acalypha vi Bee inica L., A1412. Rare. oo in 
the northeastern part of co 

Geareic hyssopifolia (L.) ea, A1326. Infre- 


nt.Old railroad tracks and ruderal areas in 


towns. 
eee ae maculata (L.) Small, 47319. Pine 
forest, roadsides, ruderal areas in towns, and 


ae 
lad ah eke ese Small, A1353. Infre- 


uderal areas in towns 


cron a itatus Michx., A1278. Common. Old 
fields, clearcut areas, cultivated areas, and 
a aes 

Croton Slane ious var. septentrionalis Mull. Arg., 
A2322. Rare. Sandy margin of the Homochitto 
Rive 

epho corollata L., A658. oe Road- 

es, pine forest, and clearc 
ae ia cyathophora Murr, oe Infrequent. 


Roadsides. 
Phyllanthus caroliniensis Walter ssp.caroliniensis, 
Al444 4, Infrequent. Ro adsides 


*Phyllanthu L., A2850. Infrequent. Rud- 
eral areas in towns. 


Tragia cordata Michx., A1146. Rare. Saffell out- 
crops. 


ee tee Shinners, A2096. aa Pine 
n the southeastern part of the county. 
ee a ifera (L.) Small [=Sapium sebiferum 
) Roxb.], A1528. Common. Riverbanks, 
encerows, old fields, and secondary forest. 
*Vernicia fordii ese Airy Shaw [=Aleurites 
fordii Hemsl.], A1865. Common. Pine forest, 
secondary woods, and roadsides, especially 

in the eastern half of the count 


FABACEAE (see LEGUMINOSAE) 


FAGACEAE 
Castanea pumila (L.) Mill.var.pumila [in the broad 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


sense, sensu Johnson (1988)], EA 32. Infre- 
Pine forest and Saffell outcrops. 
Fagus rondo Ehrh.,A1942. Abundant. Prima- 
rily ottomland hardwood forest, ravines, 
et mature hardwood forest, but also in 
pine forest and secondary forest 

Quercus alba L., EA 152. Abundant. Primarily in 
upland hardwood or mixed forest, but also 
in bottomland forest. 

Quercus X comptoniae Sarg. [=Q. lyrata Walter x 
Q. virginiana Mill ],A1259.Infrequent. Upland 
forest in the southwestern corner of the 

ounty. 

Quercus pee Munchh., A7733. Infrequent. 
Saffell o 

Quercus falcata ne FA 24/7. ena ae 
forest, primarily associated wit 

Quercus hemisphaerica W.Bartram ae A625. 
Locally common. Ridges and mid-slope of 
ravines in the Homochitto River basin. 

Quercus incana W. Bartram, A1765. Infrequent. 
Outcrops of clayey subsoil in longleaf pine 
forest of the Homochitto River basin. 

Quercus laurifolia Michx., A1402. Common. Bot- 
tomland hardwood forest and swamps. 

Quercus aie Walter, A2366. Infrequent. Swamps 
and tomland fore 

Quercus marilandica ene A1766. Locally 

Upland forest, primarily associated 
with pine; most common on ridges of the 
Homochitto ie basin and in the eastern 
half of the co 

ar a Ae A1554. Common. Bot- 

mland hardwood forest and ravines. 
cue muhlenburgii Engelm., A971. Rare. Ma- 
ture hardwood forest in ravines. 

ee ni ae L.,A2373. Abundant. Bottomland 
hardwood forest, upland forest of all types, 
anes ,and yards. 

Quercus pagoda Raf., dee Abundant. Bottom- 
land hardwood forest and ravines 

Quercus phellos L.,A736/. eee Upland flats, 
bottomland, and yards 

Quercus rubra L., A168? Rare Ravines 

Quercus tae Buckley, A1005. Infrequent. 
Upla 


ardwood or mixed forest, ravines, 

and al outcrops. 
ace tellata War IGel 1h., EA 248. Locally com- 
n.Upland forest, primarily associated with 


pine, and yards; most common in the east- 
ern half of the county. 

Quercus velutina Lam., A977. Common. Upland 
hardwood and mixed forest and Saffell 


crops. 
Quercus virginiana Mill, EA 38.Rare. Upland road- 
sides in southwestern corner of county. 
FUMARIACEAE 
Corydalis micrantha ssp. australis pany oD 
Ownbey, A404. Infrequent. Grave 
and rarely rich hardwood forest in a sate 
eastern part of the county. 
GELSEMIACEAE 
Gelsemium rankinii Small, EA 182. Infrequent. 


Swamps. 

Gelsemium sempervirens (L.) J. St.-Hil., FA 2317. 
Common. Fencerows, pine forest, and forest 
edges. 


GENTIANACEAE 

Gentiana villosa L., FA 89. \nfrequent. Pine forest. 

oe .) Pursh, Webster & Wilbur 3277 
Sy a Infrequent. Pine forest and 


ee meee Elliott, A7775.Infrequent. Pine 
forest and roadsides 

Sabatia campestris Nutt.,A1 782. Rare. Pine forest 
in the southeastern corner of the county. 


GERANIACEAE 

Geranium carolinianum L., A282. Common. Old 
fields, disturbed areas in towns, and yards. 

*Geranium dissectum L., S. Jones & CJones 4120 
(MISS). Rare. Roadsides 


GUTTIFERAE 
Hypericum crux-andreae (L.) Crantz,A 1497. Infre- 
quent. Pine forest in the southeastern part 
of the county. 
Hyperi rummondii (Grev. & Hook.) Torr. & A. 
Gray,A1293.Common.Pine forest and Saffell 


outcrops. 

Hypericum gentianoides (L.) B.S.P.,, A2324. Infre- 

uent. Sandy pine forest 

Hypericum gymnanthum Engelm.& A. Gray, A823. 
Rare. Pine forest and clearcuts in the south- 
eastern part of the county. 

Hypericum hypericoides (L.) Crantz, A1248.Com- 
mon. Pine forest 


Hypericum mutilum L., A824. Common. Pine for- 


est and clearcuts, particularly in the south- 


rtof the 
yarn rue ae ex Willd, WA&M 
7101 requent. Stream banks and 


wamps. 

oe ei walteri (.G. Gmel.) Gleason, A1457. 
Infrequent. Swamps, spring-seeps, and bot- 
tomland hardwood forest. 


HALORAGACEAE 

i ee um aquaticum Henties A798. In- 
frequent. Slow-moving strea 

Proserpinaca palustris var. caleon Fernald, 
5. Jones et al. 13944 (MISS!). Infrequent. Pond 
nang 


—, 


Cc (AO vee 


WAM 7091 8. nfieetient Moist ditches a 
bottomland hardwood fore 


HAMAMELIDACEAE 

Hamamelis virginiana L.,A871.Common. Upland 
hardwood and mixed forest. 

Liquidambar styraciflua L.,A1851.Abundant. Hard- 
wood ravines, | land hard I f 


secondary succession, and pine forest. 


HIPPOCASTANACEAE 

Aesculus pavia L., A1899. Infrequent. Mature up- 
land hardwood forest, especially at Saffel 
outcrops, creekbanks, and pine forest. 


Acces racecars 
De aria barbara L., A785. Infrequent. Bottom- 
“lake ues forest. 
lta arborescens L., S. Jones et al. 13930 
MISS!). Rare. Steep ravines and bottomland 
ro forest along the East Fork Amite 


Pina quercifolia W. Bartram, A873. Infre- 
quent. Ravines and Saffell outcrops. 


HYDROPHYLLACEA 
a ea fies Raf,, ep 71091.Rare.Beaver- 
Wea ponds in the Homochitto River 


HYPERICACEAE (see GUTTIFERAE) 


ILLICIACEAE 
Micium floridanum J. Ellis, A1884.Locally common. 
mland hardwood forest, primarily in 
the Amite and Tickfaw River basins, but at 
least one population in the Homochitto 
River basin 


BRIT.ORG/SIDA 19(3) 


ITEACEAE 
Itea virginica L., A570. Locally common. Spring- 
seeps and swamps. 


JUGLANDACEAE 

Carya aquatica (FMichx.) Nutt,A 7065. Infrequent. 
Sandy bottomland hardwood Gig prima- 
rily in the Homochitto River basi 

Carya cordiformis (Wangenh.) K. ee Al1679. 
Rare. Ravines of mature hardwood fore 

Carya glabra Aas Sweet, A/1 900. Common. in 
land hardwood or mixed forest. 

Carya illinoinensis (Wangenh.) K. Koch, A1529. 
Common. Old fields, fencerows, and house 
sites. 

Carya pallida (Ashe) Engl.& Graebn.,A1260.Com- 

Upland hardwood, mixed, or pine 


forest. 

*Carya tomentosa (Poir.) Nutt. [=C. alba (L.) Nutt. 

Elliott], A1432. Abundant. Upland hard- 

wood or mixed forest, pine forest, second- 
ary forest, and roadsides. Nomenclature 
follows D. Stone (1997). 

*Juglans nigra L., A1247. Infrequent. Fields and 
yards. Probably all introductions from else- 
where in eastern North America. 


ABIATAE 
@ *Clinopodium gracile (Benth.) Kuntze, A782. 
Rare. Bottomland hardwood forest of the 
East Fork Amite 
Collinsonia tuberosa ae A1697. la 
Upland gravelly hardwood for 
Pre ae hederacea L., A482.| ee Shady 
ards, towns, and cem 
ne eee hispida anit eo Infrequent. 
Clearcuts, disturbed fields, upland forest 
edges, and roadsides, 
Hyptis alata (Raf.) Shinners. Rare. Referenced in 
ones (1976); specimen not seen 
*!amium amplexicaule L.,A223. eonman Yards, 
towns, roadsides, and fields. 


*lLamium pals um L., A265. Infrequent. Yards 
and town 
lycopus belts Moench, A1590. aaa 
Swamp margins roadside S 
Lycopus virginicus L.,A14 pati eae 
openings, roadside arn spring-seeps, and 
riverbanks. 
Monarda fistulosa var. mollis (L.) Benth., A820. In- 


frequent. Upl 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


Monarda punctata L. var. eee A1413.\Infre- 
Pine forest and roa 
*Perilla Hutescon: (L.) Britton, A7 me inirecuiene 
Disturbed areas, eevee a forest, and 
secondary bottomland for 
*Prunella vulgaris L., A581. an Yards, 
ere towns, and occasionally along 
forest e 
ean a albescens Torr. & A. Gray, A1284. 
Infrequent. Upland pine forest, especially in 
the Homochitto River basin 
CL viele tenuifolium Schvadl FA87.Abun- 
da e forest and roadsides 
Salvi a pend A426.Common. poreet of various 
sorts and yards 
Scutellaria elliptica Muhl. ex Spreng., A764. Infre- 
Saffell outcrops and upland hard- 


od forest. 
Scutellaria incana Biehler, A1496. Rare. Pine for- 
est in the southeastern part of the county. 
Scutellaria de L.,A574. Abundant. Forest 
5 sorts, forest edges, and roadsides. 
es es Shuttlew. ex Benth.,A481.Rare. 
ards in Liberty. 
Stachys tenuifolia Willd.,A 1458.Rare.Bottomland 
hardwood forest and spring-se 
Teucrium canadense L., WA&M 70906, oe Sandy 
nk of the West Fork Amite R 
Trichostema dichotomum L., Al oe a mon 
Pine forest, sandy riverbanks, and eager 
ditches 
Trichostema setaceum Houtt.,A1639. Infrequent. 
orest in the southeastern part of the 
ne and in the Homochitto National 
orest 


oS 
< 
< 


LAMIACEAE (see LABIATAE) 

LAURACEAE 

Lindera benzoin (L.) Blume, FA 30. Locally com- 
mon. Sandy bottomland hardwood forest in 


the 
the Amite River basin. 

Persea palustris (Raf) Sarg.,FA 36.Infrequent. Bot- 
tomland paar forest, mid-slope mixed 
forest, and ravine 

Sassafras albidum ae Nees,A7740.Abundant. 

dsides, fencerows,and hardwood forest. 

LEGUMINOSAE 

al ibrissin Durazz., A943. Common, Road- 

ards. 


s, fencerows, and yar 


683 


* Alysicarpus vaginalis (L.) DC.,A2837. Infrequent. 
Roadsides 


Apios americana Medik., FA 165.Infrequent. Bot- 
tomland forest edges, swamp margins, and 
ae 
Baptisia alban var. macrophylla (Larisey) Isely [=B. 
lactea (Raf) Thieret, =B. leucantha Torr. & A. 
Gray], A579. Infrequent. Old fields and pine 
forest 


Centrosema virginianum (L.) Benth., A756. Abun- 
dant. Upland forest edges and roadsides. 

Cercis canadensis L., A267. Infrequent. Upland 
har aie forest, Saffell outcrops, and com- 
monly cu 

a. sce os ) Greene, A 1303. 
Common. Roa ds. 


dsides and old fiel 


eee nictitans (L.) var. nictitans, 
A1561.|Infrequent. Roadsides. 

ape — L.,A1252. Abundant. Pine forest, 
forest edges, and roadsides 

bees ia rotundifolia Walter ae F.Gmel., A549. 

re. Pine forest. 

Crotalaria sagittalis L., A608. Locally common. 
Pine forest, especially in the Homochitto 
River basin. 

*Crotalaria spectabilis Roth, C.Brown 18686 (LSU)). 
Rare. Roadsides. Herbarium label indicates 
Wilkinson Co., MS, but the locality data 
(‘north of Coles”) indicate Amite Co 

Desmodium canescens (L.) DC.,A1515.Infrequent. 


— 


Roadsides. 

Desmodium ciliare (Muhl. ex Willd.) DC., A2275. 
Infrequent. Pine forest. 

Desmodium glutinosum (Muhl. ex Willd.) A.W. 
Wood, A1142. Infrequent. Mature hardwood 
forest over Saffell outcrops. 

salen ium laevigatum ees DC.,A1721.Infre- 

Upland forest e 

sees eatum DC,, 87. Common. 
Forest of various sorts, roadside ditches, and 
cemeteries. 

Desmodium nudiflorum (L.) DC.,A1145.Rare. Ma- 
ture hardwood forest over Saffell outcrops. 

es imi nuttallit eae Schub.,A2308 

e. Burned pine 
beso obtusum ae Paine A1693. 
mon.Clearcuts, pine forest,and roadsides. 
L 20. Abun- 
dant. Pine forest, bottomland openings, old 
fields, and roadsides. 


eee paniculatum 


684 


sein perplexum B.G. Schub.,A1619. Infre- 
quent. Sandy bottomland river margins and 
ae . ditches 
Desmodium ere ire DC,,A1873.Infrequent. 
Ravines of mixed forest and Saffell outcrops. 
Desmodium viridiflorum (L.) DC., A2815. Infre- 
quent. Upland pine forest and roadsides in 
the eastern half of the county. 
Erythrina herbacea L., A573. Common. Forest 
dges, gravelly areas, and Saffell outcrops. 
Galactia erecta (Walter) Vail, A2702. Rare. Pine for- 
est in the southeastern part of the county. 
Galactia volubilis (L.) Britton as interpreted by 
hors [G. regularis (L.) B.S.P. sensu 
mae lincluding G. ania M.A. Curtis], 
A1301.Common. Forest e 
Gleditsia ae L,A1246. en Upland 
roadsides and forest edges 
eres vesicarium (Jacq.) R. M. Kia A2412 
Infrequent. Bottomland o 
feces striata (Thunb.) inl [=Lespe- 
deza striata (Thunb.) Hook. & Arn.j, A1722. 
Common. Upland pine forest. 
*/athyrus hirsutus L.,A777.Infrequent. Roadsides, 
old fields, and cultivated fields. 
*/espedeza cuneata (Dum.Cours.) G. Don, FA 40. 
mon. Roadsides, old fields, and pine 
forest. 
esgoseie hirta (L.) Hornem. ssp. hirta, A1499. 
Common. Pine forest and old fields 
lopedeas procumbens Michx., A622. Common. 
Up orest edges, old fields, and road- 


2s 
(va) 


ae 

Lespedeza repens (L.) W.P.C. Barton, A550. Abun- 
dant. Pine forest. 

Lespedeza violacea (L.) Pers.,A1723.Rare. Upland 
forest edges of the Homochitto River basin. 

Lespedeza virginica (L.) Britton,A1745.Abundant. 
Pine forest, forest edges, and roadsides. 

*Medicago arabica (L.) ea A361. Infrequent. 
Ruderal areas in tow 

Medicago polymorpha L. ene in Pullen et 
al. (1968); specimen not s 

Mimosa microphylla Dr eet (=Schrankia 

microphylla (Dryand.) J.F.MacbrJ,A593.Com 

mon. Pine forest and roadsides. 

Orbexilum pedunculatum (Mill) Rydb.,A513.Rare. 
Pine forest of the southeastern part of the 
county. 


*Pueraria montana (Lour.) Merr.[=Pueraria lobata 


BRIT.ORG/SIDA 19(3) 


(Willd.) Ohwi], A7 


371.\nfrequent. Roadsides 


and adjacent forest. 

Rhynchosia reniformis DC.,A694. Infrequent. Out- 
crops of clayey subsoil in longleaf pine for- 
est of the Homochitto River basin. 

Rhynchosia tomentosa (L.) Hook. & Arn., A2098. 
Infrequent. Pine forest. 

HORNA PsCudeacdeg L.,A536.Common. Upland 
f edges and roadsides 

= obtusifolia (L.) H.S. Irwin & ron a 236. 

cally common. Cultivated fiel 

es) euies (Mill) McVaugh [= S exaltata 
(Raf.) Rydb. ex A.W.Hill?], A2357. Infrequent. 
Cultivated fields 

Strophostyles helvula 

a anks of the Homochitto Rive 

Stophostye umbellata — ex Willd.) Se 

Common. Pine for 

Stylosanthes biflora (L.) 5s canes 

orest, especially in the Homochitto River 


(L.) Elliott, A2323. Rare. 


basin. 

Tephrosia spicata (Walter) Torr. & A. Gray, A646. 
Common. Pine forest. 

Tephrosia virginiana (L.) Pers.,A563. Locally com- 
mon. Pine forest, especially dry open ridges 
and frequently burned areas. 

* Trifolium arvense L.,A474. Infrequent. Roadsides. 

* Trifolium campestre Schreb., AS85. Common. 
Roadsides, yards, and fields 

Trifolium carolinianum Michx.,A1892. Rare. Edge 

upland hardwood forest in the 
ochitto River basin. 

a lium pane Sibth., A429. Common. Road- 


sides and yards. 
* Trifolium incarnatum L., A330. Abundant. Road- 
sides and fields. 
* Trifolium ashes L.,A552. Abundant. Roadsides 
fields, 
* Trifolium spina L.,A467.Infrequent. Road- 
sides an 
*Vicia sativa da nigra (L.) Ehrh. [=V. angustifolia 
L.J], A270. Common. Old fields, cultivated 
ales, a roadsides 
icia tetrasperma (I ies, A484. Infrequent. 
Old fields 
*Vicia villosa Roth ssp. villosa, A472. Common. 
Roadsides, old fields, and cultivated fields. 
* Wisteria sinensis (Sims) DC, A1879. Infrequent. 
Roadsides, secondary forest, and old home 
sites. 


*| 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


LENTIBULARIACEAE 

tUtricularia biflora Lam. [=U. gibba L., in part, 
A2063. Infrequent. Beaver-impounded 
ponds and perennially-wet ditches. Species 
taxonomy follows Godfrey & Wooten (1981) 
and Weakley (in prep.). 


LINACEAE 

Linum medium var. texanum (Planch.) Fernald, 
A645. Abundant. Pine forest and roadsides. 

Linum striatum Walter, A1044. Infrequent. Bottom- 
land open areas and wet roadsides. 


LOGANIACEAE 

+Cynoctonum mitreola (L.) Britton [=Mitreola 
petiolata (J.F. Gmel.) Torr. & A. Gray], A2283. 
Infrequent. Bottomland openings. | do not 
consider the pirated Opera Varia (1758) of 
Linnaeus, and the Genelec name Mitreola 
taken within, validly publist ording to 
Art. 34.1 of the ICBN i” 

a marilandica (L dL, A561.Infrequent. Ra- 

nes of mature hardwood forest and Saffell 

ees 


LYTHRACEAE 
*Cuphea carthagenensis Jacq.) J.F. Macbr.,A833. 
Common. Roadsides, clearcut areas, and 
sandy bottomland openings. 
*/Lagerstroemia indica L., Al177. Infrequent. 
ncerows and secondary forest. 
Rotala ramosior (L.) Koehne,A2317.Rare.Swamp 
argins in the Homochitto River basin. 


pier aoraneanarne 
a tulipifera L., ASO9. Abundant. Bot- 
mland hardwood forest, mixed forest, ra- 
vines, and Saffell outcrops. 

Magnolia acuminata (L.) L., A2105. Infrequent. 
Saffell outcrops and ravines, most common 
in the Homochitto River basin. 

ae ia a iflora L..A714.Abundant.Bottom- 

and hardwood forest, ravines, sand occasion- 
ae bee hardwood or mixed fores 

Magnolia macrophylla Michx. A565 oc com- 

mon.Ravines of the H sin. 

Magnolia virginiana L., A687. Locally common. 

mps and perennially wet roadside areas. 

MALVACEAE 

Modiola caroliniana (L.) G.Don,A317.Infrequent. 
Disturbed areas in towns, yards, and road- 
sides 


685 


Sida rhombifolia L., A1026. Common. Roadsides 
and bottomland openings 


MELASTOMATACEAE 
Rhexia mariana L. var. mariana, A940. Common. 
ond margins, margins of beaver im- 
pounded ponds, and roadside ditches. 
Rhexia nashii Small x R. virginica L., A821. Infre- 
nt. ae margins in the Homochitto 


ie bas 
Rhexia virginica a A1588. Infrequent. Roadside 
ditches. 


MELIACEAE 


hL.,A557.Common. Mesic pine 
forest, roadsides, and 


forest, mesic secondat y 
old home sites, more common in the east- 

ern half of the county. 

MENISPERMACEAE 

occulus carolinus (L.) DC., A1134. Infrequent. 
Hardwood forest and Saffell outcrops. 

MOLLUGINACEAE 

* Mollugo verticillata L., WA&M 71108. Common. 
Riverbanks and sandbars. 

MONOTROPACEAE 

Monotropa uniflora L.,A1781. Rare. Sandy lower 
slopes of ravines in the Homochitto River 


cols 


basin 

MORACEAE 

* Broussonetia papyrifera (L.) UHér.ex Vent., EA 18. 
Infrequent. Disturbed areas in towns. 


Morus rubra L., A436. Infrequent. Upland forest, 
Saffell outcrops, and yards. 


MYRICACEAE 

Morella cerifera (L.) Small [=Myrica cerifera LJ], 
A220. Abundant. Pine forest, roadsides, vari- 
ous forest edges, and occasionally along 
swamp margins. 

NELUMBONACEAE 

Nelumbo lutea Willd., S. Jones et al. 19892 (GA!, 
MISSI). Rare. Pond in Liberty; apparently now 
extirpated 

NYCTAGINACEAE 

* Mirabilis jalapa L.,A1511.\nfrequent. Roadsides 

d old homes in the southeastern 

part of the county. 

NYMPHAEACEAE 

Nuphar lutea ssp. advena (Aiton) Kartesz & 


Gandhi [=N. advena (Aiton) W.T. Aiton],A577. 
Infrequent. Beaver-impounded ponds and 
stagnant sloughs 

*Nymphaea odorata Aiton, A1 170. Rare. Natural- 
izing in ponds from introductions from other 
parts of the state. 


OLEACEAE 
Fraxinus americana L., A915. Infrequent. Upland 
ixed forest. 


Fraxinus Rete ai ca Marshall, Al077. Com- 
mland hardwood forest and 
moist ro a. 

*Ligustrum lucidum W.T.Aiton,A 1096. Infrequent. 
Seconda y pit e forest, fencerows,and r road- 
sides. 

*ligustrum sinense Lour.,, A547. Abundant. Sec- 
ondary pine forest, bottomland forest, for- 
est edaes. fencerows, and riverbanks. 

Osmanthus americanus (L.) Benth.& Hook.f.ex A. 

ray, A2415.Infrequent. Swamps in the south- 
eastern part of the county. 


ONAGRACEAE 
Circaea lutetiana ssp. canadensis (L.) Asch. & 
Magnus, A767. Rare. Bottomland hardwood 
forest along the West Fork Amite River. 
Gaura oe. Small, S. Jones 5345 (MISS!). 
Rare. Roadside in the northeastern part of 
ay At 
Ludwigia alternifolia L.,A1344.Common.Roadside 
ditches, pond margins, riverbanks, and dis- 
turbe | 
Ludwigia ee Walter, A1360. Infrequent. 
Riverbanks and pond margins. 
Ludwigia ‘sms Walter sa aes 
Infrequent. Pond mar 
Ludwigia pee Raf, A/ 781. Rare. ft aleeene 
forest. 
Ludwigia leptocarpa (Nutt.) H. Hara, A1838. Rare. 
Pond margins. 
Ludwigia linearis Walter, A1659. Rare. Pond mar- 
gins in the southeastern part of the a 
Ludwigia palustris (L.) Elliott, WA&M 70920. 
Locally common. Pond margins, hee 
open swamps, and river mar 
Ludwigia peploides ssp.g cee Kae) PH. 
Raven, A1097. Infrequent. Ponds and open 
stream margins 
Oenothera biennis L.,A1410.Common.Roadsides 
and clearcuts 


BRIT.ORG/SIDA 19(3) 


Oenothera fruticosa L. oi fruticosa, A1975. Pine 
forest and clearc 
Oenothera laciniata Wa A473. Common. 
Clearcuts, roadsides, towns, and yards. 
Oenothera linifolia Nutt., A2095. Rare. Open pine 
he southeastern part of the county. 
Moonee speciosa Nutt., A604. Locally com- 
n dsides 


OROBANCHACEAE 
Epi poe virginiana (L.) W.P-C. Barton, A97. Infre- 
nt. Mature hardwood forest 


OXALIDACEAE 

Oxalis corniculata L., A274. Infrequent. Pine for- 
est, fields, and yards 

*Oxalis debilis var. corymbosa (DC.) Lourteig, 
A244. Infrequent. cs ruderal areas 
in towns, and yar 

Oxalis zi enii ore je Abundant. Pine forest 

affell outcrops. 
on ae A. ies 8&.Infrequent. Towns 
yards 
Oxalis violacea L., A390. Infrequent. Saffell out- 
rops and upland hardwood forest 
PASSIFLORACEAE 
Passifora incarnata L.,A1270.Common.Clearcuts, 
oadsides, fencerows, and forest edges. 
Passiflora lutea L., A1399. Infrequent. Mesic for- 


PENTHORACEAE 
Penthorum sedoides L., A1365. Infrequent. Bot- 
tomland hardwood forest. 


PHYTOLACCACEAE 
Phytolacca americana L., A673. Common. Road- 
sides and pasture edges. 


PLANTAGINACEAE 

Plantago aristata Michx., A596. Common. Pine 
forest, roadsides, old fields, and disturbed 
areas in towns. 

Plan pare een Nutt., A7896. Rare. Cem- 


ee virginica L., A555. Common. Clearcuts, 
fields, roadsides, and cemeteries. 

PLATANACEAE 

Platanus Sc bea L., A1233. Common. River 
mar mind hardwood forest, and 
ee arene 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


POLEMONIACEAE 

Phlox divaricata L., A204. Infrequent. Saffell out- 
crops and hardwood ravines. 

Phlox pilosa L., A431. Infrequent. Pine forest and 
roadsides. 


POLYGALACEAE 

Polygala incarnata L., A758. Rare. Outcrops of 
clayey subsoil in longleaf pine forest of the 
Homochitto National Forest 

Polygala mariana Mill., A606. Common. Pine 


forest. 

Polygala nana (Michx.) DC., A1935. Infrequent. 
Outcrops of clayey ssl in meng pine 
forest of the Homochitto National Forest and 
pine forest of the sgurneasicin part of the 


county. 
Polygala Haas L.,A885.Rare.Cemetery in the 
south n part of the county. 
POLYGONACEAE 
Brunnichia ovata (Walter) Shinners, A 1400. Infre 
nt. Sandy river margins and associated 


*Polyqontut pe, itosum var. 
Stewar ae nfequent. Soroniand oe 
est anes Woo 

Polygonum seis ides Michx. [=Persicaria 

asso ides be Small], A805. poe 
margins, moist bottom 

a fields and Hone roadsides, and woods 


roads. 

Polygonum lapathifolium L., A2823. Rare. Pond 
margin in the eastern part of the county. 

Polygonum pensylvanicum L. [=Persi-caria 
pensylvanica (L.) M.Gomez, including P. 
bicornis Raf], EA 66. Infrequent. Bottomland 
ditches and forest edges. 

Polygonum punctatum Elliott [=Persicaria 
punctata (Elliott) Small],A1595.Locally com- 
mon. Roadside ditches, swamp margins, river 
margins, and bottomland forest edges. 

Polygonum scandens L. [=Fallopia scandens (L.) 
Holub], A1786. Infrequent. Moist roadside 
ditches. 

Polygonum setaceum Baldwin [=Persicaria 

setacea (Baldwin) Small], A/069. Infrequent. 
Swamps and spung SEEDS 


P olygonum VIFGIETTALIOTEE CL. [=P ChoiCGliad 


(L.) Gaertn., Antenoron virginianum (L.) 
Roberty &Vautier, fOovard virginiana (L. Raf.], 


virgini Ghia 


tL 


687 


A1308. Infrequent. Upland mixed forest es- 
pecially in the Homochitto River basin, bot- 
tomland forest, and spring-se 

*Rumex crispus L., A179. nn Old fields, 
cultivated fields, fencerows, and roadsides. 

nS hastatulus Baldwin,A1877.Common, Old 
fields, roadsides, and yards. 

ee pulcher L., A779. Infrequent. Old fields 
and cultivated fields. 


PORTULACACEAE 

Claytonia virginica L.,A477.Rare.Edge of bottom- 
land hardwood forest. 

*Portulaca oleracea L., FA 126. Infrequent. Side- 
walk cracks in towns. 


PRIMULACEAE 
Anagallis minima (L.) E.H.L. Krause, A7870. Infre- 
ent. Pine forest and roadsides. 

Lysimachia lanceolata Walter, A844. Infrequent. 
Roadside ditches. 

Samolus valerandi ssp.parviflorus (Raf.) Hultén [=S. 
parviflorus Raf], A1946. Infrequent. Bottom- 
land hardwood forest. 


RANUNCULACEAE 

*Clematis terniflora DC., A1354. Infrequent. 
Fencerows, roadsides, and house sites. 

Clematis virginiana L., A1696. Common. Forest 
edges. 


riliic Qbortive 


sat 


AINIIN 


is L., A279. Infrequent. Moist 


ope dsides and yards. 

Ranunculus as Muhl. ex Bigelow, A268. In- 
frequent. Pine forest, clearcuts, and road- 
sides 


* Ranunculus parviflorus L.,A540. Infrequent. Old 
fields. 
Ranunculus pusillus Poir., A324. Common. Moist 


SS 
mw 
ond 


Ranunculus recurvatus Poir., A652. Infrequent. 
Sandy and gravelly areas, especially near 
bottomland. 

*Ranunculus sardous Crantz, A222. Abundant. 
Pastures, old fields, yards, and roadsides. 

RHAMNACEAE 

Berchemia scandens (Hill) K. Koch, AS58.Common. 
co of various sorts ie forest edges. 

5 L., A659. Infrequent. Pine 


st and associated wn ides. 
Se caroliniana (Walter) A.Gray [=Rhamnus 
caroliniana Walter], A609. Infrequent. Mature 


688 


hardwood forest of ravines and Saffell out- 
crops 


ROSACEAE 

Agrimonia microcarpa Wallr,A1555.Rare. Mature 
upland hardwood te . Saffell outcrops. 

gel a fea Wallr., A7389. Rare. Mature 

ardwood sales of Saffell outcrops. 

mee ae (F. Michx.) Fernald, A203. In- 
fre . Forest of various sorts. 

mage marshallii Eggl., EA 246.Common.For- 

arious sorts 
_ Heal les Ashe, WA&M 70982. Infre- 
pland, unburned longleaf pine for- 
ee . Homochitto River basin. 

Crataegus spathulata Michx., A547. Rare. Upland 
uote longleaf ne forest of the 
Homochitto River basi 

ee indica one Focke, A206. Infre- 

uent. Yards and ruderal a 

Geum canadense Jacq.,A 1057. i Si bottom- 
land forest of the Homochitto R 

Malus angustifolia (Aiton) Michx., ao Infre- 
quent. Upland forest, especially in the Homo- 
chitto River basin. 

Photinia pyrifolia (Lam,) KR. HOBETSON & J.B. 
Phipps [ tifolia (L.) Pers], A2024. 
fae Spring- cen and swamp 


¢ oe ann (Desf.) Kalkman, ee Rare. 
Upland and bottomland mixed for 
Potentilla simplex Michx., A503. Rare. a out- 


Tops. 

Pit angustifolia Marshall, EA 196. Common. 
Pine forest and dry roadsides. 

Prunus caroliniana (Mill) Aiton, FA 224. Infre- 
quent. Saffell outcrops, upland mature hard- 
wood forest, and bottomland hardwood 
forest 

Prunus mexicana S.Watson, A205. Common. For- 

es of various sorts and Saffell out 
crops. 

Prunus serotina Ehrh., A745. eae Forest of 

various sorts and fenc 

Prunus eo ata Elliott, Pe 
and bottomland forest edges 

ie calleryana Decne., A383. care Old oil well 


eee Upland 


oe ae J.C. Wendl.,A776. Rare. Roadside 
north of Liberty. 


BRIT.ORG/SIDA 19(3) 


Rosa carolina L., A1944. Common. Upland pine 
forest and roadsides 

*Rosa laevigata Michy,, 406. Common. 
Fencerows, roadsides, forest edges of 


various sorts. 
*Rosa multiflora Thunb.ex A. Murray, A543. Infre- 


u S 

*Rosa wichuraiana Crép., A797. Infrequent. Old 
pastures and fields, fencerows, and roadside 
ditches 

Rubus ort Link, A438. Abundant. Pine forest, 
forest edges, and clearcuts. 

Rubus pie Willd. [including R. enslenii 
Tratt.J], A2518. Infrequent. Pine forest and 
gravelly areas 

Rubus trivialis Michx.,A218. Abundant. Old fields, 
pastures, and open forest of various sorts. 

RUBIACEAE 


Ceph qlanthtiye 
a 


a 


“identalis L., A1098. Infrequent 
Pond margins, riverbanks, and swamp 
margins. 

Diodia teres Walter, A1282. Common. Clearcuts, 

ruderal areas, and pine forest. 
oP virginiana L., A719. Infrequent. Clearcuts 
and disturbed 

ie aparine . oe Infrequent. Ruderal 
areas and yar 

Galium circaezans viene Referenced in Jones 
(1976); specimen not se 

Galium obtusum ete ssp. eee A479. 
Rare. Bottomlands 

Galium orizabense ssp. laevicaule (Weath. & 
se py ee eleee, eles Saffell 
outcro 


& 


oadsides 


Galium es nee var n (Michx.) Torr. 
A. Gray, A1503. Rare. Pine forest 

Galium triflorum Michx., A848. Rare Mette hard- 
wood forest. 


Galium uniflorum Michx.,A1018. Rare. Pine forest. 
Houstonia micrantha nba Terrell, A287. In- 
requent. Yards, cemeteries, and fields. 


Houstonia purpurea L. var. purpurea, A613. Infre- 
quent. Forest drains. 

Houstonia pase sae A209. |nfrequent. 
Yards d fields. 


eteries 


a rosea (Raf.) el A294. Infrequent. 
Yards, cemeteries, and fields 


Mitchella repens jase Corammon bares of vari- 
sorts. 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


Oldenlandia boscii Mae Chapm., WA&M 71205. 
Infrequent San and gravelly opel Ings. 
Oldenlandia uni a : A2289. Infrequent. Bot- 

miand fields. 
Richardia ea L., A2353. Infrequent. Sandy 
river banks. 
* Sherardia cee L.,A656. Infrequent. Roadside 
ditches and yards. 


UTACEAE 
BONGO moat se ) na Al pase NS SLSHE 


Bide ed lene var. moll Torr. & A. Gray, A1201. 
Rare. Upland hardwood forest in the western 
part of the count 

Zanthoxylum clava-herculis L.,A1200.|\nfrequent 
Upland forest or sandy bottomland forest. 


SALICACEAE 

* Populus alba L.,A1176. Rare. Old home sites in 
the eastern half of the c 

Populus deltoides ae ex Sir ial Al225. 
Infrequent. Forest edges or bottomland 


openings. 
Salix nigra Marshall, A575.Common. Riverbanks 
pond margins, and moist roadsides. 


SAPOTACEAE 

Sideroxylon lycioides L. [=Bumelia lycioides (L.) 
Pers.],A2576. Rare. Upland hardwood forest 
in the Homochitto River basin. 


co 


SAURURACEAE 
Saururus cernuus L.,A737.Locally common. Road- 
side ditches open swamps spring-seeps, and 


besteanmpeundee ponds. 


SCHISANDRACEAE 

Schisandra glabra (Brickell) Rehder, A994. Infre- 
quent. Hardwood ravines and Saffell out- 
crops 


SCROPHULARIACEAE 
Agalinis fasciculata (Elliott) Raf,,A 1553. Abundant. 
Old fields. 
aul is plukenetii (Elliott) Raf,,A7764.Infrequent. 
oes forest of the Homochitto River 


~ 
eae ; nih (Vahl) Raf., A2254. Rare. Rocky 
bluffs overlooking the East Fork Amite River. 
+Aureolaria dispersa (Small) Pennell [=A. virginica 
(L.) Pennell, in part], A7632. Infrequent. Pine 
forest in the southeastern part of the county. 
Species laxOnOoMy follows Pennell (1935). 


689 


Aureolaria pectinata (Nutt.) Pennell, EA 96. Infre- 
quent. Pine forest. 

Buchnera americana L. ae B. floridana 
Gandoger], A588. Common. Fields. 

Gratiola floridana Nutt., ee Locally common. 

pring-seeps and swamp margins. 

aa. neglecta Torr., A517. Rare. Bottomland 
roadside in the southeastern part of the 
county. 

Gratiola pilosa Michx., WA&M 71195. Infrequent. 

Pine forest and roadsides 

Gratiola virginiana L., A532. Eien Bottom- 
land forest and river m 

*lindernia crustacea (L.) eae A1355. Infre- 


Lindernia dubia (L.) Pennell, A1045. Common. 
Bottomland openings and river margins. 

*Mazus pumilus (Burm.f) Steenis, EA 227. Infre- 
quent. Bottomland fields and yards. 

Mecardonia acuminata (Walter) Small, A1502. 
Infrequent. Pine forest 

Micranthemum umbrosum (Walter ex J.F.Gmel.) 
S.F. Blake, A763. Locally common. Pond mar- 
gins, wet ditches, swamps, and bottomland 
hardwood forest. 

Mimulus alatus Aiton, A1330. Swamp margins, 
river margins, and edges of bottomland 


——. 


hardwood forest. 
Nuttallanthus canadensis (L.) D.A. Sutton 
nes canadensis (L.) Chaz.], A269. Com- 
n. Pastures, roadsides, and yards. 
ea aris canadensis L.,A643. Rare. Upland for- 
mochitto River basin. 
Penstemon renee Nutt. ex Sims, A843. Infre- 
quent. Pine forest and roadsides 
Penstemon laxiflorus Pennell, A1883. Infrequent 
side 
Scoparia dulcis L.,A2290. Rare. Gravelly clearcuts. 
Seymeria cassioides (Walter ex J.F. Gmel.) S-F. 
Blake, A1743. Common. Pine forest. 
*Verbascum thapsus L.,A1132. Infrequent. Saffell 
outoRs ee areas, and roadsides. 
L.,A286.Common. Roadsides 


and yards. 
Veronica peregrina L.,A1882.Common.Roadsides 
nd clearcuts. 
*Veronica persica Poir., A295. Common. Road- 
sides 


SIMAROUBACEAE 
* Ajlanthus altissima (Mill. 


7 


Swingle, A2109. Infre- 


quent. Ditches and roadsides in the western 
part of the county. 


SOLANACEAE 

Physalis angulata L.,A1237.Common. Cultivated 
fields. 

Physalis carpenteri Riddell, A10517. 
Saffell outcrops. 

Physalis heterophylla Nees, A700. Infrequent. 
Saffell outcrops and sandy bottomland. 
Physalis pubescens L.,A1240.Common.Clearcuts 

and roadsides 


Infrequent. 


Solanum elaeagnifolium Cav., M. Whitson 1142 
DUKE!). Rare. Gravelly upland in the 
Homochitto River basin 

Solanum carolinense L., A666. Common. Road- 
sides, cultivated fields, and pine forest. 

* Solanum pseudocapsicum L.,A2346. Infrequent. 
Sandy bottomland forest along the forks of 
the Amite Riv 

Solanum ptycha oe Dunal, A720. Common. 
Clearcuts, open fields, and sandy roadsides. 

* Solanum viarum Dunal, A1025. Rare. Cattle 
barns. Noxious weed; all individuals ob- 
served were collected or destroyed. 

STERCULIACEAE 

Melochia corchorifolia L.,A1795.Rare. Riverbanks 
and sandbars of the Amite River basin. 


STYRACACEAE 
Halesia oo. bundant.Bottomland 
est, swamp margins, ravines, and Saffell 


ve groom Aiton, AT100. Infrequent. oa 
r fell 


avines.andS 


outcrops. 


SYMPLOCACEAE 
sel ai eae (L.) UHér., A402. Common. 
and hardwood forest, upland hard- 

eed or sli forest, and ravines. 

THEACEAE 

Stewartia ect ane L., A559. Infrequent. 
Bottomland hardwood forest, hardwood ra- 
vines, bees Saffell outcrops. 

TILIACEAE 

Tilia americana var. caroliniana (Mill) Castigl., 
A2075. Infrequent. Hardwood forest of Saffell 
outcrops 


BRIT.ORG/SIDA 19(3) 


ULMACEAE 
~ seieecs Willd., A7061. Infrequent. Sandy 
mland of the Homochitto River basin. 

re alata Michx., A935. Abundant. Upland 
hardwood forest, pine forest, and bottom- 
land hardwood forest. 

Ulmus americana L., FA 200. Infrequent. River mar- 
gins and bottomland forest. 

Ulmus rubra Muhl., A870. Rare. Bottomland hard- 
wood forest. 


UMBELLIFERAE 

Chaerophyllum aan Hook,, A332.Common, 
Roadsides and fie 

Ciclospermum heii (Pers.) Sprague ex 
Britton & Wilson [=Apium leptophylium (Pers.) 
Benth.], A636. Infrequent. Roadsides, upland 
forest over loose soil, and Saffell outcrops. 

Cicuta maculata L., A807. Infrequent. Moist road- 
sides and open bottomland 

* Daucus carota L.,FA 262. Infrequent. Roadsides 

ee ree Nt ex re as Abun- 

st fields and open 

. tto i nd. 

Eryngium yuccifolium Michx., WA&M 71053. Infre- 
quent. Upland pine forest and sandy river 

argin 


Bivdirocatyie ranunculoides L.f., Al837. Rare. 
Muddy swamp edge in the West Fork Amite 
eran age. 

Hydrocotyle umbellata L.,A2059. Common. Pond 
margins, swamp margins, sloughs, and wet 

‘elen es. 

ydrocotyle verticillata Thunb,, ee Infrequent. 
Balam hardwood fore 

—— capillaceum (Michx.) ieee. 


soni canadensis L.var.canadensis, A859.Com- 
— forest and dry, upland hard- 


co forest. 
Sanicula al, E.P. Bicknell, A615. Infrequent. 
Mesic hardwood ravines and Saffell out- 


rops. 
Tho ium um ifol aa JA. oe Infrequent. 
hardwood 


ravines, 

URTICACEAE 

Boehmeria cylindrica (L.) Swartz,A1350.Common. 
Bottomland forest or other mesic forest. 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 


Laportea canadensis (L.) Wedd., A2112. Infre- 
quent. Hardwood forest of the Homochitto 
River basin 

Pilea pumila (L.) A.Gray,A1823.Rare. Abandoned 

xbow swales 

Urtica chamaedryoides Pursh,A2118. Infrequent. 
Hardwood forest on thick loess in the Homo- 
chitto River basin and disturbed forest in the 
northeastern corner of the county. 

VALERIANACEAE 

Valerianella radiata (L.) Dufr., A272. Common. 
Roadsides, yards, and fields. 

VERBENACEAE 

Callicarpa americana L., A1029. Abundant. Pine 
— ee ie and Saffell outcrops. 

m (L.) Kuntze, A2282. Rare 


oadsides. 
Pe pulchella (Sweet) Tronc. Referenced 
in Pullen et al. (1968); specimen not seen. 
*/ antana camara L.,A2306. Rare. Roadsides 
Phryma leptostachya L., A1143. oe Rich 


hardwood forest at Saffell outcr 
*V/erbena brasiliensis Vell, A724. jaeeee Road- 
sides, upland forest edges, and old fields. 


Verbena haleisinall Aa7o Infrequent. Roadsides 
* Verbena rigida Spreng., A597. Locally common. 
adsides. 

* Vitex agnus-castus L., A996. Rare. Ruderal lot in 
Liberty. 

VIOLACEAE 

Viola affinis Leconte, A247. Common. Ravines, 
upland hardwood and mixed forest, and 
Saffell outcrops. 

Viola bicolor Pursh [=V. rafinesquii Greene], A328. 
Infrequent. Yards and old fields. 


691 


Viola missouriensis Greene (sensu lato, including 
V. floridana Brainerd and possibly V.langloisii 
Greene;see Gilad 1995),A291.Infrequent.Old 
fields and yards 

Viola pedata L., A334. Locally common. Pine for- 
est and roadsides in the eastern half of the 


unty. 
ioe aie L,A217. Abundant. Moist open- 
ings, streambanks, bottomland fields, and 
roadside ditches 
Viola triloba Schwein., A708. Common. Forest of 
various sorts and Saffell outcrops 
Viola walteri House, A260. Common. Upland 
hardwood forest, ravines, and Saffell out- 
crops. 


VISCACEAE 

Phoradendron leucarpum (Raf.) Reveal & M.C. 
Johnst., 41534. Common. Parasitic and epi- 
phytic upon various hardwoods, primarily 
Prunus serotina and Quercus spp. 


VITACEAE 
Ampelopsis arborea (L.) Koehne,A1305.Common. 
Open forest or forest edges of various sorts. 
Parthenocissus quinquefolia (L.) Planch., A1155. 
nt. Pine forest, upland hardwood or 
mixed forest, bottomland hardwood forest, 
ps, yards, and towns. 


Saffell outcrops, swam 
Vitis aestivalis Michx. var. aestivalis, A2081.Abun- 
orest and forest edges of various sorts. 
Vitis cinerea (Engelm.) Engelm.ex Millardet var. ci- 
rea, A888. Commen, Forest and forest edges 
ariOus S : 
Vitis ai ee Abundant. Forest 
and forest edges of various sorts, 


DISCUSSION 


The survey for vascular plants in Amite County, Mississippi yielded 923 spe- 
cies, which is about 31% of the total number of species found in Mississippi 
(Kartesz 1999). Table 3 provides a summary of the taxa found in the county. The 
largest plant families are Compositae (116 spp.), Gramineae (99 spp.), Cyperaceae 
(74 spp.), and Leguminosae (63 spp.), and the three largest genera are Carex G4 
spp.), Panicum s.l. (19 spp.), and Quercus (18 spp. + | common hybrid). Intro- 
duced species make up about 16% of the flora, which is a proportion similar to 
neighboring areas (e.g., Clewell 1985: 16%). 

The survey re-confirmed the existence of several rare species in the county 


692 BRIT.ORG/SIDA 19(3) 


Taste 3. Synopsis of vascular plant taxa recorded for Amite County, Mississippi 


Species Genera Families 

Lycopodiophyta | | 1 
Polypodiophyta 26 20 11 
Coniferophyta 7 3 2 
Magnoliophyta 889 442 139 

(Magnoliopsida 634 343 109) 

(Liliopsida 255 99 30) 
TOTAL 923 466 153 
Indigenous 777 389 142 
Introduced 146 (16%) 77 11 


Largest families: Compositae (116 spp.), Gramineae (99 spp.), Cyperaceae (74 spp.), Leguminosae 
(63 spp. 
Largest genera: Carex (34 spp.), Panicum s.|.(19 spp.), and Quercus (18 spp.) 


and added a few new records to the state. Rarity is measured in accordance with 
the Mississippi Natural Heritage Program (1994), where “G” and a number in- 
dicate worldwide status and “S” and a number indicate state status. The rarest 
species are given a number I, and the more secure given 5. Rare species indi- 
cated by the Mississippi Natural Heritage Program (1995) to occur in Amite 
County that were again encountered in the present survey include Antennaria 
solitaria (G5/S3?), Chromolaena ivifolium (G5/S2?), Epidendrum conopseum 
(G3G4/52), Luzula acuminata (G5/S3), Mikania cordifolia (G5/S354), Pachys- 
andra procumbens (G4G5/S3), Schisandra glabra (G4/S3?), Stewartia malaco- 
dendron (G4/S354), Trichomanes petersii (G3/S1), and Trillium foetidissimum 
(G3G4/S3). Stewartia malacodendron and Trillium foetidissimum, although 
listed as state rare species, are actually not uncommon in the county. 

In addition to previous records, the present survey recorded the existence 
of Carex decomposita (G3G4/S3?), Dryopteris ludoviciana (G4/S1), Iris brevi- 
caulis (G4/S?), Lobelia appendiculata (G4G5/S2S3), Matelea carolinensis (G4/ 
S253), Melanthium virginicum (G5/82S3), Sabatia campestris (G5?/S2S3), and 
Spiranthes ovalis (G5/S2S3). 

New records for the state are Alstroemeria psitticina (Alstroemeriaceae), 
Clinopodium gracile (Labiatae), Ipomoea indica (Convolvulaceae), Photinia 
serratifolia (Rosaceae), and Solidago auriculata (Compositae). Alstroemeria 
psitticina is an introduced species from Brazil (Bailey 1949). One small popula- 
tion was discovered within the city limits of Liberty under a tree with much 
leaf mulch. Neither the property owner nor his neighbors recognized the spe- 
cies,and none claimed to have cultivated such a species in the past. Clinopodium 
gracile is an introduced species from Japan (Burkhalter 1984). It was first noted 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 693 


to occur in the United States in Louisiana in 1963 (Thieret 1964) and has subse- 
quently been collected in Florida (Burkhalter 1984) and reported for Alabama 
J.V.Ward, pers. comm. to Kartesz 1999). C. gracile was actually discovered in 
the United States as early as 1934 but apparently was never reported (Roland 
Harper, s.n,, lberia Parish, LA, 16 July 1934, BH1). It has also been known to occur 
in Mississippi John R. MacDonald 9771, Copiah Co., MS, 19 July 1996, MO!) but 
has not been previously reported. Ipomoea indica is a pantropical weed and was 
found in several roadside ditches near Liberty. Photinia serratifolia isa shrubby 
species native to China (Bailey 1949) and has been frequently cultivated in the 
southern United States. Although I found no clear evidence of self-established 
lines, the species was found in areas with no evidence suggesting former culti- 
vation either. Solidago auriculata is the only new record of a species native to 
the southeastern United States. It has a large distribution but seldom occurs 
with frequency. 

Also collected were Physalis carpenteri and Dryopteris ludoviciana, both 
of which have only recently been reported to occur in Mississippi (L.M. McCook, 
pers. comm. to Kartesz 1999, Sorrie & Leonard 1999, respectively). At least one 
specimen of Physalis carpenteri had been collected in Mississippi before (War- 
ren County, MISS!), was misidentified, and was later annotated correctly by Ja- 
net Sullivan, then working on her part of the Flora of the Southeastern United 
States. Unfortunately, that treatment was not subsequently published. Consid- 
ering the limited range of Physalis carpenteri to Florida, Louisiana, and Missis- 
sippi, it will likely be added to the Mississippi Natural Heritage Program list of 
rare species. 

The flora also yielded a number of champion trees and shrubs. Especially 
large trees and shrubs were noted and contributed to the Mississippi Forestry 
Commission champion tree program. Amite County was already known to be 
home to the largest individuals of Frangula (Rhamnus) caroliniana, Hamame- 
lis virginiana, Liriodendron tulipifera, Magnolia acuminata, Nyssa sylvatica, 
Pyrus communis, and Triadica ae sebifera in Mississippi and is now 
known to be home to the largest Halesia diptera, Ilex verticillata, Ilex vomitoria, 
Kalmia latifolia, and Morella cerifera. 

The flora of Amite County reveals the complexity of plant distributions in 
the southeastern United States. The flora is rich in species and vegetation types, 
with prominent variations resulting from physiographic, topegiapme and 
edaphic factors. The eastern half of the county shows floristic similarity to other 
upland regions of the lower coastal plain. Pines are the dominant woody plants, 
surrounded by a herbaceous vegetation rich in composites, legumes, grasses, 
and sedges. Surprisingly, there is more similarity between the highlands of the 
Homochitto River basin and the southeastern corner of the county than with 
the area in between (see Fig. 2). Perhaps this is a result of the erosion of loess 
from Homochitto ridges and the exposure of the underlying Miocene clays. In 


694 BRIT.ORG/SIDA 19(3) 


these Homochitto areas, many coastal plain species are found which are not 
even found in the southeastern corner of the county, for example, Drosera 
brevifolia, lonactis linariifolius, Liatris elegans, and Symphyotrichum adnatus. 

With the introduction of loess to the substrate and the large Mississippi 
Embayment just to the west, Amite County naturally serves as the westernmost 
distribution point for several coastal plain species, such as Gelsemium rankinii 
and Illicium floridanum. This is almost true for Packera anonyma, Gaylussacia 
dumosa, and Kalmia latifolia as well, but there are rare reports of their occur- 
rence west of the Mississippi River (MacRoberts 1989). The introduction of loess 
also serves to harbor the residual flora of Pleistocene migrations from the north 
(Delcourt & Delcourt 1975), and thus, Amite County is home to several species 
indicative of the mesophytic forests of mid-eastern North America, including 
Adiantum pedatum, Cynoglossum virginianum, Hydrangea arborescens, and 
Pachysandra procumbens. There is also a weaker botanical association with the 
West (e.g., Cuscuta cuspidata, Gaura brachycarpa, Liatris pycnostachya, Sabatia 
campestris, and Vernonia texana) and with the neotropics (e.g., Chromolaena 
ivifolia and Mikania cordifolia). Other phytogeographical conclusions have 
been discussed in greater detail in Allen et al. 1975). 


ACKNOWLEDGMENTS 


Professor Robert L. Wilbur, who encouraged me to complete this master’s project 
and challenged me at various stages of its completion, is heartily thanked. He 
is also responsible for the treatment of Hydrocotyle (Apiaceae) in the flora and 
caught several errors of identification and lapses of taxonomic judgment before 
this publication. My parents, Earl and Pauline Alford, also provided immense 
support for this project by making collections of the flora, mailing specimens, 
asking around for certain kinds of collection sites, and somehow both accept- 
ing and persisting through my need to collect and travel the county while at 
home, even during short holidays. Several people are responsible for treatments 
in this flora, and they deserve immense thanks, especially since these taxa are 
some of the most perplexing and difficult groups in the Southeast. Charles T. 
Bryson (USDA Southern Weed Science Research Unit, Stoneville, Mississippi), 
Jay W. Horn (DUKE), Gerry Moore (BKL), and Guy Nesom (NCU) identified the 
Carex (Cyperaceae), Xyris (Xyridaceae), Rhynchospora (Cyperaceae) and Pani- 
cum subgenus Dichanthelium (Gramineae), and Aster and Boltonia 
(Compositae), respectively. All errors of taxonomic judgment or identification, 
however, | accept as my own. Charles M. Allen (NLU) supplied spreadsheets of 
plant collections in the neighboring Louisiana parishes and most importantly 
provided excellent background work and a model to the present survey through 
his own thesis of a similar type. lalso thank Kenneth L. Gordon (USFS), the late 
John Allen Smith, Catherine Keever, John MacDonald CUBE), Lucile McCook 
(MISS), Steven R. Hill GLLS), Carl E. Lewis (BH), Maggie Whitson (DUKE), Fred 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 695 


and David Anderson, Robert and Bettye Causey, Mike Garrett, the late Jewell 
Lambert, John William Powell, Charlotte Reynolds, Sam and Gayle Tumey, 
Reginald Richardson, Harry Wells, and the curators of BH, GA, IBE/MISSA, LSU, 
MICH, and MISS. The Homochitto National Forest and the Mississippi Depart- 
ment of Wildlife, Fisheries, @ Parks are both thanked for permits to collect 
plants. A combination of Duke University’s Catherine Keever and Lawrence 
Giles Awards provided necessary money for herbarium visits, and extra funds 
from the James B. Duke Fellowship and the Harvey Fellows Program of the Mus- 
tard Seed Foundation allowed me numerous collecting opportunities. 


REFERENCES 


Atrorb, M.H. 1998. New records of Allocapnia (Plecoptera: Capniidae) from Mississippi and 
Louisiana, with accompanying scanning electron micrographs. Entomol. News 109: 
183-188. 

Auten, C.M. 1972. A flora of the vascular plants of St. Helena Parish, Louisiana. M.S. Thesis, 
Louisiana State University. 

Atten, CM. 1992. Grasses of Louisiana, 2nd ed. Cajun Prairie Habitat Preservation Society, 
Eunice, LA. 

Atten, C.M.,M.G. Curry, and B.F. Martin. 1975.A vascular flora of St. Helena and West Feliciana 
Parishes, Louisiana. Univ. of Southw. Louisiana Res. Ser. No. 39. 

BacktUND, M., B. Oxetman, and B. Bremer. 2000. Phylogenetic relationships within the 
Gentianales based on ndhF and rbcLl sequences, with particular reference to the 
Loganiaceae. Amer. J. Bot. 87:1029-1043. 

Baitey, LH. 1949, Manual of cultivated plants. MacMillan, New York. 

Bicker, A.R., Jr, compiler. 1969. Geologic map of Mississippi. 

BLAISDELL, R.S., J. Wooten, and R.K. Goorrey. 1974. The role of magnolia and beech in forest 
processes in the Tallahassee, Florida, Thomasville, Georgia area. Proc. Annual Tall Tim- 
bers Fire Ecol. Conf. 13:363-397. 

Bourpo, E.A., Jr. 1956. A review of the general land office survey and of its use in quantita- 
tive studies of former forests. Ecology 37:/54—/68. 

Braun, E.L. 1950. Deciduous forest of eastern North America. Free Press, New York. 

Brummitt, R.K. and C.E. Powe, eds. 1992. Authors of plant names. Royal Bot. Gardens, Kew. 

Bryson, C.T. and R. Carter. 1992. Notes on Cyperus and Kyllinga (Cyperaceae) in Mississippi 
with records of six species new to the state. Sida 15:119-124. 

Bryson, C.T., J.R. MacDonato, R. Carter, and S.D. Jones. 1996. Noteworthy Carex, Cyperus, 
Eleocharis, Kyllinga, and Oxycaryum (Cyperaceae) from Alabama, Arkansas, Georgia, 
Louisiana, Mississippi, North Carolina, Tennessee, and Texas. Sida 17:501-518. 

BurkHatter, J.R. 1984. Additions to the vascular flora of Florida. Castanea 49:180-186. 

Campselt, C.S. 1983. Systematics of the Andropogon virginicus complex (Gramineae). J. Ar- 
nold Arbor.64:171—254. 

Captenor, D. 1968.Forest composition on loessal and non-loessal soils in west-central Mis- 
sissippi. Ecology 49:322-331. 


696 BRIT.ORG/SIDA 19(3) 


Casey, A.E. 1957. Amite County, Mississippi, 1699-1890. Vol. Ill. The environs. Privately pub- 
lished by the Amite County Historical Fund, Birmingham 

Casey, A.E. and F.P. Orken. 1948. Amite County, Mississippi, 1699-1865. Vol. |. Birmingham: 
Privately published by the Amite County Historical Fund. 

Casey, A.E., FP. Orken, M.T. Loch, and C.G. Nortr. 1950. Amite County, Mississippi, 1699-1865. 
Vol.Il. The churches. Privately published by the Amite County Historical Fund, Birmingham 

CHAPMAN, A.W. 1897. Flora of the southern United States. 3rd ed. Cambridge Botanical Supply 
Co., Cambridge, MA. 

CHRISTENSEN, N.L. 2000. Vegetation of the southeastern coastal plain. In: M.G. Barbour and 
W.D.Billings,eds. North American terrestrial vegetation, 2nd ed. Cambridge Univ.Press, 
Cambridge, U.K. Pp. 397-448. 

Ciark, E. 1997. Mississippi official and statistical register 1996-2000. Jackson, MS. 

Crewelt, A.F. 1985.Guide to the vascular plants of the Florida panhandle. Tallahassee: Florida 
State UP. 

Corrett, D.S. and M.C. Jounston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner. 


Cronauist, A. 1980. Asteraceae. Vascular flora of the southeastern United States. Vol.1. Univ. 
of North Carolina Press, Chapel Hill. 

DaHiGREN, R.M.T., H.T. Cuirrorp, and P.F. Yeo. 1985. The families of the monocotyledons: struc- 
ture, evolution, and taxonomy. Springer-Verlag, Berlin. 

Darsy, W. 1817.A geographical description of the state of Louisiana, the southern part of 
the state of Mississippi, and territory of Alabama .... James Olmstead, New York. 

Darsy, W. and T. DwicHt, Jr. 1836.A new gazetteer of the United States of America ....Rev. 
ed. Edward Hopkins, Hartford. 

Devcourt, H.R. and PA. Deicourt. 1974. Primeval magnolia-holly-beech climax in Louisiana. 
Ecology 55:638-644. 

Detcourt, H.R. and PA. Detcourt. 1975. The blufflands: Pleistocene pathway in the Tunica 
Hills. Amer. Mid. Naturalist 94:385—400. 

Devcourt, H.R. and PA. Detcourt. 1977. Presettlement magnolia-beech climax of the Gulf 
coastal plain: quantitative evidence from the Apalachicola River Bluffs, north central 
Florida. Ecology 58:1085-1093. 

Devcourt, H.R.and PA. De_courr. 2000. Eastern deciduous forests. In: M.G. Barbour and W.D, 
Billings, eds. North American terrestrial vegetation, 2nd ed. Cambridge Univ. Press, Cam- 
bridge, U.K. Pp. 357-395. 

Doerine, J.A. 1935. Post-Fleming surface formations of coastal southeast Texas and south 
Louisiana. Bull. Amer. Assoc. Petrol. Geol. 19:65 1-688. 

Doerine, J.A.. 1956. Review of Quaternary surface formations of Gulf Coast region. Bull. 
Amer. Assoc. Petrol. Geol. 40:1816-1862. 

DonocHue, MJ. 1983. The phylogenetic relationships of Viburnum. In: N.I. Platnick and V.A. 
Funk, eds. Advances in cladistics, vol. 2. Columbia Univ. Press, New York. Pp. 143-166. 
DonocHue, M.J., R.G. Osten, J.F. Smith, and J.D. Patmer. 1992. Phylogenetic relationships of 

Dipsacales based on rbcl sequences. Ann. Missouri Bot. Gard. 79:333-345. 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 697 


Duncan, W.H. 1953. Taxonomic collections of vascular plants in the southeastern United 
States—their abundance and relation to production of floras. Rhodora 55:353-358. 

Evans, A.M. 1978. Mississippi flora: a guide to the ferns and fern allies. Sida 7:282-297. 

Fenneman, N.M. 1938. Physiography of eastern United States. McGraw-Hill, New York. 

FLORA oF NortH America EprroriaL Committee, eds. 1997.Flora of North America north of Mexico. 
Volume Ill. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford Univ. Press, New 
York. 

GANDHI, K.N. and R.D. THomas. 1989. Asteraceae of Louisiana. Sida, Bot. Misc. 4:1-202. 

Git-ap, N.L. 1995. Systematics of Viola subsection Boreali-Americanae. Boissiera 53:5—130. 

Gitus, N.E. 1963. Early inhabitants of the Natchez District. Gillis, Baton Rouge, LA. 

GLeason, H.A. 1931.Book review: Johnson's taxonomy of flowering plants. Torreya 31:77-84. 

Goorrey, R.K. 1988. Trees, shrubs, and woody vines of northern Florida and adjacent Georgia 
and Alabama. Univ. of Georgia Press, Athens. 

Goorrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United 
States. Monocotyledons. Univ. of Georgia Press, Athens. 

Goorrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of southeastern United 
States. Dicotyledons. Univ. of Georgia Press, Athens. 

Harper, F.,ed. 1958. The travels of William Bartram: naturalist’s edition.Yale Univ. Press, New 
Haven, CT. 

Haskins, M.L. and WJ. Hayoen. 1987. Anatomy and affinities of Penthorum. Amer. J. Bot. 74: 
164-177 

Hopckins, E.J. 1958. Effects of fire on undergrowth vegetation in upland southern pine 
forests. Ecology 58:36-46. 

Houmes, J.S.and J.H. Foster. 1908. A study of forest conditions of southwestern Mississippi. 
Mississippi State Geol. Surv. Bull. No. 5. 

Houmeren, PK., N.H. Houmoren, and L.C. Barnertt, eds. 1990. Index herbariorum Part 1: the 
herbaria of the world, 8th ed. New York Botanical Garden, Bronx. 

JOHNSON, G.P. 1988. Revision of Castanea sect. Balanocastanon (Fagaceae).J. Arnold Arbor. 
69:25-49, 

Jones, S.B., Jr. 1974a. Mississippi flora. |. Monocotyledon families with aquatic or wetland 
species. Gulf Res. Rep. 4:357-379. 

Jones, S.B., Jr. 1974b. Mississippi flora. Il. Distribution and identification of the Onagraceae. 
Castanea 39:370-379. 

Jones, S.B,, Jr. 1975. Mississippi flora. Ill. Distribution and identification of the Brassicaceae. 
Castanea 40:238-252. 

Jones, 5.B., Jr. 1976. Mississippi flora. VI. Miscellaneous families. Castanea 41:189-21 2. 

Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vas- 
cular flora of the United States, Canada, and Greenland. 1st ed. in Kartesz, J.T.,and CA. 
Meacham. Synthesis of the North American flora, Version 1.0.North Carolina Botanical 
Garden, Chapel Hill. 

Krinitzsky, E.L. and W.J. Turnsutt. 1967. Loess deposits of Mississippi. Special Pap. Geol. Soc. 
Amer. 94. 


698 BRIT.ORG/SIDA 19(3) 


Kurz, H. 1944. Secondary forest succession in the Tallahassee Red Hills. Proc. Florida Acad. 
Sci. 7:59-100. 

Leccert, A.R., A.C. Miceranor, L.B. Hate, and PJ. Bartow. 1968. Soil survey of Walthall County, 
Mississippi. USDA, Washington, D.C. 

LeLonc, M.G. 1986. A taxonomic treatment of the genus Panicum (Poaceae) in Mississippi. 
Phytologia 61:251-269. 

LINNAEUS, C. 1758. Opera varia. Lucca. 

Lowe, E.N. 1913. Forest conditions of Mississippi. Mississippi State Geol. Surv, Bull. No. 11 (a 
reprint with additions of Bulletins Nos.5 and 7). 

Lowe, E.N. 1921. Plants of Mississippi:A list of flowering plants and ferns. Mississippi State 
Geological Survey Bulletin No. 17. Hederman Brothers, Jackson, MS 

MacRoserts, D.T.1989.A documented checklist and atlas of the vascular flora of Louisiana. 
Bull. Mus. Life Sci. Louisiana State Univ. 7-9. 

McAree,W.L. 1956.A review of the Nearctic Viburnum. Published by the author, Chapel Hill, NC. 

McCook, L.M. 1982. The vascular flora of the Clark Creek Natural Area, Wilkinson County, 
Mississippi. M.S. Thesis, University of New Orleans. 

Miveranor, A.C. 1976. Soil survey of Amite County, Mississippi. USDA, Washington, D.C. 

Miveranor, A.C. and L.B. Hate. 1968. Soil survey of Pike County, Mississippi. USDA, Washing- 
ton, D.C. 

Mississipp! FORESTRY COMMISSION. 1998. Facts at a glance:July 1997—June 1998.MFC Publication #3. 

Mississipp| NATURAL HeriTAGE PRocrAM. 1994, Special plant list. Museum of Natural Science, 
Mississippi Department of Wildlife, Fisheries, & Parks, Jackson. 

Mississipp| NaTuRAL HeriTace Procram. 1995. Special plants & animals: by county. Museum of 
Natural Science, Mississippi Department of Wildlife, Fisheries, & Parks, Jackson. 

Monk, C.D. 1965. Southern mixed hardwood forest of north central Florida. Ecol. Monogr 
35:335-354. 

Monk, C.D. 1968. Successional and environmental relationships of the forest vegetation 
of north central Florida. Amer. Midl. Naturalist 79:441-457, 

NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION (NOAA). 1962-1997. Annual climatological 
summary, Liberty 5 W station (22075070). National Climatic Data Center, Asheville, NC. 

Nesom, G.L. and M. Treiber. 1977. Beech-mixed hardwoods communities: A topo-edaphic 
climax on the North Carolina coastal plain. Castanea 42:119-140. 

Pattison, W.D. 1957. Beginnings of the American rectangular land survey system, 1784— 
1800. Univ. of Chicago Press, Chicago, IL. 

Pennett, FW. 1935. The Scrophulariaceae of eastern temperate North America. Acad. Nat. 
Sci. Philadelphia Monogr. 1:1-650. 

Pessin, LJ. 1933. Forest associations on the uplands of the lower Gulf coastal plain (longleaf 
pine belt). Ecology 14:1-13. 

Putten, T.M. 1966. A preliminary check-list of the Orchidaceae of Mississippi. Castanea 
31:153-154. 

Putten, T.M.,S.B. Jones, Jr, and J.R. Watson, Jr. 1968. Additions to the flora of Mississippi. Cas- 
tanea 33:326-334. 


ALFORD, VASCULAR FLORA OF AMITE COUNTY, MISSISSIPPI 699 


Quarterman, E. and C. Keever. 1962. Southern mixed hardwood forest: climax in the south- 
eastern coastal plain, U.S.A. Ecol. Monogr. 32:167-185. 

Raororo, A.E., H.E. AHLes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
Univ. of North Carolina Press, Chapel Hill. 

SHinners, L.H. 1962. Annual Sisyrinchiums (Iridaceae) in the United States. Sida 1:32-42. 

SmaALt, J.K. 1933, Manual of the southeastern flora. Science Press Printing Co., New York 

Soutis, D.E.and PS. Soutis. 1997.Phylogenetic relationships in Saxifragaceae sensu lato:a com- 
parison of topologies based on 18S rDNA and rbcL sequences. Amer. J. Bot.84:504-522. 

Sorrie, B.A.and S.W. Leonaro. 1999. Noteworthy records of Mississippi vascular plants. Sida 
18:889-908. 

SPEARING, D. 1995. Roadside geology of Louisiana. Mountain Press, Missoula, MT. 

Stern, E. 1976. The freshwater mussels (Unionidae) of the Lake Maurepas—Pontchartrain— 
Borgne drainage system, Louisiana and Mississippi. Ph.D. dissertation, Louisiana State 
University. 

STEYERMARK, J.A. 1963. Flora of Missouri. lowa State Univ. Press, Ames. 

Stone, D. 1997. Carya.|n: Flora of North America Editorial Committee, eds. Flora of North 
America north of Mexico. Volume III. Magnoliophyta: Magnoliidae and Hamamelidae. 
Oxford Univ. Press, New York. Pp. 417-425 

STRONG, M. and R. Karat. 1999. (1413) Proposal to conserve the name Scirpus miliaceus 
(Cyperaceae) with a conserved type. Taxon 48:387-389. 

Struwe, L., V.A. Atpert, and B. Bremer. 1994. Cladistics and family level classification of the 
Gentianales. Cladistics 10:1 75-206. 

Temete, L.C. and T.M. PuLLen. 1968. A preliminary checklist of the Compositae of Mississippi. 
Castanea 32:106-115. 

Tuieret, J.W. 1964. Lysimachia japonica (Primulaceae) and Clinopodium gracile (Labiatae) in 
Louisiana: new to the United States. Sida 1:249-250. 

THwaites, R.G. 1904. Early western travels 1748-1846. Vol. IV: Cuming's tour to the western 
country (1807-1809). A.H. Clark Company, Cleveland, OH. 

TreLease, W. 1902. The Yucceae. Thirteenth Annual Report of the Missouri Botanical Gar- 
den, St. Louis. 

Uniteo States Census Bureau. 1820. Amite County, Mississippi, 1820 in United States histori- 
cal census data browser [computer file, 1992]. Univ. of Virginia Press, Charlottesville. 
Uniteo Stares Census Bureau. 1840. Amite County, Mississippi, 1840 in United States histori- 
cal census data browser [computer file, 1992]. Univ. of Virginia Press, Charlottesville. 
Uniteo States Forest Service. 1995. Ecological units of the eastern United States: first ap- 

proximation. USFS, Atlanta. 

Weak.ey, A.S. [in prep.] Flora of the Carolinas and Virginia. The Nature Conservancy. 

WeLts, B.W. 1942. Ecological problems of the southeastern United States coastal plain. Bot. 
Rev. (Lancaster) 8:533-561. 

ZuLoaca, FJ. Duscovsky,and O. Morrone. 1993. Infrageneric phenetic relations in New World 
Panicum (Poaceae: Panicoideae: Paniceae): a numerical analysis. Canad. J. Bot. 71: 
1312-1327. 


BRIT.ORG/SIDA 19(3) 


Book REVIEW 


RICHARD P. WUNDERLIN and Bruce FE Hansen. 2000. Flora of Florida, Volume I. 
Pteridophytes and Gymnosperms. (ISBN 0-8130-1805-6, hbk.). University 
Press of Florida, 15 NW 15" Street, Gainesville, FL 32611-2079, US.A. 
(Orders: wwwupf.com). $49.95, 384 pp, 8 b&w photos, 68 drawings, bib- 
liography, index, 7" x 10" 

“Flora of Florida, Volume I, is the first of a proposed eight-volume comprehensive reference to the 

more than 3,800 vascular plants, native and non-native, known to occur growing wild in the state.” 

This provides standard botanical treatments for all families, genera, and species of the ferns and 

gymnosperms, including keys, nae nomenclature, illustrations, and summary information 


on habitat ee graphic distributi atroductory chapters, good for the whole Flora of Florida 


series, pro views of the ect. setting, vegetation, and botanical exploration of the state. A 
number of excellent biographical capsules form part of the ‘biographical exploration’ chapter. At 
the ete a iterature Cited, General Index, Index toCommon Names, and Index to Scientific Names. 
ously unusual that two separate volumes dealing primarily with the ferns and fern 

anit of F loi appear the same > yea, but the treatments are largely complementary and the au- 
as Gil Nelson provided some of the dust jacket PR commen- 


en, 


tary for the W&H volume: “This important addition to the botanical literature of Florida and the 
eastern United States will be welcomed by professional and amateur botanists and field biologists 
throughout the region.” W&H include 152 species and 9 hybrids; there are various differences in 
taxonomic interpretation at the species, genus, and family levels between the two treatments, but 


they cover essentially the same species. 

1 contrast to the Nelson volume, W&H provide longer and more detailed technical descrip- 
tions, detailed and formal synonymy, keys to all genera with more than a single species, and a line 
drawing (habit and details, placed with the text) of one species for each genus. Think of the W&tH 
volume as the technical companion to the Nelson volume—a serious pteridologist in the Southeast 
surely will want both books. Of course, it would have been good (from my point of view) to have all 
the information in a single volume. Gymnosperms are covered only in the W&H volume.—Guy | 
Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 


SIDA 19(3): 700. 2001 


NOTES ON THE INCREASING PROPORTION OF 
NON-NATIVE ANGIOSPERMS IN THE MISSOURI FLORA, 
WITH REPORTS OF THREE NEW GENERA FOR THE STATE 


George Yatskievych Jay A. Raveill 
Missouri Department of Conservation/ Department of Biology 
Missouri Botanical Garden Central Missouri State University 
40) Warrensburg, MO 64093, U.S.A. 
St. Louis, MO 63166, U.S.A. jar8812@cmsu2.cmsu.edu 


george. yatskievych@mobot.org 


ABSTRACT 
Three additions to the Missouri flora are reported: Fatoua vi (Moraceae), Oenanthe javanica 
oo and Ottelia alismoides (Hydrocharitaceae). Each of these represents a nonin ie 
genus new to the state. The number of introduced taxa of ae plants in Missouri continu 
increase more rapidly than that of native taxa, and the non-native component of the state's rie 
diversity has risen from ca. 22.8 percent to 27.7 percent during the past 35 years. 


RESUMEN 
Se citan tres nuevas especies para la flora de Missouri: Fatoua villosa (Moraceae), Oenanthe javanica 
(Apiaceae), y Ottelia alismoides (Hydrocharitaceae). Cada una de éstas representa un género nuevo 
no autéctono para el estado. El nimero de taxones introducidos continua incrementandose mas 
rapidamente que el de los taxones nativos, y el componente no nativo de la diversidad floristica del 
estado ha aumentado del 22.8 por ciento al 27.7 por ciento en los ultimos 35 anos 


There are many good reasons for establishing permanent programs to collect 
data on floristic diversity in various regions. Among these, perhaps the most 
surprising to non-botanists is that field and herbarium studies continue to yield 
major new distributional records, even in supposedly well-botanized areas. The 
notion that plants, unlike animals, are generally rooted in place has led to the 
flawed perception among some scientists, administrators, and politicians that 
plant distributions generally are static, and that inventory and monitoring ac- 
tivities reasonably may be curtailed following attainment of some qualitatively 
established benchmark, such as publication of a state floristic manual. 

Actual facts counter this perception. For example, for many years, knowl- 
edge of the vascular flora of Missouri was considered by many botanists to be 
relatively complete, at least compared with that of most other states. Missouri 
had an exemplary floristic manual (Steyermark 1963), praised by botanists for 
its completeness and attention to detail. Steyermark’s own extensive body of 
meticulously documented field work and more than 60,000 Missouri collec- 
tions, along with the activities of a large number of other prolific collectors, 
resulted in one of the most thoroughly d regional floras in the country. 


SIDA 19(3): 701 — 709. 2001 


702 BRIT.ORG/SIDA 19(3) 


Missouri’s landlocked midcontinental position also contributed to the notion 
of stability of the state’s flora. 

Even before the publication of Flora of Missouri, however, additional species 
records began to accumulate. In fact, Steyermark (1963) was forced to add a 3- page 
addendum to his book to include mention of 20 species that been confirmed to 
occur in Missouri after his text had been finalized for the publisher, based on 
specimens newly accessioned in herbaria. Over time, a remarkable number of 
other additions to the register of the state’s flora has been reported by a large 
number of workers. Yatskievych (1999) summarized these for the 35-year period 
following publication of Steyermark’s Flora as comprising 292 species, 25 in- 
fraspecific taxa, and 21 hybrids, a net increase of 11.9 percent. He further noted 
that the amazing rate of nearly ten new taxa discovered per year showed no 
sign of slowing. The new records represent a mixture of relatively localized taxa 
overlooked by earlier botanists, recently established populations reflecting rap- 
idly shifting or expanding ranges of some plants, and a few novelties only recently 
described as new to science. Turner and Yatskievych (1992) detailed the Missouri 
distributions of new records recorded from 1963 to 1991 and some additional 
records were discussed by Yatskievych (1999). 

At the species level, 37 percent of these reports were of taxa considered to 
be native components of the flora, with the remaining 63 percent representing 
non-native taxa (Yatskievych 1999). Thus, during the three and a half decades 
following the publication of Steyermark’s (1963) Flora, the non-native compo- 
nent of the state's floristic diversity rose from ca. 22.8 percent to 27.7 percent. It is 
worth noting that although most of the non-native additions to the flora have 
remained relative rarities, some of them have become quite common and wide- 
spread, including such weedy species as Alliaria petiolata (M. Bieb.) Cavara & 
Grande (Brassicaceae, garlic mustard), Dipsacus laciniatus L. (Dipsacaceae, cut- 
leaved teasel), Elaeagnus umbellata Thunb. (Elaeagnaceae, autumn olive), 
Lonicera maackii (Rupr.) Maxim. (Caprifoliaceae, Amur honeysuckle), and 
Ligustrum obtusifolium Sieb. & Zucc. and L. sinense Lour. (Oleaceae, privets). 
Some of the worst invasive exotics troubling property owners, land managers, 
and conservationists today were not known to grow in Missouri in 1963. 

With this in mind, we present data to justify the inclusion of three recently 
found species in the flora of Missouri. The taxa are notable in that they represent 
new generic records for the state as well, and in one case, to our knowledge, the 
species previously has not been recorded for the North American flora. We believe 
that each of these species has the potential to become more widespread in Missouri 
and possibly to become invasive in some native plant communities in the state. 


Fatoua villosa (Thunb.) Nakai (Moraceae, mulberry weed).—This monoecious 
annual is native to eastern Asia, where it often occurs in disturbed habitats. It was 
first reported for North America from Louisiana by Thieret (1964), who indicated 


that it may have been present in the New Orleans area for 15 or more years prior 
to his report. Since that time, it has become widespread in the southeastern and 
midwestern states west to Texas and has appeared sporadically in Utah, Cali- 
fornia and Washington, as well as Hawaii, Puerto Rico, and the Bahamas. For 
reviews of the literature on its geographic spread, see Vincent (1993) and Kartesz 
and Meacham (1999). Reports of occurrences in Indiana (Wunderlin 1997, 
Kartesz & Meacham 1999) require confirmation. 

Fatoua villosa isa nondescript herb with alternate, petiolate, stipulate, ovate 
to triangular leaves having trunctate to cordate bases, crenate margins, and at- 
tenuate tips. The vegetative portions strongly resemble seedlings or root sprouts 
of mulberries (Morus spp.), acne the common name UN weed.” The 
dense, flattened, brownish purple cymes containing both te and pistil- 
late flowers somewhat resemble a reduced version of a Dorstenia (Moraceae) 
inflorescence. The flowers mature at different times, so the tiny projectile seeds 
are dispersed over at least a two month period (pers. obs. in Missouri). Wunderlin 
(1997) provided a description and keys to separate Fatoua from other genera of 
North American Moraceae. 

Mulberry weed is most commonly found in greenhouses, flower beds, and 
similar highly disturbed sites. Much of its interstate spread may be in the form 
of seeds residing as contaminants in nursery stock and soil or mulch. The strong 
superficial similarities between the herbage to that of Morus suggests that plants 
of Fatoua may be overlooked by some collectors and that the species range might 
be greater than has been documented thus far. 

In Missouri, Fatoua villosa was first brought to the authors’ attention by 
Dr. Michael Vincent of Miami University, who had studied the species’ distri- 
bution in Ohio (Vincent 1993), and who noted plants in flower beds at the Mis- 
souri Botanical Garden while attending a symposium in October, 1993. Subse- 
quent herbarium studies disclosed that the species was present as early as 1990 
as a greenhouse weed at the Botanical Garden. By 1994, the plant had become a 
troublesome pest in planted areas around the property and the horticulture staff 
began an aggressive campaign to weed it out. Despite the efforts of staff and 
volunteers since then, the plants have merely decreased in numbers and size 
and not been eliminated, presumably because of a soil seed bank. Indeed, by 
1996 the species had spread to the Garden's Shaw Nature Reserve, in Franklin 
County, where it became established in tended woodland areas. Other weedy 
occurrences noted in flower beds at various locations in and around St. Louis 
City and County presumably resulted from independent introductions into the 
state. However, based on a survey of herbarium specimens, the species was 
present as early as 1972 asa greenhouse weed in Cape Girardeau. In recent years, 
localized infestations have been documented from additional counties, and the 
species eventually may be found in most metropolitan areas around the state. 
Its invasive potential remains to be determined, but it is worth noting that it 


704 BRIT.ORG/SIDA 19(3) 


has been collected along disturbed margins of at least one mesic upland forest 
site in St. Louis County. 


Specimens examined: U.S.A. Missouri. Boone Co.: Columbia, 608 E. Cherry Street, common weed 
along edge of perennial evergreen shrubbery at NW corner of Federal anerts 24 Aug 1995, McK- 
enzie 1629 (MO, UMO). Butler Co.: Poplar Bluff, 252 S Barron Road, weed in flower bed of residence, 
present for at least 2 years, 17 Sep 1994, Hudson s.n. (MO); Poplar Bluff, near steps leading from E 
she oe of eit Clinic to parking lot, at edge of lawn, 26 Aug 1996, Hudson 956 (MO). Cape 
pe aus Brooks Gardens, I1L0 N Cape Rock Drive, weed in greenhouse, 24 Sep 
ic eee sn. ae SEMO). Cole Co.: Jefferson City, Missouri Department of Conservation Head- 
quarters, in mmaulened seen of courtyard within office complex, 7 Aug 2000, Smith 3605 (MO). 
Franklin Co.: Gray Summit, Shaw Arboretum, Whitmire Wildflower Garden in shaded bed near ga- 
zebo at Send of small pond, scattered weeds in planting, 14 Sep 1996, Yatskievych & Yatskievych 96- 
82 (MO). St. Louis Co.: St. Louis City, Missouri Botanical Garden, weed in greenhouse, 19 Sep 1990, 
Miller, Merello, & Schmidt 5608 (MO); Missouri Botanical Garden, weed in flower bed in front of 
Climatron, 9 Oct 1993, Vincent 6443 (MO, MU); Missouri Botanical Garden, along edge of main walk 
from Ridgway Center to Climatron, uncommon weed in bed, presumably introduced as a seed con- 
taminant in Zinnia planting, 9 Oct 1993, Yatskievych & Yatskievych 93-371 (MO); Missouri Botanical 
Garden, weed on W side of a Administration Building, 30 Aug 1995, Lievens, Yatskievych, 
& Sitch ae 5770 (MO); Clayton; SW corner of Forsyth Blvd. and Bemiston Rd., scattered weeds in 
planting along bank building, 7 Sep 1996, Yatskievych & Yatskievych 96-80 (MO); Manchester, 957 
oft Woods Ct, uncommon along disturbed margin of mesic upland forest adjacent to back yard, 
25 Oct 2000, Yatskievych & Yatskievych 00-99 (MO). 


= 


Oenanthe javanica (Blume) DC. (Apiaceae, water celery).—American horti- 
culturalists continue to search abroad for hardy new plants to promote for use 
in gardens in the United States. This is especially true in water gardening, where 
there is a strong trend to utilize “specimen plants” exhibiting unusual growth 
forms. The water dropwort genus, Oenanthe, contains perhaps 40 species of 
mostly Old World perennials, a few of which have made their way into plant 
commerce (Mabberley 1997). Most of the species contain a number of toxic com- 
pounds (mostly polyacetylenes), and the vernacular “dropwort” generally has 
been used to denote plants responsible for animal or human poisoning. Thus, 
some species have been cultivated on a small scale for medicinal, pharmaceuti- 
cal, or biochemical investigations. 

The most commonly cultivated species of Oenanthe have been recorded 
as localized escapes: O. aquatica (L.) Poir. (water fennel), from Franklin County, 
Ohio (Cooperrider 1995); and O. pimpinelloides L., from Humboldt County, Cali- 
fornia (Constance 1993). To these, we now add O. javanica, from Johnson County, 
Missouri, apparently the first report of this species’ establishment outside of 
cultivation in North America. 

Oenanthe javanica is a native of southeastern Asia and the Indo-Malay- 
sian region. As North American botanists may be unfamiliar with the species, 
the following description has been adapted from the treatment in the forth- 
coming dicot volume of Steyermark’s Flora of Missouri (Yatskievych, in prep.): 

Plants perennial, glabrous, with fibrous roots, lacking tubers. Stems 30-150 


cm or more long, spreading with ascending branches and tips, somewhat in- 
flated, rooting at the lower nodes. Leaves alternate and sometimes also basal, 
short- to long-petiolate, the petioles somewhat inflated. Leaf blades 3-20 cm 
long, ovate to triangular-ovate in outline, pinnately 1-2(-3) times compound, 
the leaflets 10-50 mm long, narrowly lanceolate to broadly ovate, rounded, 
narrowed, or tapered (sometimes unequally so) at the base, finely to more com- 
monly coarsely toothed along the margins, occasionally with 1 or 2 basal lobes. 
Inflorescence an umbel, opposite the leaves and occasionally also terminal, 
mostly long-pedunculate. Involucre absent or less commonly of | or 2 bracts, 
these shorter than the rays, spreading to ascending at flowering, linear, with 
sharply pointed tips. Rays (4-)6-20, 0.5-3.0 cm long, strongly angled and with 
entire or minutely toothed, pale angles or narrow wings. Involucel of 7-13 
bractlets, these shorter than to more commonly longer than the pedicels, linear, 
sometimes with thin, white, papery margins, tapered to sharply pointed tips. 
Flowers 5 to numerous in each umbellet, all perfect, epigynous, the pedicels 1-5 
mm long. Sepals 5, minute triangular teeth. Petals 5, obovate, appearing shal- 
lowly notched apically but narrowed abruptly to a slender pointed extension 
at the tip, white. Ovary inferior, 2-carpellate, glabrous. Fruit a schizocarp, some- 
times shed while still green, 2-3 mm long, oblong in outline, somewhat flat- 
tened laterally, glabrous, the 2 mericarps each with 5 ribs, these blunt, and broad, 
tan or light yellow to straw-colored, all or mostly obscuring the reddish brown 
surfaces between them. 

This species was first collected in Missouri in 1996 by Lisa Wilson, a student 
in the plant taxonomy class at Central Missouri State University, but initially 
went unrecognized as a new record. Confirmation of its identity required col- 
lection of fruiting material during the 2000 field season. All collections originated 
from Race Horse Lake, a 0.73 ha (8 acre) artificial lake in the Pertle Springs 
Biological Study Area, owned by Central Missouri State University and located 
in Warrensburg, Johnson County. The population, which apparently originated 
near a bridge on the southern side of the lake, has been monitored since 1997 
and has spread to occupy the entire shoreline of the lake. There are no records 
of its intentional cultivation at the site and the circumstances of its establish- 
ment there are not known. The peo occurs as - emergent eee with com 
mon associates including Polygonum hyd 
atrovirens, Boehmeria cylindrica,and Bidens cernua. Its potential for eae a 
serious weed is apparent in a 20 x 50 m, seasonally-inundated area along the 
southern side of the lake, where it forms a near monoculture. However, it isa 
minor vegetational component where steep slopes limit the littoral zone to a 
narrow strip. Oenanthe javanica has not spread to other ponds and lakes in the 
drainage, but is found in pockets of soil in the concrete spillway that drains 
Race Horse Lake and leads directly to the adjacent 6 ha (15 acre) Cena Lake. 
Future spread will be monitored and a plan to eliminate the population of 


706 BRIT.ORG/SIDA 19(3) 


Oenanthe is being formulated by the CMSU Facilities and Grounds personnel 
in consultation with the Biology Department faculty. 

Water celery is unusual in the genus Oenanthe in its edible herbage, and 
the species has a long history of cultivation in southeastern Asia and Malesia 
as a vegetable and potherb. Its adoption into North American horticulture ap- 
parently has been relatively recent, and the species was not included in Hortus 
Third (Liberty Hyde Bailey Hortorium staff 1976). Its popularity has risen in 
the last few years and cultivars have begun appearing on the market, such as 
‘Flamingo’ with pinkish foliage. One indication of its popularity is that a recent 
search of the World Wide Web yielded about 65 sites including mention of the 
genus Oenanthe, principally online nursery catalogs. Some of these nursery 
catalogs include comments on the potential aggressiveness and vegetative 
spread of the species in some garden situations. That it can be dispersed by seed 
is evidenced from our conversations with horticulture staff at two botanical 
gardens in Indiana and Missouri, where plants grew spontaneously as contami- 
nants in plantings of other aquatic species. 

It should be noted that there is no modern monograph of Oenanthe. Sev- 
eral taxa originally treated as separate species have been reduced to infraspe- 
cific status by more recent workers, (e.g, Murata 1973). If one accepts the taxo- 
nomic validity of multiple varieties in this morphologically plastic taxon, then 
Missouri materials (and horticultural materials in the United States in general) 
should be referred to var. javanica, as currently circumscribed. Additionally, in 
working to determine the Missouri materials, we were surpised at the similari- 
ties between O. javanica and the North American native, O. sarmentosa C. Presl 
ex DC. (Pacific water dropwort), which occurs in wetlands mostly near the coast 
from Alaska to California. This species has been brought into cultivation lo- 
cally within its native range, but apparently has not been marketed outside the 
Pacific Northwest. Regrettably, the only treatment thus far to attempt to con- 
trast the two species is that of Hiroe (1979), whose key separated them on the 
basis of subtle differences in involucre and involucel size and shape, which are 
variable in both taxa. Future monographers may determine that O. javanica and 
O. sarmentosa should be combined into a single species with a disjunct distri- 
bution around the Pacific Rim. 


Specimens examined: U.S.A. Missouri. Johnson Co.: Warrensburg, Pertle Springs, Race Horse Lake, 6 
Oct 1996, Wilson 113 (MO); same locality, 17 Aug 2000, Raveill 3175 (MO, NEMO, SEMO, UMO, WARM): 
same locality, 26 Aug 2000, Raveill 3176 (MO, NEMO, SEMO, UMO, WARM). 


Ottelia alismoides (1_.) Pers. Hydrocharitaceae, duck lettuce) —The pantropical 
genus Ottelia comprises 21 mostly Old World species of aquatic annuals and 
perennials (Cook et al. 1984; Cook & Urmi-Konig 1984). One species, O. 
alismoides, which is native from Asia to Australia, has become widely estab- 
lished as an aquatic weed in parts of Africa, Europe, and North America. In the 


707 


United States, where the U.S. Department of Agriculture has listed the species 
as a federal noxious weed, it was first reported from southwestern Louisiana 
(Holmes 1978), where it was collected in 1939 (Haynes, 2000) and apparently 
remains well-naturalized in lakes and reservoirs in five parishes (Thomas & 
Allen 1993). Hatch et al. (1990) reported it from adjacent southeastern Texas 
without citation of localities or vouchers. A population in Butte County, Cali- 
fornia, in a ditch associated with rice fields was exterminated even before it 
could be reported in the literature (Turner 1980). The report from Missouri is 
the first example of a population occurring at a significantly inland location. 

Ottelia alismoides isa short- oe robust, submerged aquatic with large, 
long-petiolate, mostly ovate-cordat bling those of a plantain (Plantago) 
or water plantain (Alisma). The long-pedunculate, 1-flowered, emergent inflo- 
rescences are enclosed basally in a spathe with several undulate wings or ribs. 
The usually perfect flowers have 3 showy (2-3 cm long) white to pale pink petals. 
For more complete descriptions of this species and keys to its determination, see 
Godfrey and Wooten (1979), Cook and Urmi-Kénig (1984), and Haynes (2000). 

In July 2000, a population of Ottelia was located in southeastern Missouri 
by herpetologist Jeff Briggler, who, with other biologists from the Missouri De- 
partment of Conservation, was conducting reptile and amphibian surveys in 
two adjacent artificial wetlands at the Big Cane Conservation Area, in Butler 
County. These sites had been excavated in 1998 to create marsh habitat for the 
state-endangered western chicken turtle (Deirochelys reticularia miaria). There 
is no evidence that propagules of Ottelia were transported to the site acciden- 
tally on the equipment used for the excavations, so the plants are presumed to 
have been introduced into the area by migratory waterfowl. The shallow de- 
pressions in sandy soil presently contain a young successional wetland plant 
community, including individuals and patches of such associates as Alisma 
triviale, Azolla mexicana, Echinodorus cordifolius, Eleocharis spp., 
Heteranthera spp., Ludwigia peploides, Myriophyllum pinnatum, Potamogeton 
foliosus, Sagittaria calycina, and Sphenoclea zeylanica. Plants of Ottelia origi- 
nally were noted flowering in the northernmost of the two areas, but a subse- 
quent visit by local botanist Stanton Hudson (who is completing a floristic sur- 
vey of Butler County) disclosed a few plants in the more southern area as well. 
Other sites in the vicinity have not been searched yet for the presence of duck 
lettuce. The Missouri Department of Conservation is preparing to attempt the 
eradication of plants at the known sites and to survey for the occurrence of 
duck lettuce in adjacent areas. In addition to the possibility that O. alismoides 
may spread to natural wetlands in the southern portion of the state, the poten- 
tial exists for this species to invade rice fields in Missouri’s Bootheel. 


1: U.S.A. Missouri. Butler Co.: Big Cane Conservation Area, ca. 3 mi S of hee 
ca. 21/3 mi NNW of parking lot on County Rd 278, in water of shallow man-made s, 27 Ju 
2000, Briggler, Pelton, Johnson & Urichs.n.(MO, UMO); same locality, 12 Sep 2000, Se ane O). 


708 BRIT.ORG/SIDA 19(3) 


ACKNOWLEDGMENTS 


The authors wish to thank the following individuals, who provided informa- 
tion and/or review toward one or more of the reports in this paper: Missouri 
Department of Conservation Biologists Jeff Brigler, Karen Kramer, Tim Smith, 
and David Urich; US. Fish @ Wildlife Service Biologist Paul McKenzie; Stan 
Hudson, formerly of Poplar Bluff; Becky Sucher, plant recorder at Missouri Bo- 
tanical Garden; David Bauman, botanist at White River Gardens; and Mike Vin- 
cent, of Miami University. 


REFERENCES 


Constance, L. 1993. Apiaceae (Umbelliferae], carrot family. In: J.C. Hickman, ed. The Jepson 
manual, higher plants of California. University of California Press, Berkeley. Pp. 136-166. 

Cook, C.D.K. and K. Urmi-Konie. 1984. A revision of the genus Ottelia (Hydrocharitaceae) 2. 
The species of Eurasia, Australasia, and America. Aquatic Bot. 20:131-177. 

Cook, C.D.K., J.-J. Symoens, and K. Urmi-Konic. 1984. A revision of the genus Ottelia 
(Hydrocharitaceae) |. Generic considerations. Aquatic Bot. 18:263-274. 

Coorerriver, 1.S.1995. The Dicotyledoneae of Ohio. Part 2:Linaceae through Campanulaceae 
Ohio State University Press, Columbus. 

Goorrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United 
States. Monocotyledons. University of Georgia Press, Athens. 

Harct, S.L., KN. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas 
Agric. Exp. Stat. Publ. MP-1655, College Station. 

Haynes, R.R. 2000. Hydrocharitaceae Jussieu, tape-grass or frog-bit family. In: Flora of North 
America Editorial Committee, eds. Flora of North America north of Mexico. Volume 22. 
Magnoliophyta: Alismatidae, Arecidae, Commelinidae (in Part), and Zingiberidae. Ox- 
ford University Press, New York. Pp. 26-38. 

Hiroe, M. 1979. Umbelliferae of World. Ariake Book Co., Tokyo. 

Hoimes, W.C. 1978. Range extension for Ottelia alismoides (L.) Pers. (Hydrocharitaceae). 
Castanea 43:193-194. 

Karresz, J.T. and CA. Meacham. 1999. Synthesis of the North American flora, version 1.0. 
North Carolina Botanical Garden, Chapel Hill [CD-ROM]. 

LiBertY Hype BaiLey HorTorium Starr. 1976. Hortus third. MacMillan Publishing Co., New York. 

Masseriey, D.J. 1997. The plant book, a portable dictionary of the vascular plants, second 
edition. Cambridge University Press, Cambridge, Great Britain. 

Murata, G. 1973. New or interesting plants from Southeast Asia 1. Acta Phytotax. Geobot. 
25:97-106. 

STEYERMARK, J.A. 1963. Flora of Missouri. lowa State University Press, Ames [errata added at 
second printing, 1968]. 

THieRET, J.W. 1964. Fatoua villosa (Moraceae) in Louisiana:new to North America. Sida 1:248. 

THomas, R.D. and C.M. Atten. 1993. Atlas of the vascular flora of Louisiana. Volume I: Ferns 
and fern allies, conifers, & monocotyledons. Louisiana Department of Wildlife and Fish- 


eries, Baton Rouge. 


Turner, C.E. 1980. Noteworthy collections. Ottelia alismoides (L.) Pers. (Hydrocharitaceae). 
Madrofo 27:177. 

Turner, J.and G. Yarskigvycu. 1992. County record vouchers for vascular plant species newly 
recorded for Missouri since 1963. Missouriensis 13:1-24. 

Vincent, M.A. 1993. Fatoua villosa (Morac ede), Mt ilbert y weed, in Ohio.Ohio J.Sci.93:147-149, 

Wunper tin, R.P. 1997. Moraceae Link, mulberry family. In: Flora of North America Editorial 
Committee, eds. Flora of North America north of Mexico. Volume 3. Magnoliophyta: 
Magnoliidae and Hamamelidae. Oxford University Press, New York. Pp. 388-399. 

YatskievYcH, G. 1999. Steyermark’s flora of Missouri, revised edition, volume 1. Missouri 
Department of Conservation, Jefferson City. 


BRIT.ORG/SIDA 19(3) 


Book REVIEW 


JEAN ANbREWS. 1998. The Pepper Lady’s Pocket Pepper Primer. (ISBN 0-292-70467- 
4, hbk. 0-292-70483-6, pbk.). University of Texas Press, PO. Box 7819, Austin, 
TX 78713-7819, U.S.A. (Orders: http://www.utexas.edu/utpress, 512-471- 
4032). $17.95, 184 pp, 94 full color photographs, b/w illustrated glossary, 


" 


x 


d . 


the pungent pods 


“This little book is intended to clarify some of the li undil 
indigenous to ae mbus’ New World,” says the author. The book may be iw but like the tiny 
Chilpequjn, Capsicum annum var. glabriusculum, it packs a lot in its small size! This time, author/ 
artist ce eee shows us her equally adept skill as photographer in this slick and colorful 


hand book. 
I ick ref identificati id he titl a ‘pocket primer’). 


Chapters can be found on nauieneauie how to re ae domesticated erie what makes a pep- 
per hot, discussions on color, aroma, flavor and nutritional value, how to select and store peppers, 
how to roast and rehydrate, and how to cultivate them 

rty-five peppers are identified and discussed in the chapter simply titled: The Peppers. Each 


are Tee reer by common type/group name with sclenutic names aaa peed the com- 
} 


mon name. Each are described and a aaaaiues color } 
+h 


Th | I | peppers in the 
Addition al ar is consistent “a uniform categories: SIZE, COLOR, FRUIT SHAPE, 
CY, SUBSTITUTES, OTHER NAMES, SOURCES, USES, AND REMARKS. 
i. sre is an extensive Illustrated Glossary showing simple line drawings of fruit shape at pe- 
fruit shape at blossom end or apex, and fruit cross-section. An explana- 


a 


book. 
FLESH 


duncle (stem 
tion of fruit ia: and a Pungency Rating/Heat Scale are also provided. Sources for locating 
oe dried su ten a aaa prepared products and seeds are included. In the back isa list of Seed 

a few addresses and, of course, an index. Plus, Jean adds some personal remarks 


a 

oe on her years of ee on Hie eect Spits of apse: 

oif youare looking for 

de los a strikes again aes a an new creation. a his book is designed for pepper lovers of all 

types, professional or not. Brava, Senora Andrews! —Linny Heagy, Linny/Designer, Illustrator, Arling- 
ton, 1X, U.S.A. a0005835@adirmail.net. 


} Jnce again b tra Sen 


SIDA 19(3): 710. 2001 


GLOCHIDION PUBERUM (EUPHORBIACEAE) 
NATURALIZED IN SOUTHERN ALABAMA 


Miriam L. Fearn Lowell E. Urbatsch 
Depa tment t of Eartt tn Sciences oe Sciences Department 
University of South Alabama Louisiana State University 
Mobile, AL 36688-0002, U.S.A. Baton as LA 70803-1705, U.S.A. 


During the month of July 2000, a colony of about ten unusual, small trees was 
observed growing in a wooded area near Halls Mill Creek in Mobile, Alabama. 
They have supscquently been identified as Glochidion puberum (L.) Hutch., a 
member of Euphorbiaceae (Fig. 1). The largest tree was approximately 4.5 m 
tall with a diameter of nearly 7.5 cm at its base while the mance was a seed- 
ling less than 5 dm tall. Simple, alternate, distichous leaves terized these 
plants. Numerous, axillary clusters of small, yellowish flowers and young fruit 
were evident on the larger individuals. On subsequent visits to the site mature 
fruits were observed. 

The Alabama occurrence appears to be the first known record of this spe- 
cies naturalized in the United States. Specimen data from Harvard University 
Herbaria (HUH) indicate that this taxon is widespread in southeastern China 
where its occurrence is documented by specimens for the following provinces: 
Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hong Kong, Hubei, 
Hunan, Jiangsu, Sichuan, Taiwan, Yunnan, Zhejiang (Anthony Brach, pers. 
comm.). This species is known to be cultivated in Florida. University of Florida 
Herbarium (FLAS) specimen data indicate its cultivation on the University of 
Florida campus in Gainesville, Alachua County. According to label data, a single 
tree reported near St. Augustine in St. Johns County, Florida was obtained from 
the University of Florida. 

The source of this material in Alabama is unknown. To our knowledge, G. 
puberum is not cultivated in the region; area plant nursery personnel and state 
agriculture extension staff were not familiar with this species. No record of it 
for Alabama was found in the Freeman Herbarium, Auburn University. This 
plant has few attractive features and, therefore, is probably not sought for orna- 
mental use. Certain species of Glochidion are available in the nursery trade in 
tropical regions of the world where they are used for hedge plantings and refor- 
estation (Alstonville Tree Farm 2000). Glochidion puberum reportedly has 
many ethno-botanical medicinal uses in China in the treatment of abscess, 
amenorrhea, arteritis, snake bite, dysentery, enteritis, flu, laryngitis, malaria, 
sores, and trauma (Duke 1997). A Chinese name for the plant is “Suan p’an tzu” 


SIDA 19(3): 711 — 714. 2001 


712 BRIT.ORG/SIDA 19(3) 


or “abacus plant” because the seeds resemble small abacus beads (Kam-biu Liu, 
pers. comm.). 

The area in Alabama where the plant was discovered is beside a roadway 
through a degraded wetland on the south side of Halls Mill Creek. The 
Glochidion plants are growing aproximately five meters from the edge of the 
asphalt road and about two meters from the wetland itself. They are on better- 
drained soil about halfway up the roadside embankment, one meter above 
poorly drained, acidic, wetland soils and standing water. Overstory vegetation 
consists primarily of scattered Pinus taeda L. near the road and a few Acer 
rubrum L. near the creek itself. In close proximity to the G. puberum colony are 
other non-natives suchas Sapium sebiferum (L.) Roxb., Ligustrum sinense Lour, 
and Lonicera japonica Thunb. Nearby native plants include Cyrilla racemiflora 
L. and Myrica cerifera L. The site is relatively shady due to the surrounding 
Pinus and relatively tall Sapium. 

Glochidion puberum are large shrubs or trees. According to label data, one 
individual on the University of Florida campus was multi-trunked, 10 meters 
tall. Based on the material from Alabama the bark is brown with closely spaced 
fine longitudinal furrows; milly sap or exudate absent; twigs tan, densely pu- 
bescent; hairs uniseriate, spreading or tangled. Leaves simple, alternate, disti- 
chous, deciduous, 5-7 cm long, 2-3 cm wide, abaxially pubescent; blades nar- 
rowly elliptic, somewhat coriaceous; apex acute; bases rounded, asymmetric: 
margin entire; venation pinnate, secondary veins ca. 9 pairs, evenly spaced, 
prominent, arcuate, yellowish; petiole ca. 4 mm long, rusty-brown, densely pu- 
bescent; stipules laterally placed, free of one another, scale-like, acute 1-2 mm 
long. Inflorescences axillary, 10-20 flowers per cluster; pedicels ca. 2mm long. 
Flowers at least some unisexual with staminate and pistillate flowers on the 
same plant, actinomorphic; ca. 5 mm in diameter, perianth consisting of 6 se- 
pals, distinct, persistent in fruit; petals absent; stamens ca. 8; filaments joined, 
free of the perianth; ovary superior; carpels 4-5, bilocular; styles bi-lobed. Fruit 
capsular, pale green to yellow, ca. 14 mm in diameter. Seeds reddish-orange, ca. 
4 mim long (Fig. | 

Glochidion is mainly a tropical genus of more than 200 Asian, Australasian, 
and Polynesian species (Webster 1994). The native range of G. puberum, in China 
and its occurrence in the Gainesville, Florida area suggest that the taxon pos- 
sesses some frost tolerance and may otherwise be pre-adapted to conditions in 
the southeastern U.S. 


Voucher specimens: ALABAMA: Mobile Co. naturalized in a wooded area near roadside on west side 


of Demetropolis Road immediately south of Halls Mill Creek Bridge, small tree approximately 7.5 
cm in diameter at the base and 3.5-4 m tall growing with others, 30° 36.35' N, 88’ 9.42’ W: 12 Jul 2000, 
Fearn s.n. (LSU). Additional specimens have been distributed to AUA, DAV, HUH, and L. 


2 didintiandlinianencns 


R h £ a 
Fig. 1. Bran I | h A leaf 
I g twig - 


714 BRIT.ORG/SIDA 19(3) 


ACKNOWLEDGMENTS 


Thanks to Grady Webster, University of California, Davis and Jeremy Bruhl, 
Director, N.C.W. Beadle Herbarium (NE) University of New England, Armidale, 
Australia for their assistance with identification. Kent Perkins, University of 
Florida Herbarium, Florida Museum of Natural History, Gainesville and Bruce 
Hansen, Herbarium University of South Florida, Tampa provided specimen data 
for this species in Florida. Distribution data for this species in China was pro- 
vided by Anthony Brach and David Boufford, Harvard University Herbaria. 
Kam-biu Liu and Roland Roberts, Louisiana State University, provided anec- 
dotal information about the taxon, and comments on the manuscript, respec- 
tively. Support from National Science Foundation grant DBI-9987491 and from 
the Departments of Earth Sciences and Biological Sciences at our respective 
institutions is gratefully acknowledged. 


REFERENCES 


AtsTONviLLe TREE Farm. 2000. Catalogue. Alstonville, New South Whales. Australia http:// 
om.au. (7 February 2001). 

Duke, J.A. 1997. Ethnobotanical uses of Glochidion puberum (EUPHORBIACEAE). Phy- 
tochemical and Ethno Botanical Databases, Agricultural Research Service, Phytochemi- 
cal Database, USDA. http://www.ars-grin.gov/cgi-bin/duke/ethnobot.pl?Glochidion 
%20puberum. (7 February 2001). 

Weester, G.L. 1994. Classification of the Euphorbiaceae. Ann. Missouri Bot.Garden 81:3-32. 


RADIATE AND ERADIATE INDIVIDUALS IN 
GRINDELIA NUDA (ASTERACEAE) 


R.T. Harms 
Department of Linguistics 
University of Texas at Austin 
Austin, TX 78712-1196, U.S.A. 


Grindelia nuda Wood var. nuda is a common rayless taxon of central Texas. 
However, a population exhibiting both radiate and eradiate heads exists in NW 
Hays County in central Texas. | first noted this when plants cultivated from 
seeds collected from rayed plants produced both rayed and rayless individuals 
(Figs. 1 and 3; scales represent mm). Subsequent inspection of the locale, along 
a quarter mile of county road, revealed roughly a dozen specimens of each type. 
The two types typically did not grow together. Additionally, one individual from 
my planting was essentially “subradiate,” with rays varying in length from 1-4 
mm (Fig. 2), in contrast to rays of 10-14 mm on regular radiate plants. 

Voucher specimens (verified by B. L. Turner): TEXAS. Hays Co.: NW part of county on roadside 
immediately above Deadman’s Hole, 27 Oct 2000, Harms 11 (rayless) & Harms 12 (rayed) (TEX). 
Since the presence or absence of rays is emphasized in recent keys for Texas 
Grindelia (eg,, Diggs et al. Shinners & Mahler's illustrated flora of North Cen- 
tral Texas, 1999), a revision for the key seems in order, perhaps utilizing achene 
morphology asa distinguishing feature following G. Nesom, Phytologia 68:304, 
1990: 


Besides the presence of ray flowers, at least the Great Plains forms of G. squarrosa differ from G. 
nuda in the production of dimorphic achenes: the ray and outer disc achenes of G. squarrosa are 
smooth and compressed but slightly 3-4 angled, while the inner disc achenes are longer than the 
outer, strongly compressed and 2 angled, and have numerous superficial, longitudinal nerves, the 
achenes of G. nuda are monomorphic, all smooth or developing shallow furrows late in their matu- 
ration. 


ACKNOWLEDGMENTS 


Iam grateful to Tom Wendt and BL. Turner for their assistance in preparing 
this report. 


SIDA 19(3): 715 — 717. 2001 


) 


BRIT.ORG/SIDA 19(3 


716 


Fic. 1. Radiate Grindelia nuda. 


Fic. 2.”Subradiate” Grindelia nuda. 


717 


HARMS 


Eradiate Grindelia nuda. 


Fic. 3 


718 BRIT.ORG/SIDA 19(3) 


Book REVIEW 


Epwarb FE: ANDERSON. 2001. The Cactus Family. (ISBN 0-88192-498-9, hbk.). Timber 
Press, 133 S.W. Second Ave., Suite 450, Portland, OR 97204, USA. (Orders: 
www.tim berpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). 
$99.95, 776 pp, L008 color photos, 6 b/w photos, 3 drawings, 8 maps, hard- 
cover, 8" x 11" 

This is another fine publication from Timber Press, this time by author Edward F Anderson, who 

has spent over 45 years researching cacti and is currently in residence at the Desert Botanical Gar- 

den, Phoenix, Arizona. The author brings a number of talents to the table, an ethnobotanical inter- 


st, a conservationist ethic, and exceptional photography. The Cactus Family is an encyclopedic ret- 


erence to the genera of the oe sannly and is nothing short of landmark 


e book begins with fi g introductory information on nie science of cacti. 
nea one covers ae seoduuee le features of cacti, such as growth forms, stems, roots, flowers, as 
tributi vues eens ethnobotany of cacti. This 


chapter ets covers peyote the San Pedro cactus, Saguaro, cacti for food and medicine, cer- 
emonial and religious uses, as sources of dyes, and its role in horticulture. This chapter is highlighted 
by color photographs of indigenous people preparing cacti for a number of uses. Chapter three is 
devoted to the conservation of cacti, specifically in situ and ex situ conservation and the legal pro- 
tection of cacti. Chapter four is contributed by Roger Brown and covers the cultivation of cacti. Light, 
containers, potting media, water, fertilizer, air circulation, pests, and propagation are all covered in 
this chapter. Chapter five covers the classification of cacti, the problems with it, its history, and an 
overview of the cactus classification of se saasnanaaiah Cactaceae Systematics Group. These chap- 


ters set a bulk of the book, tl the famil 
vered in the treatm 125 genera and 1810 species. For each taxon, its latin and common 


names are ae ,along with a description of the plant and a color photograph where available. There 
are 1000 color photos in all making this an invaluable resource for cactus lovers and investigators. 


An ere of maps and botanic ais and 2 rbaria with se pensain collections of cacti, a glos- 
sary, literature cited, and indices of scienti book. 

There is no other book that eee discusses the botany, conservation, cultivation 
and apnopotany of sac than The Gactils Family. This Sua of a lifetime of research this 


a =e eae Ly | 


beautifully f the Cactaceae family that will be use- 


ful to the systematist, eihnobotnict or conservationist among us. Kev nD. Janni, Botanical Re- 
search Institute of Texas, Fort Worth, TX 76102-4060, U.S.A, kjanni@britorg. 


SIDA 19(3): 718. 2001 


CUPRESSUS ARIZONICA (CUPRESSACEAE) NEW TO THE 
DAVIS MOUNTAINS OF WEST TEXAS 


John P. Karges James C. Zech 
The Nature Conservancy Department of Biology 
Box 2078 Sul Ross State University 
Fort Davis, TX 79734, U.S.A. Alpine, TX 79832, U.S.A. 
jkarges@tnc.org jzech@sulross.edu. 


The Arizona Cypress, Cupressus arizonica Greene (Cupressaceae) has previ- 
ously been reported to occur natively asa well-publicized population ata single 
Texas location in the Chisos Mountains (Vines 1960; Correll & Johnston 1979; 
Powell 1988; Simpson 1988; Cox & Leslie 1997; Powell 1998). This species is con- 
sidered to be widely distributed throughout northern Mexico and the south- 
western United States, including southern Arizona, southwestern New Mexico, 
and southern California (Vines 1960; Correll & Johnston 1979; Powell 1988; Cox 
&r Leslie 1997; Powell 1998). In Mexico, Arizona Cypress is known from the Si- 
erra Madre Occidental into extreme northeastern Sonora and Chihuahua in the 
west and southward on the east to southern Coahuila to the Sierra de Parras 
(Peattie 1950), and into Zacatecas. It is well known in the Maderas del Carmen 
Mountains in northern Coahuila, approximately 40 miles southeast of the 
Chisos (Wood et al. 1999) in montane conifer forests. In Texas, C. arizonica has 
been thought to be restricted to a similar forest type occurring in Boot Canyon 
(e.g., Warnock & Hinckley 7138, SRSC) within the Chisos Mountains of Big Bend 
National Park in Brewster County (Vines 1960; Correll & Johnston 1979; Powell 
1988: Simpson 1988; Cox & Leslie 1997; Powell 1998). This limited distribution 
has puzzled phytogeographers (Simpson 1988) as to why C. arizonica occurs in 
the Chisos Mountains but not in other high Trans-Pecos Mountain ranges (¢.g., 
Davis and Guadalupe). 

Cupressus arizonica has been recently collected from The Nature 
Conservancy’s Davis Mountains Preserve in the Davis Mountains of Jeff Davis 
County. The location is a steep, forested canyon at an elevation between 2072 
and 2103 m. It was found just below Bridge Spring (Mt. Livermore, 7.5 min, USGS 
topographic quad). The canyon floor is mesic with permanently saturated soils 
from the spring. This drainage is an eastern headwater tributary of Madera 
Creek and Canyon which drains the north slopes of Mount Livermore, the highest 
summit of the Davis Mountains. The canyon is north trending, shaded and steep 
enough to maintain a cool, mesic microclimate compared to the much more 
xeric south and west facing drainages nearby. The primary community type is 
pinyon-oak-juniper forest with gray oak (Quercus griseus), alligator juniper 


SIDA 19(3): 719 — 721. 2001 


720 BRIT.ORG/SIDA 19(3) 


(YJuniperus deppeana), Texas madrone (Arbutus xalapensis), some Gambel’s oak 
(Q. gambeli), silverleaf oak (Q. hypoleucoides), southwestern chokecherry 
(Prunus serotina), and three species of pine, Mexican pinyon (Pinus cembroides), 
ponderosa (P. ponderosa), and southwestern white pine (P. strobiformis). Our 
collection, from an isolated tree, indicates a more widespread distribution (pos- 
sibly dating back to Pleistocene) as suggested by Cox and Leslie (1997), as well 
as a more continuous distribution of C. arizonica with both Texan and Mexican 
populations to the south and New Mexico, Arizona, and California populations 
to the north and west. This newest specimen agrees in habitat with previous 
collections. Isolated, forested, canyon bottoms at high elevations, associated with 
springs, are habitat characteristics which appear common for this species 
(Elmore 1976; Correll & Johnston 1979; Powell 1988; Cox & Leslie 1997: Powell 
1998: Watson & Eckenwalder 1993). Cupressus arizonica has been extensively 
planted as ornamentals and wind-breaks throughout the Trans-Pecos (Powell 
1988, 1998), and may be considered the most widely planted tree in west Texas 
(Simpson 1988). While introduction through ornamental stock remains possible, 
the height, overall size, and habitat of the tree is comparable to those within the 
Chisos. This may indicate that the Davis Mountain collection isa post-Pleistocene 
relictual fragment of a more extensive Texas population resulting from the ex- 
tension of this species’ natural distribution. Our on-going investigations in the 
Davis Mountains and other insular West Texas mountain ranges will continue 
to elucidate the pattern of distribution and differentiation among species of 
Cupressaceae including Cupressus and Juniperus. 


Voucher specimen: TEXAS. Jeff Davis Co.: Bridge Gap Spring, ca. 75 ft downstream from spring 
box, ca. 40 ft tall, 20 Jun 1998 JP Karges & L.K. Hedges 2480 (SRSC). 


ACKNOWLEDGMENTS 


The authors thank the Nature Conservancy of Texas; Linda K. Hedges for field 

assistance; A. Michael Powell for specimen preparation; Sharon Yarborough of 

SRSC; and the reviewers for helpful comments. 

REFERENCES 

Cornett, D.S.and M.C. Jounston. 1979. Manual of the vascular plants of Texas. The University 
of Texas at Dallas, Richardson. 

Cox, PW. and P. Lestit 1997. Texas trees a friendly guide. Corona Publishing Company, San 
Antonio. 

E:more, F.H.1976, Shrubs and trees of the southwest uplands. Southwest Parks and Monu- 
ment Association, Tucson. 

Peartic, D.C. 1950. A natural history of Western trees. Houghton Mifflin Co, Boston, 

Powett, A.M. 1988. Trees & shrubs of Trans-Pecos Texas including Big Bend and Guadalupe 
Mountains National Parks. Big Bend Natural History Association, Inc., Big Bend National 
Park, 


KARGES AND ZECH, CUPRESSUS ARIZONICA IN TEXAS 72) 


Powett, A.M. 1998. Trees and shrubs of the Trans-Pecos and adjacent areas. University of 
Texas Press, Austin. 

Simpson, B.J. 1988. A field guide to Texas tress. Gulf Publishing Company, Houston. 

Vines, R.A. 1960. Trees, shrubs, and woody vines of the southwest. University of Texas Press, 
Austin. 

Watson, F.D. and J.E. Eckenwatoer. 1993. Cupressaceae. In: Flora of North America Editorial 
Committee, eds. Fl. North Amer. Oxford Univ. Press, New York and Oxford. Vol. 2:399- 


Woop, S., G. Harper, E. Mucpavin, and P. Nevitte. 1999. Vegetation map of the Sierra del Car- 
men, U.S.A. and Mexico. Final Report. U.S. Geological Survey, National Wetlands Res. 
Center, and National Park Service. 


722 BRIT.ORG/SIDA 19(3) 


Book REVIEW 
MicHaet J. BALICK, MICHAEL H. Ner,and Daniet E. AtHA. 2000. Checklist of the Vascular 
Plants of Belize with Common Names & Uses. (ISBN 0-89327-440-2, pbk.). 
Memoirs of the New York Botanical Garden, Volume 85. New York Botanical 
Garden Press, Bronx, New York 10457-5126, U.S.A. $42.34, 246 pp, 6" x 9". 


This is the first floristic checkli I (less than 200,000) in Cen- 
tral America, Belize. The first product to come from the collaboration between ji New 30K pos 
le a ] ¥ 1s - 4 ] } ja Bey ores | 1 


1997). Another volume, The Ethnol ical Di itvof Belize, isi i fs chiswolemeseos 


species in 1219 genera and 209 families are recognized with l, 2% en for this subtropical envi- 
ronment 


The design follows that of other eae of the Garden with a preface, abstract, introduction 
d end 


(covering: geography, climate, vegetatio1 and endemism), a section covering 
classification and family sequence, and sections Gee as such: Psilophyta, eee ta 

Polypodiophyta, Pinophyta, Cycadophyta, Magnoliophyta: Magnoliopsida, and Magnol iophyes 
L eisai as sections are es by a pyblogra Day and general index. For each plant covered 


synonyms, uses, references 


notes (e.g. taxonomic problems), h habit, vouchers, types, and cultivation notes. The authors recognize 


nineteen categories for plant uses het beverages, construction, dyes, fibers, edibl | ani- 
mal forage, fuel, fumitory, gums, latex, medicines, oils, ornamentals poisons, resin eal spices and 


tannins. A major strength of this eae is its erudite foundation in the historical botany of Belize, 
acknowledging the contributions of major collectors (indigenous and Western) over time and its 
framework for designing the Belize Beniebolny Bree and eae rese 


This isa must-l a a ogy, Hee ecol- 
ogy, economic botany and saan i of Meso- fae I wou ld a caaeaa a udents and 


professionals alike for its cl tion to botanical and biogeographical detail ee pee on 


local culture and uses. All libraries ae a Latin American biodiversity focus should have a copy, as 
well as all institutions supporting tropical botany programs. 
REFERENCES 

SHELDON, J.W., M.J. Batick, and S.A. Lairp. 1997. Medicinal Plants: Can Utilization & Conserva- 
tion Coexist? Advances in Economic Botany vol. 12, New York Botanical Garden Press, 
Bronx, New York. 

—Kevin D. Janni, Botanical Research Institute of Texas, Fort Worth, TX 76102-4060, 
U.S.A, kjanni@brit.org 


SIDA 19(3): 722. 2001 


DIPSACUS FULLONUM (DIPSACACEAE) AND 
VERBESINA WALTERI (ASTERACEAE), NEW TO TEXAS 


Jason R. Singhurst Walter C. Holmes 
Wildlife Diversity Program Department of Biology 
Texas Parks and Wildlife Department Baylor University 
ustin, TX 78704, U.S.A. Waco, TX 76798-7388, U.S.A. 
jason.singhurst@tpwd state. tx.us walter_holmes @baylor.edu 


ABSTRACT 


] reg beet A \ q 


Dipsacus fullonum L. (Dipsacaceae) and Verbesina walte1 


to Texas. 


RESUMEN 


Se citan Dipsacus fullonum L. (Dipsacaceae) y Verbesina walteri Shinners (Asteraceae) nuevas para 
Texas. 


Continued fieldwork and herbarium study conducted by the authors have re- 
sulted in the following records of two species here reported as new to Texas. 
Neither species is included in the standard references treating the vascular 
plants of the state (Correll & Johnson 1970; Hatch et al. 1990; Johnston 1990; 
Jones et al. 1997), nor is the first mentioned species, which is from the area treated 
in Shinners & Mahler’s Illustrated Flora of North Central Texas (Diggs et al. 
1999), included in that work. 


Dipsacus fullonum L. (Dipsacaceae) 

TEXAS. Collin Co.: 3.0 mi W of Anna at jct. of U.S. Hwy 75 and FR 455, 30 Jun 2000, Singhurst 9379 
(BAYLU, TEX). 

Dipsacus fullonum, commonly called Fuller's teasel, is an erect, coarse, prickly 
biennial about 0.5-2(-3) m tall. The flowers are violet and borne in oblong-ovoid 
capitula 2-4 cm tall that are subtended by numerous linear to linear-lanceolate 
prickly involucral bracts 3-10 cm or more long. Gray (1895) mentions that 
Dipsacus fullonum has “awn-like tips of the rigid chaff [paleae] hooked at the 
end, which makes the teasel useful for carding woolen [sic] cloth; cultivated in 
central N.Y. for this purpose ...” Bobrov (1957), however, states that “because of 
the flexibility of the paleae, this plant [D. fullonum] is not used for napping in 
the woolen industry.” In commenting on the complicated nomenclature of the 
species, Bobrov (1957) notes that D. fullonum is a Linnean name that cannot be 
abolished because it is the type species of the genus, consequently the plant, 
“which is of no use to the textile industry, retains the [common] name Fuller’s 


SIDA 19(3): 723 — 725. 2001 


724 BRIT.ORG/SIDA 19(3) 


teasel.” It is probable that the plant referred to by Gray (1895) as D. fullonum is 
actually D. sativas (L.) Honckeny. 

The species is native to the Old World from Atlantic and central Europe to 
Asia Minor and Iran (Bobrov 1957). It has been reported from all of the contigu- 
ous states of the United States except Delaware, Florida, Georgia, Louisiana, 
Maine, Minnesota, North Dakota, and South Carolina (USDA, NRCS 1999). The 
plant normally occurs in waste places, rights-of-way, margins of cultivated 
fields, and similar areas. At the collection locality cited above, two “clumps” of 
the species were found in the immediate area. One consisted of about 20 stems 
originating from the same basal area while the other had the same growth form 
and approximately 35 stems. The occurrence of only two plants suggests that 
the species has only recently colonized the area. 

Dipsacus fullonum is the second species of this Old World family to be in- 
troduced to Texas, the other being Scabiosa atropurpurea L., which isa garden 
escape becoming an invasive weed in north central Texas (Diggs et al. 1999). 
The following key distinguishes the two genera. 


pe 


Plants prickly; heads 2-4 cm long; involucral bracts 3-10 cm long; corollas 4-lobed Dipsacus 
1. Plants not prickly; heads to about 1 cm long; involucral bracts to 1 cm long;corollas 
5-lobed Scabiosa 


Verbesina walteri Shinners (Asteraceae) 


TEXAS. Jasper Co.: Floodplain of Walnut Run Creek near U.S. Hwy 96 ca.2 mi S of jct. with U.S. Hwy. 
190 in Jasper, 23 Sep 2000,Holmes 10984 & tea ee U, TEX). Newton Co.: Big Cow Creek at jct. 
with U.S. Hwy 190,W of Newton, 23 Sep 2000, Holmes 11029 & Si Inghurst (BAYLU,TEX); Big Cow Creek 
at jct. with FR 363, E of Bleakwood, 23 Sep 2000, na 11053 & Singhurst (BAYLU). 


Verbesina walteri is described by Cronquist (1980) as being “much like V. 
alternifolia, but the heads discoid, with white flowers.” Verbesina alternifolia 
has both ray and disc flowers that are yellow. Cronquist gives the distribution 
as South Carolina to Georgia and Louisiana, with outliers in the Piedmont Pla- 
teau of North Carolina and the Ouachita Mountains of Arkansas. In their treat- 
ment of the family for Louisiana, Gandhi and Thomas (1989) considered the 
species to be widespread and abundant east of the Mississippi River but un- 
common in the remainder of the state. The known Texas distribution is basi- 
cally contiguous with the western distribution of the species in Beauregard, 
Vernon, and Sabine parishes, Louisiana, presented by Thomas and Allen (1996). 
Ateach of the locations mentioned in the list of exsiccatae, the species occurred 
in the flat floodplains in two or more “clumps” about 25-100 m or more distant 
from each other. Plants numbered from few (2-5) to 25 or more in each clump. 


ACKNOWLEDGMENT 


We are grateful to John Pruski (US) for review of the manuscript. 


SINGHURST AND HOLMES, NEW RECORDS FOR TEXAS 725 


REFERENCES 


Bosrov, E.G. 1957. Dipsacaceae. In:B.K. Shishkin, ed., Flora of the U.S.S.R. vol. 24, Dipsacaceae, 
Cucurbitaceae, Campanulaceae, Izdarel’stvo Akademii Naulk SSR, Moskva-Leningrad. 
[Translated from Russian in 1972 by J. Lorch and U. Plitman, Keter Press, Jerusalem, 
Israel.] 

Correll, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner. 

Cronauist, A. 1980. Vascular flora of southeastern United States. Vol. 1. Asteraceae. Univer- 
sity of North Carolina Press, Chapel Hill. 

Diccs, G.M,JR., B.L. Liescome, and R.J.O’Kennon. 1999. Shinners & Mahler’s illustrated flora of 
north central Texas. Botanical Research Institute of Texas, Fort Worth. 

GANDHI, K.N. and R.D. THomas. 1989. Asteraceae of Louisiana. Sida, Bot. Misc. 4:xii + 202. 

Gray, A. 1895. Gray’s field, forest, and garden botany. American Book Company, New York. 

Hatcu, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas 
Agric. Exp. Sta. Publ. MP-1655, Texas A&M University, College Station. 

Joxnston, M.C. 1990. The vascular plants of Texas. A list, up-dating the manual of the vas- 
cular plants of Texas. Published by the author, Austin. 

Jones, 8.D., J.K. Wipre, and PM. Montcomery. 1997. Vascular plants of Texas: a comprehensive 
checklist including synonymy, bibliography, and index. University of Texas Press, Austin. 

THomas, R.D.and C.M. Alten. 1996. Atlas of the vascular plants of Louisiana. Vol 2. Dicotyle- 
dons. Acanthaceae-Euphorbiaceae. Louisiana Department of Wildlife and Fisheries, 


Baton Rouge. 
Uspa, Nacs. 1999. The PLANTS database (http://plants.usda.gov/plants). National Plant Data 
Center, Baton Rouge, LA. 


726 BRIT.ORG/SIDA 19(3) 


Book REVIEW 
FRANKLIN W. MartIN, RUTH M. RuBerte, and Laura S. MEItzNer. 1998. Edible Leaves 
of the Tropics. Third Edition. (ISBN 0-9653360-1-8, pbk.). ECHO, 17430 
Durrance Road, North Fort Myers, FL 33917, U.S.A. $14.95, 194 pp, baw 
plates, 6" x 9". 


This third edition of Edible Leaves of the Tropics puts back in print this classic that has been unavail- 
able for 15 years. Published by the Educational Concerns for Hunger Organization, Inc. or ECHO, this 
text is a primary resource used by the staff in answering questions of development workers in the 
tropics and subtropics. This edition updates the 1981 edition with new information from scientific 
publications and the insights of the ECHO staff and at the same time retains much of the informa- 
tion and a from Martins original. 

es are a high quality food in the tropics and “developing” nations, not only because of their 
act value but also the little effort required in their production. This text describes tropical 
plants with edible leaves, discussing their eee onan habit, cultivation guidelines, nutritional 
value, preparation, multiple uses, and cautions. The boo ivided into ten chapters, follow 
list of seed sources and resources, a hice iy an as on tropical plants with edible leaves 


(over 1500 species), and an index to genera and common names. Chapters one through ten cover 89 
pages and discuss the place of green leaves in the diet, the principal green leaf herbs of the tropics, 
common weeds with edible leaves, temperate zone green leaves, lettuce, poisonous leaves, and cul- 
ture and care of green-leafed vegetables. Following chapter ten are 56 pages of “Illustrations.” How- 
ever there are no — but black and iiaite paolograpes of plants, the quality e we ae 


from poor to goo 
hardly visible. The appt on ae pine with edible leaves is this oe strong point and de- 
fines it as a “source” book. This appendix, six pages in length, covers over 1500 species, their 
latin name, source «Cons Tropica es ae etc.), and type (shrub, herb, vine). It would be use- 
ful for the common names to be included in this table for future esullons. 

Edible L aecoithe Tropics is a useful book for anyone i tropical botany, gardening, 
and the nutrition of i iets Ne necessar uy apprepuate tor classes, hiss is primarily a source 
book a ould y tinue to justify its existence. It 
is well written and researched and available for a modest price. | “would recommend this book for 


Z 


anyone interested in \ finding an inventory of edible tropical plants for either their own research o 
consumpti ion.—Kevin D. Janni, Botanical Research Institute of Texas, Fort Worth, TX ie. ae 
iim} 


oO 


SIDA 19(3): 726. 2001 


CAREX ABSCONDITA (CYPERACEAE: CAREYANAE) IN 
LOUISIANA 


David J.Rosen 
U.S. Army Corps ees 
Galvecton NMictri t, PO. 1229 
Galveston, TX 77553, ‘i 


During fieldwork southeast of Baton Rouge, Louisiana, I collected a Carex, Sect. 
Careyanae in alluvial mixed woods overlooking the Amite River. A tentative 
identification of Carex abscondita Mackenzie was made. A duplicate specimen 
(Rosen 1032) was sent to Stanley Jones (BRCH), who confirmed that the sedge 
was indeed C. abscondita, and provided information on his own collection of C. 
abscondita from Louisiana (Jones 13392). In a subsequent review of Thomas and 
Allen’s Atlas of the vascular flora of Louisiana (1993), C. abscondita was listed 
under excluded species. This paper confirms the occurrence of C. abscondita in 
Louisiana’s flora based on field collections and herbarium specimens. 

Ina revision of Carex, Sect. Laxiflorae, Bryson (1980) cites collections of C. 
abscondita from Orleans (GH), St. Tammany (US), and Washington (TENN, 
VDB) parishes. According to Jones (1999), the correct section is Sect. Careyanae. 
Queries to several herbaria (LSU, MCN, MICH, and NO) revealed additional col- 
lections of C. abscondita from Louisiana. Carex abscondita is a sciophyte of mesic 
to wet alluvial woodlands, often overlooked due to its well-concealed inflores- 
cences hidden among the leaf bases. This could explain the scarcity of collec- 
tions from Louisiana. Sufficient collections reported herewith lessen the gap in 
distribution (Fig. 1) from the specimens cited by Bryson (1980) for eastern Loui- 
siana, to Texas where C. abscondita is also a recent addition (Naczi & Bryson 
1990; Jones et al. 1997). Additional fieldwork and herbarium | needed 
to complete the distribution of C. abscondita in Louisiana. 


Specimen citations: LOUISIANA. Calcasieu Parish: Woods adjacent to Camp WI-Ta-Wenton in Sect. 
2,785, R7W, 1 Apr 1998, Neyland 1726 (MCN). Catahoula Parish: Catahoula Ranger District of the 
Kisatchie National Forest, Georgetown vicinity, about 6 mi W, floodplain of Indian Creek, 20 Apr 
1996, Hyatt 6960 (LSU). Claiborne Parish: Corney fire tower vicinity, near Corney Lake on LA Hwy.9, 
common, mainly in floodplains along streams, 17 Apr 1998, Hyatt 8003 (MICH). East Baton Rouge 
Parish: Behind residence at 19452 Indian Ridge Avenue in SE Baton Rouge, frequent in sloping 
woodland above Amite River, 13 Apr 2000, Rosen 1032 (BRCH, NO). Grant Parish: Packton, 1.0 mi S 
of US Hwy. 167 in Sect. 4, T9N, R2W, in floodplain woods on sandy soil, locally Gofninane on Ores! 
floor, 19 May 1995, Hyatt 6434 (MICH); Catahoula Ranger District of the Kisatchie Natio 

Creek near what is called and reported to be a Spanish Mound at the end of the ead into aGe 
Spanish Mound, 18 Apr 1996, Hyatt 6958 (LSU). Natchitoches Parish: Winn Ranger District of the 
Kisatchie National Forest; Ashland vicinity, about 3 air mi SW, mature floodplain deciduous forest; 
common along medium to large streams, 14 May 1996, Hyatt 7066 (LSU). Rapides Parish: Evangeline 


SIDA 19(3): 727 — 729, 2001 


728 BRIT.ORG/SIDA 19(3) 


Laren] eer 


/109Nn\ YL (+ \ ISU by 
yy 7 


4 nce cee ; i Ao ee ae ' 
closed circles (@), MCN by closed diamonds ( # ), MICH by open diamonds ( © ), and NO by open circles (©). 


Ranger District of the Kisatchie National Forest; Alexandria Vicinity; about 8 mi WSW; Timber Com- 
partments 14/78's north edge, S bank of Bayou Beouf vicinity,07 May 1996, Hyatt 7052 (LSU); Brushy 
Creek Riparian site, Sect. 11,T2N, R3W, common, local in floodplains and adjoining slopes, 23 May 
1998, Hyatt 7912 (MICH); Kisatchie National Forest;0.7 mi SW on forest rd. 188 from its jct.with forest 
rd. 155; then 0.7 mi SW on a logging rd. to Camp Pond Creek bottomland, NW of Williana; bottom- 
land forest with a few pines, lower slopes, occasional, 22 April 1998, Jones 13392 (SAT). West Feliciana 
Parish: Ravines in woods near St. Francisville, 14 Apr 1936, Penfound s.n. (NO); Along W side of Rt.61, 
just S of St. Francis Hotel, 0.2 mi S of jct. of Rts.61 & 10, common, 23 May 1994, Naczi 3988 (MICH); W 
side of US Rt.61,S of Wakefield near fire tower, Sect. 2,T1S, R2W, 20 April 1998, Hill 30011 (MICH). 


a 


— 


ACKNOWLEDGMENTS 
Thanks to Stanley D. Jones (BRCH), Roland P. Roberts (LSU), Ray Neyland 
(MCN), Tony Reznicek (MICH), and Anne S. Bradburn (NO) for their assistance. 
lam also extremely grateful to Robert EC. Naczi and Charles T. Bryson for their 
illuminating review of this manuscript. 


ROSEN, CAREX ABSCONDITA IN LOUISIANA 729 


REFERENCES 

Bryson, C.T. 1980. A revision of the North American Carex section Laxiflorae (Cyperaceae). 
Ph.D. dissertation, Mississippi State Univ., Starksville 

Jones, S.D. 1999. Carex. In. Shinners and Mahler's illustrated flora of North Central Texas. 
G.M. Diggs, Jr, B.L. Lipscomb, and RJ. O’Kennon. Botanical Research Institute of Texas, 
Fort Worth. 

Jones, S.D., J.K. Wiprr, and PRM. Montcomery. 1997. Vascular plants of Texas: A comprehensive 
checklist including synonymy, bibliography, and index. University of Texas Press, Austin. 

Naczi, R.F.C. and C.T. Bryson. 1990. Noteworthy records of Carex (Cyperaceae) from the 
southeastern United States. Bartonia 56:49-58. 

THomas, R.D. and C.M. Atten. 1993. Atlas of the vascular flora of Louisiana, Volume |: fern 
allies, conifers, and monocotyledons. The Natural Heritage Program, Wildlife Div., Loui- 
siana Dept. of Wildlife & Fisheries, Baton Rouge. 


730 BRIT.ORG/SIDA 19(3) 


Book REVIEW 


PAUL Martin, David YETMAN, MARK FIsHBEIN, PHIL JENKINS, THOMAS VAN DEVENDER, and 
REBECCA WILSON (eds.). 1998. Gentry’s Rio Mayo Plants: The Tropical Decidu- 
ous Forest & Environs of Northwest Mexico. (ISBN 0-8165-1726-6 hbk.). 
University of Arizona Press, 330 S. Toole Ave., Suite 200, Tucson, AZ, 85701- 
1814. (Orders: www.uapress.arizona.edu, 520-621-1441, 520-621-8899 fax). 
$75.00, 558 pp, 2 maps, photos, index, 7" x 10" 

The interface of Neotropical and Sonoran Desert ecozones has been a region of great interest to bota- 

nists and biologists. Howard Scott Gentry was drawn to it over 60 years ago, specifically the Rio 

Mayo region of northwestern MEME) and in 1942 he published the first version . this ae new 

version of the classic text uy and amends major portions of the orig S 

the original list of taxa. This was undertaken with the ea and ee tof Gentry until his death in 

1993. Gentry also authored a number of books on Aga rth America, Sonora, and 

Baja, California during his lifetime. The list of almost 3,000 taxa isaecompanied by a new historical 
dee ease a review of geography and vegetation, ch nd and rivers asa result of agricul- 
and lumbering, and an nea on i6eal ae and use of aes 

The oe is divided into four parts, covering: the Rio Mayo at the end of the ty 

he original, and an annotated list of Rio Mayo Vasco 


rentiet Sepytrar 


a list of Rio Mayo localities, excerpts from t 
plants. Part one offers an overview of the region and a contemporary overlook of the regional vegeta- 
tion. Part two isa list of see and an nae of the list. Part three covers oo (rivers, 
mountains, climate, etc) and veg , plant habitats, successions in rl as). Part 
four is the annotated et ist (345 pages) preceded - an explanation of the format. 

F names, local distributions, habitat, eer information, and indigenous 
uses are provided. There are no English common names included in this list, but some plants have 
the loc al Rio mae name. era two comments concerning this editorial decision: |. the local names 
are not sist j their existence and 2. why leave out the English common names??? 
Recognized as eee in several parts of Latin America, the taxa Sonchus asper (L.) Hill and 
Sonchus oleraceus L. are listed without a Rio Mayo name, I found this surprising. Three to five local 
participants could probably accomplish ane by eh nai a aa afternoons ina local herbarium re- 
cording local names. T English pl ould offer an interesting 


— perspective and possibly open the book up to a wider audience. iotleed Latin names are of 


a les ESsaiends to most iy eed His nee in ae section in ear atlas s ce be pleasing to 
I tiv 
and wild plants, the latte . ludi d checklist of useful fruits, seeds, roots and herbage, 
and construction, fuel, and cee aacoue uses. 
hese sections are followed by Gentry’s bibli literature cited, and an index. There are two 
maps jacluded’ one being a large removable vas map from the back cover. This isa pleasing f eature 
' the book | Hl utilit sefulness. However, tl 
review copy we received here at BRIT advertises a sees oe a Thiss same advertisement 
appears on the press w books/bid1204.htm. A color map would b 
a great addition, eee ee to errehasize biodiversity or biogeography, and also ia 
the price. 


aes 


Overall, this is a beautiful book. It is well designed, academically and aesthetically. Moreover, 
his is a major contribution to our so ane Tlogeographical literature. The authors should be 
commended for their efforts. F biogeography, conservation, and eth- 
nobotany of nort ae as should find this ees oie Bc D.Janni, Botanical Recearch In- 
stitute of Texas, Fort Worth, TX 76102-4060, U.S.A 
SIDA 19(3): 730. 2001 


PHYLLOSTACHYS BAMBUSOIDES (POACEAE: BAMBUSEAE) 
PREVIOUSLY UNREPORTED FROM LOUISIANA 


David J.Rosen Stanley D. Jones 
U.S. Army Corps eae Botanical Research Center 
Galveston District, PO. Box 1229 PO. Box 6717 
Galveston, TX 77553, on Bryan, TX 77805-6717, U.S.A. 


Joseph K. Wipft 
Pure Seed Testing, Inc. 
PO. Box 449 
Hubbard, OR 97032, U.S.A. 


ABSTRACT 


Phyllostachys bam busoides, native of China, is reported from the floodplain of Thompson Creek, West 
Feliciana Parish, Louisiana. A key to Louisiana Phyllostachys and a description of the genus and spe- 


cies are included. 
RESUMEN 


Phyllostachys bambusoides, nativo de C me, se cita del llano inundado de Thompson Creek, West 
Feliciana Parish, Louisiana. $ para los Phyllostachys de Louisiana y una descripcion 


de P. bambusoides. 


Phyllostachys bambusoides Sieb. & Zucc.,a native of China (McClure 1957), has 
been found growing outside cultivation in West Feliciana Parish. Hitchcock 
(1950) describes P bambusoidesas an exotic introduced through cultivation and 
persisting in areas with mild winter temperatures, but listed no distribution. 
Phyllostachys bambusoides has previously been reported to occur in cultiva- 
tion for Texas Jones et al. 1997), but has not been reported from Louisiana (Allen 
1992: Thomas & Allen 1993). 


Phyllostachys Sieb. & Zucc. 

Perennial with extensive rhizomes. Culms straight and stiffly erect; thick 
hallow, glabrous, and D-shaped in cross section; culms branching freely from 
the middle and upper nodes, with usually 2 unequal branches at each node, 
sometimes a third smaller one. Culm sheaths deciduous and subcoriaceous. 
Leaves 1-7 per branchlet; sheaths ciliate along one side; ligules a short ciliolate 
membrane; blades flat, tessellate, lanceolate, abruptly acuminate, base round 
and connected to a short flattened pseudopetiole. Pseudospikelets numerous, 
often with small rigid bracts at their bases, spikelet concealed by inflated ex- 
ternal, coriaceous bract; bract bearing reduced leaf blade, glabrous but with 


SIDA 19(3): 731 — 734. 2001 


732 BRIT.ORG/SIDA 19(3) 


ciliate margin. Spikelet florets 1-2; glumes 2(1), lanceolate; lemmas similar to 
glumes; palea bilobed at apex. 

e characteristics most generally useful in the identification of bamboo 
species (once the genus is known) are found in the culm sheaths that clothe the 
young shoot. These characteristics reveal themselves most vividly while the 
sheaths are in a fresh state. As they dry, their delicate, often very distinctive 
colors fade to light or dark straw, though any spots of dark pigment usually 
persist. The tissues shrink and sometimes become more or less warped, and 
some of the delicate parts, such as the auricles and the ligule, become brittle 
and easily broken when dry.” (McClure 1957). When collecting culm sheaths, 
Bay nome sales bea along the length of the culm. Phyllostachys 
rom P. aured using the following key modi- 
fied from McClure (1957) This key must be used in the spring when the young 
shoots are actively growing and the culm sheaths are still present. 


A KEY TO PHYLLOSTACHYS IN LOUISIANA 


Auricles and well soa oe) setae present on the upper culm sheaths es me 


ermost culm she ally without auricles); culms 6-22 m tall; base of culm 
m in diameter. eee eee Sieb. & Zucc. 
Auricles and oral setae absent on the upper Cl Il cheathe (ri ted Ree r); 
culms 3-10 m tall; base of culm 1-5 cm in diameter. hys aurea 


E. ea ex A, Riv. & C. Riv. 


Phyllostachys aurea FE. Carr. ex A. Riv. &C. Riv. 

Culms 3-10 m, straight and stiffly erect; base of culms 1-5 cm in diameter; 
thick hollow, glabrous, D-shaped in cross section; green, visibly to strongly glau- 
cous soon after sheath fall, becoming green to yellowish green when mature: 
culms branching freely from the middle and upper nodes, usually with 2 un- 
equal pranchies at each node, sometimes a third branch also present. Culm 
sheathsd , subocoriaceous, pale olive green to pale rosy buff, with bur- 
gundy or pale-green veins, rather sparsely strewn with small brown spots, not 
glaucous, glabrous except for a line of minute white hairs along the base. Au- 
ricles and oral setae absent. Ligules short, apex slightly convex or rarely slightly 
concave and ciliate on the margin in the lower several sheaths; slightly longer 
and long ciliate to fimbriate on the margin in mid-culms sheaths. Culm sheath 
blades subulate to lanceolate, more or less crinkled below, becoming long, nar- 
rowly strap-shaped and pendulous above. Leaves leaf sheaths 2.5-3.5 cm long, 
ciliate along one side, with auricles and oral setae well-developed (occasionally 
absent), conspicuous in spring when leaves are young; ligules a short ciliolate 
membrane; leaf blades flat, tessellate, 4-12 cm long, 5-16 mm wide, lanceolate, 
abruptly acuminate, base round to a short flattened pseudopetiole, 
pseudopetiole 2-4 mm ra Bein or eraescent tieiets scabrous; 1-7 
leaves per branchlet. P asc g overlapping, 35-50 


ROSEN ET AL 733 


cm long, often with small rigid bracts at their bases, spikelet concealed by in- 
flated external, coriaceous, bract ca. 22 mm long, bearing reduced leaf blade, 
glabrous, ciliate margins. Spikelets l-several in each pseudospikelet. Spikelet 
florets 1-2; glumes 2(-L), lanceolate; lemmas similar to glumes, ca. 19 mm long, 
glabrous with a stiff awn tip, palea ca. 15 mm long, glabrous, bilobed at apex, 
not keeled. Native to southern China. 


Phyllostachys bambusoides Sieb.& Zucc. 

Culms 6-22 m, straight and stiffly erect; base of culms 5-15 cm in diam- 
eter; thick hollow, glabrous, D-shaped in cross section; not glaucous at culm 
sheath fall, green, golden yellow or golden yellow with a variable amount of 
green striping; culms branching freely from the middle and upper nodes, usu- 
ally with 2 unequal branches at each node, sometimes a third branch also 
present. Culm sheaths deciduous, subocoriaceous, greenish to ruddy buff, more 
or less densely spotted throughout with dark brown spots; the lowest several 
culm sheaths usually without auricles and oral setae, but the rest bearing two 
(rarely one) auricles; auricles conspicuous, narrow to broadly ovate or falcate 
(more or less staghorn in shape), conspicuously fringed with several greenish 
crinkled bristles (oral setae). Ligules well-developed, the apex more or less 
strongly convex with ciliolate margins in smaller culms to truncate and ciliate 
with coarse bristles in older culms. Culm sheath blades short, lanceolate, re- 
flexed, and crinkled in lower sheaths to strap shaped and recurved in the upper 
ones; green or colored with burgundy, green and cream stripes to 9 cm long. 
Leaves 1-7 leaves per branchlet. Leaf sheaths 5.2-25 mm long, ciliate along one 
side, auricles, when present, conspicuous, with oral setae well developed (occa- 
sionally absent), oral setae conspicuous in spring when leaves are young; ligules 
convex, a short ciliolate membrane; glabrous, ciliate along one margin; leaf 
blades flat, tessellate, 10-15 cm long, 15-20 mm wide, lanceolate, abruptly 
acuminate, base round to a short flattened pseudopetiole; adaxially glabrous, 
abaxially glaucous, densely spinulous scabrous on the basal portion of mid- 
vein, scabrous along one margin of blade. Pseudopetiole 3-5 mm long. 
Pseudospikelets numerous, l-several per node on leafy or leafless branchlets; 
ascending and overlapping, 40-80 cm long, often with small rigid bracts at their 
bases, spikelet concealed by inflated external, coriaceous, bract 10-40 mm long, 
bearing reduced leaf blade. Spikelets 2-5 in each pseudospikelet. Spikelet flo- 
rets 1-2; glumes 2, lanceolate ca. 17 mm long; lemmas similar to glumes, 18-22 
mm long, densely pubescent, ciliate margins, palea 17-20 mm long, bilobed at 
apex, keeled, 3-veins between the keel, 3-5 veins along margin, keels pubes- 
cent. Native to Japan and mainland China, extending westwards to the Hima- 
layan Mountains. 

In Louisiana, Phyllostachys bambusoides was observed growing aggressively 
in sandy, alluvial soils, out of cultivation, among riparian hardwoods, and covering 


734 BRIT.ORG/SIDA 19(3) 


an area of ca. 0.5 ha. A random sampling (n=9) of culm DBH yielded a mean of 
6.6 cm ranging to 12.25 cm with an estimated mean culm height of 9 m. 
Voucher sisal tk LOUISIANA. West Feliciana Parish: W of the town of Jackson, N of Hwy. 10 on 
the west bank a ae. in deep se sands with Plata identalis, Betula nigra, 
Quercus phellos, Salix nigra,a 30 Apr 2000, Rosen 1098, det. .K. Wipff(BRCH, jkw- 
pers. herb., NLU, SAT, and a 


REFERENCES 

Atten, C.M. 1992. Grasses of Louisiana, 2nd ed. Cajun Prairie Habitat Preservation Society, 
Eunice, LA. 

HitcHcock, A.S. 1950. Manual of the grasses of the United States. U.S. Dept. Agric., Washington. 

Jones, S.D., J.K. WirrF, and P.M. Montcomery. 1997.Vascular plants of Texas: A comprehensive 
checklist including synonymy, bibliography, and index. University of Texas Press, Austin. 

McCiure, F.A. 1957. Bamboos of the genus Phyllostachys under cultivation in the United 
States. U.S. Dept. of Agric. Handb. 114, Washington, D.C 

THomas, R.D. and C.M. Aten. 1993. Atlas of the vascular flora of Louisiana, Volume |: fern 
allies, conifers, and monocotyledons. The Natural Heritage Program, Wildlife Div., Loui- 
siana Dept. of Wildlife & Fisheries, Baton Rouge. 


A SCUTELLARIA (LAMIACEAE) NEW TO 
NORTH CAROLINAAND A KEY TO THE 
SMALL-FLOWERED CAROLINA CONGENERS 


Alexander Krings Joseph C. Neal 
Herbarium Department of Botany Department of Horticultural Science 
North Carolina State University North Carolina State University 
Raleigh, NC 27695-7612, U.S.A. Ralei 2 NC 27695-7609, U.S.A. 
Alexander_Krings@ncsu.edu e_Neal@ncsu.edu 


Native to South America, Scutellaria racemosa Pers. (South American skullcap) 
has been collected sporadically in the southeastern United States but has not 
been reported for North Carolina by Radford et al. (1968) or Kral (1981). Listed 
as an obligate wetland species by Reed (1988), S. racemosa has primarily been 
found in coastal plain communities and was previously only known from Texas, 
Louisiana, Alabama, Florida, Georgia, and South Carolina (Godfrey & Wooten 
1981; Kral 1981; Allen 1983: Jones & Coile 1988; Tobe et al. 1998; Wunderlin & 
Hansen 2000). During field work, Joe Neal discovered a population of an un- 
known weed on the grounds of a nursery in Chatham County, in the lower Pied- 
mont of North Carolina. Live plants, collected from the site on 4 Dec 1998, were 
grown out in a greenhouse at North Carolina State University and identified as 
Scutellaria racemosa based on fertile material. Representative material of this 
collection was prepared for deposit at NCSC in Dec 2000. A subsequent site 
visit, also in Dec 2000, yielded additional voucher material (see voucher speci- 
mens below). Based on observations by Neal, the population has been estab- 
lished since at least 1996. Plants were found spreading at the edges of nursery 
ground cloths and greenhouses, as well as around a nearby pond. Soils appear 
to be persistently moist to wet, due to regular irrigation. A search of nearby 
forests revealed that the population has started encroaching in paca areas 
only within reach of the irrigation system. Voucher speci deposited at 
BHO, NCSC, and USCH. 

Voucher specimens: U.S.A. North Carolina. Chatham Co.: Specimen from potted plant collected 4 
Dec 1998 from 2925 NC Hwy 751 and grown in North Carolina State University greenhouse by Jo- 
seph Neal, 1 Dec 2000, Krings & Neal 365 (NCSC), 2925 NC Hwy 751, field collection from nursery, 
specifically from fertilized pots of shrubs kept in enclosed greenhouse space and receiving regular 
irrigation, 6 Dec 2000, Krings & Neal 366 (BHO, NCSC, USCH) 


Scutellaria racemosa Pers. (Fig. 1) can be distinguished from its southeastern 
congenerics by the hastate leaves (Fig. 2, C). In the Carolinas, it is one of only 
four species exhibiting corollas 7 mm long or less (see key below). Worldwide, 
itis apparently one of only two skullcaps displaying consistently hastate leaves 


SIDA 19(3): 735 — 739. 2001 


736 BRIT.ORG/SIDA 19(3) 


(Paton 1990). The other species, S. hastifolia L., is native to W Europe and exhib- 
its an erect habit, flowers greater than 2 cm long, and brown nutlets with a 
prominent median band. Scutellaria racemosa on the other hand, exhibits a 
trailing to weakly ascending habit, flowers typically 4 mm or less long, and 
brown nutlets lacking a median band (Fig. 1; description). 


KEY TO SMALL-~FLOWERED SCUTELLARIA 
(COROLLAS S 7 MM LONG) IN THE CAROLINAS 


1. Larger leaves hastate, 1.3-3.2 cm long; corollas white to lavender or purple, 2-4 
m long. S. racemosa Pers. 
1. Larger leaves not hastate, 0.6—12 cm long; corollas white to lavender or violet, 3-7 
mm lon 
2. Petioles > 4mm long;leaves 3-12 cm long, the api ypically a fl 
in axillary racemes, the bracts much reduced. S. laterifolia L. 
2. Petioles < 3 mm long; leaves = 3 cm long, the apices acute to blunt; flowers 
solitary in the axils, or somewhat racemose, the bracts resembling stem leaves 
and only somewhat reduced 


3. Principal leaves ovate, rounded to slightly cordate, = 2.5 cm long. S.nervosa Pursh 
3. Principal leaves ovate to lance-ovate, rounded to slightly cordate or not, < 
cm long or less S. parvula Michx. 


SYNONYMY AND SPECIES DESCRIPTION 
iia racemosa Pers., Syn. Pl. 2:136. 1807. (Fig. 1). Type: collected near 
ontevideo, Paraguay Commerson s.n. (HOLOTYPE: P). 
Scutellaria bonariensis Willd. ex Benth., Linnaea 11:345. 1837. 
Scutellaria hastata Larranaga, Escritos de Don Damaso Antonio Larrafaga 2:87. 1923. 
Scutellaria heterophylla Willd. ex Benth., Linnaea 11:345, 1837. 
Scutellaria rojasii Briq., Bull. de l'Herbier Boissier, sér. 2, 7:600. 1907. 
Scutellaria rumicifolia Kunth, Nov. Gen. Sp. 2:324. 1817. 


Colonial, low herbaceous perennials with slender rhizomes; stems 4-angulate, 
glabrous to puberulent, trailing to weakly ascending (to 1 ft.), typically several 
from base; leaves opposite, 3-5 veined from base, glabrous or minutely puberu- 
lent on both surfaces, the uppermost typically narrowly- lanceolate or deltate 
and toothed or lobed basally, 1-4 mm broad, 5-12 mm long, the lower narrowly 
to broadly hastate, 5-20 mm broad, 13-32 mm long, apices retuse or, less fre- 
quently, rounded; flowers solitary, axillary, pedicels 2-4 mm long, puberulent; 
calyces campanulate, puberulent to glabrate with age, green to purplish, with 
a protul n the distal surface (scutellum), 1.8-2 mm long, enlarging to 3 
mum long following anthesis and splitting horizontally; corolla white to laven- 
der or purple, 2-4 mm long, 2-lipped, the upper concave, hoodlike, the lower 
spreading, purple or pink spotted; stamens 4; nutlets brown, obovoid, lacking 
median band, ca. 0.5 mm in diam. 


KRINGS AND NEAL, SCUTELLARIA NEW TO NORTH CAROLINA 737 


Pers. (Lamiaceae): A. Habit; B. i-ii. Leaf variation; C. Flower; D. Calyx; E. Dehisced calyx and 
be A-B based on Krings 371 (NCSC). C based on Radford 46268 (FLAS). D—E based on Godfrey 73437 (FLAS). 


ACKNOWLEDGMENTS 


We thank Guy Nesom and Bruce Sorrie for their review and thoughtful com- 
ments on the manuscript. 


BRIT.ORG/SIDA 19(3) 


738 
3.2 cm 
| 
4.7 cm 
1.2 cm 
3.2 cm 
1.2 cm 
a. 2. Comparative leaf Hanae de of the four Carolina species of Scutell hibiting Il | 
Suter. M.E. Wharton 33317 (NCSO); ii: S ifolia L., W.H. D 23451 (NCSC . Scutellaria 


rsh., arena ae 1016 (NCSC). ¢ i: auivane racemosa a Tiles & Ned 366 (NCSC); ii: 
Sap racemosa a. Palacios-Cuezzo 497 (NCSC). D 


REFERENCES 

Auten, C. 1983. Murdannia keisak (Hassk.) Hand.-Mazz. (Commelinaceae), Bothriochloa 
hybrida (Gould) Gould (Poaceae), and Scutellaria racemosa Pers. (Lamiaceae) new to 
Louisiana. Sida 10:189-190 

Goprrey,R.K. and J.W.Wooten. 1981.Aquatic and wetland plants of the southeastern United 
States: Dicotyledons. University of Georgia Press, Athens. 

Jones, S.B., Jr.and N.C. Coie. 1988. The distribution of the vascular flora of Georgia. Depart- 
ment of Botany, University of Georgia, Athens. 

Kral, R. 1981. Some distributional reports of weedy or naturalized foreign species of 
vascular plants for the southern states, particularly Alabama and middle Tennessee. 
Castanea 46:334-339. 


KRINGS AND NEAL, SCUTELLARIA NEW TO NORTH CAROLINA 739 


Paton, A. 1990. A global taxonomic investigation of Scutellaria (Labiatae). Kew Bull. 45: 
399-450. 

Raproro, A.E., H.E. Axes, and C.R. Beit. 1968. Manual of the vascular flora of the Carolinas. 
The University of North Carolina Press, Chapel Hill. 

Reeo, PB., Jr. 1988. National list of plant species that occur in wetlands: national summary. 
U.S. Fisheries and Wildlife Service Biol. Rep. 88 (24):125. 

Tore, J.D., K.C. Burks, R.W. Cantrett, M.A. Gartano, M.E. Sweeney, D.W. HALL, P. WALLACE, G. ANGLIN, 
G. Newson, J.R. Cooper, D. Bickner, K. Gitgert, N. AYMOND, K. GREENWOOD, and N. RayMonb. 1998. 
Florida wetland plants: an identification manual. Florida Department of Environmen- 
tal Protection, Tallahassee. 

Wunoertin, R.P. and B.F. Hansen. 2000. Atlas of Florida vascular plants (http:// 
www. plantatlas.usf.edu/). [S.M.Landry and K.N. Campbell (application development), 
Florida Center for Community Design and Research.] Institute for Systematic Botany, 
University of South Florida, Tampa. 


740 BRIT.ORG/SIDA 19(3) 


Book REVIEW 


Rosert H. Ropicuaux and Davin A. YETMAN (eds.). 2000. The Tropical Deciduous 
Forest of Alamos: Biodiversity of a Threatened Ecosystem in Mexico. (ISBN 
0-8165-1922-6, hbk.) The University of Arizona Press. 355 S. Euclid Ave, Suite 
103, Tucson, AZ 85719. (Orders: www.uapress.arizona.edu, 520-621-1441, 
520-621-8899 fax). $50.00, 260 pp, 19 halftones, 5 line illustrations, 4 maps, 


61/8" x 91/4". 
Hopes orests follow a phytogeogr aphic and eeeeca continuum of wet to dry. In wet climates 
lly taller trees with medium-| ndry climates trees are usually shorter 
with siialler ibaves The tropical deciduous forest of Alamos M here in the middle of 


this gradient. Most people, those of us in academia not withstanding, do not picture this part of North 


America as being tropical, but nonetheless, “there are more than 100 ae of tropical trees in 36 
families, 48 species of orchids, 6 species of parrots, mountain lions, jaguars, and according to local 
legend, a big cat called t peaues® Pgs. The Tropical Deciduous Forest of Alamos: es ae Threat- 
ened Ecosystem in Mexi f artic 


OQ. 


cles covering aspects of botany, ecology, ethnobotany, 
and even the diversity and distribution of reptiles, amphibians, ee and birds in the region. 
This volume is a satisfying continuation of Gentry’s Rio Mayo Plants (h tal 1998: Reviewed 


this issue: pg 730) also published by the University of Arizona Press. 

The book comes in seven chapters, followed by an index. The introductory chapter discusses 
the ecosystem and biodiversity of the region and the human use of these resources. In this chapter, 
the authors not only give highway and road directions to the area (only a day’s drive south of the 
Mexico-U.S. border) but also ater a ve cecomnnencaltons on soliciting plant knowledge from the 
local people. (Dispenseme. C 5 ?). Chapter two compares the structure and func- 
tioning of the tropical deciduous forest of Alginos with the other areas of western Mexico. Chapter 
three analyzes the vegetation and flora of the tropical deciduous forest, specifically the Rio 
Cuchujaqui. Included in this chapter is an annotated checklist of the river area that illustrates well 
the diversity of tree species in the oS In chapter four, the authors examine the tree species of the 
forest that are used by the Mayo people of Mexico. The Mayo are linguistic brethren of the Yaqui of 
western Sonora. The Mayo and others a developed several varieties of domesticated plants that 


— 
_— 


flourish in the area. Two appendices close chapter four, one ethan: ethnobotanically useful trees 
and columnar cacti of the Mayo region and the other providesa list of the trees in the Mayo region by 

scientific, Mayo, and Spanish names. Chapter five discusses further the crop diversity among the 
ees nous farming cultures and includes an appendi ees ees names for native wild and do- 


esticated crops of the northern Sierra Madre Occidental mae pte examines the diversity and 
- 


ree butions of amphibians, reptiles - ter seven sravidesan com- 
I 
saath Cae the bide of the Alamos area, tes oe distributions and sea- 
nal beha 
na 


setae with the majority of books devoted to biodiversity conservation, this volume has a 


remem oy (ie. on ene! to it. The people of the region are dependent on several plant 
g this aspect makes the need for conservation even more compel- 


ie The editors of this ee and a ee of each chapter have done a fine job in broadly 


outlining this region and elucidating the need for biocultural conservation. I enthusiastically rec- 
ommend this book to anyone interested in the botany, ecology, ethnobotany, and conservation of 
this region in niet Enwees cae —Kevin D. Janni, Botanical Research Institute of Texas, Fort Worth, 
TX 76102-4060, U.S.A. kjanni@ 


SIDA 19(3): 740. 2001 


LESPEDEZA CUNEATA (FABACEAE), A FIRST RECORD 
OF ITS OCCURRENCE IN MEXICO 


Eduardo Estrada C. and Carmen Yen M. 
Facultad de Ciencias Forestales 
LInivercidad Arita do N/ f] 


P41,67700, Linares, Nuevo Leé6n, MEXICO 


ABSTRACT 


Lespedeza cuneata (Dum.Cours.) G. Don, a native species from eastern Asia and introduced in the 
U.S.A.,, is reported for a time to nee Receu por) anical me Se ections ip petween Cusunienes 
and La Junta, in Chihua 


its previously known range, in eastern Texas. 


RESUMEN 


Lespedeza cuneata (Dum.Cours) G. Don, una especie del este de Asia e introd 


se cita por primera vez para México. Colecciones recientes de material botanico entre Cuauhtémoc y 
La Junta, en el estado nortefio de Chihuahua, México, permiten extender su rango de distribucion 


cerca de 800 km al sur de su rango de distribucion conocido, en el este de Texas 


The genus Lespedeza is present in Australia, eastern Asia and eastern North 
America (Clewell 1966; Turner 1959; Correll & Johnston 1979). Three species of 
Lespedeza, L. striata (Thunb.) H. & A,, L. stipulacea Maxim. and L. cuneata 
(Dum.cours.) G. Don, have been introduced in eastern U.S.A. to prevent soil ero- 
sion, for green manure and for forage for cattle and wildlife. (Clewell 1966). Les- 
pedeza cuneata is one of many plants used for soil and water conservation; it 
provides good ground cover on soils that are not adequate for other plants, and 
it helps build the soil by replenishing the nitrogen supply (Guernsey 1970). In 
Mexico there are only isolated records for this genus: L. virginica (L.) Britton 
and L. texana Britton are reported for Coahuila (Clewell 1966), and Estrada and 
Marroquin (1992) reported L. repens for central part of Nuevo Leon. There are 
no records of Lespedeza species for Sonora (Shreve & Wiggins 1964), Arizona 
(Kearney & Pebbles 1960), and Chihuahua (Estrada @ Martinez 2000). Figure | 
shows its historical and recent range distribution in U.S. A. and Mexico. 


Lespedeza cuneata (Dum.Cours.) G. Don 

Perennial herb, 25-45 cm tall; stems several from the base, erect and, ascend- 
ing, branched above, longitudinally ribbed, appressed pubescent, the trichomes 
0.2-0.4 mm long, white, most abundant in ribs; stipules persistent, linear lan- 
ceolate, 2.5-5 mm long, striate, sparsely pubescent; leaves abundant, crowded, 
trifoliolate, petioles 2-3.5 mm long; leaflets cuneate 7-10(-18) mm long, 2-3 mm 
wide, apex retuse, truncate or mucronate, Byabrous above, sparse Papescent below, 
lateral veins ascendant, branched, t inf] y, shorter 


SIDA 19(3): 741 — 743. 2001 


742 BRIT.ORG/SIDA 19(3) 


than the foliage, solitary or in pairs; pedicels | mm long; bractlets 2, ovate, apex 
attenuate, L.2-19 mm long; calyx narrowly campanulate, 4-4(-5) mm long, 1-3 
mm wide, 5-dentate, the throat 1.3—1.5 mm long, the teeth 2.7-3 mm long, lan- 
ceolate, the two upper notched less deeply than three lower ones; corollas white, 
papilionate; banner obovate or spatulate, 6-7.3 mm long, the base thiny, au- 
riculate and unguiculate, with a purple throat, the wings narrowly oblong, 
inequilateral, 6-6.2 mm long, 1.3 mm wide, the keel incurved, 6-7 mm long, 1-3 
mm wide, apex retuse; stamens 10, diadelphous; ovary ovate; style linear fili- 
form; fruit in lower branches originated from cleistogamous flowers, rounded, 
2mm in diameter, reticulate, the style tightly recurved, sparse pubescent; fruits 
of chasmogamous flowers undeveloped. 


Specimens examinated: Chihuahua:km 135 carretera Cuauhtémoc-La Junta,a la altura de Paramo 
de Morelos, Mpio. Guerrero, vegetacion de pastizal mediano abierto de Bouteloua gracilis, Brickellia 
spp., Mimosa aculeaticarpa, 1950 m.s.n.m., 25.Jul. 1994, C. Yen, F. Estrada 2693 (ANSM, CFNL, TEX) 


Fid.1M t . hy ee) ere) ra 4 #7 {fy Wa Don inUsA A fhhayi (Cert £CFhil | \ 
} t \ AI . 


ESTRADA AND YEN, LESPEDEZA CUNEATA IN MEXICO 743 


The presence of Lespedeza cuneata is sporadic, and patchy along roadsides, it 
does not appear to invade adjacent abandoned croplands. This species has not 
been internationally introduced in Chihuahua (people of the area, personal 
comments). As far as we observed, its distribution comprises small patches in 
an area of 26 km in this area. (from km 120 to km 146 Cuauhtémoc-La Junta 
highway). 


ACKNOWLEDGMENTS 


We thank B.L. Turner for plant verification, John L. Anderson, James Henrickson, 
and Barney Lipscomb for helpful suggestions, Mariela Pando for English revi- 
sion, and Miguel Gonzalez B. for the map. 


REFERENCES 


Crewelt, A.F. 1966. Native North American species of Lespedeza (Leguminosae) Rhodora 
68:359-405. 

Corrett, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner. 

Estrapa, E.and J.S. Marroauin. 1992.Leguminosas del centro sur de Nuevo Leén. Bol. Cient. 
No. Especial 10. Facultad de Ciencias Forestales, U.A.N.L. 

Estrapa, E.and A. Martinez. 2000.Legumes from the central part of the state of Chihuahua, 
Mexico. Sida 19:351-360. 

Guernsey, WJ. 1970. Sericea Lespedeza its use and management. U.S.D.A. Farm Bull. No 
2245.Washington, D.C 

Kearney, T.H.and R.H. Peestes. 1960. Arizona flora. University of California Press, Los Angeles. 

SHreve F.and |. Wicains. 1964. Vegetation and flora of the Sonoran Desert. Stanford Univer- 
sity Press. Stanford, CA. 

Turner, B.L. 1959. The legumes of Texas. University of Texas Press. Austin. 


744 BRIT.ORG/SIDA 19(3) 


Book REVIEW 


Ropert H. MacArtHur and Epwarb O. WiLson. 2001. The Theory of Island Bioge- 
ography. (ISBN 0-691-08836-5, pbk.). Princeton oS Press, 41 Will- 
iam Street, Princeton, NJ 08540, U.S.A. (Orders: www.pupress. pri edu, 
609-258-5714, 609-258-1335 fax). $19.95, 203 pp, 60 cae oe wo 


This newest edition in the Princeton series of b andmarks in Biology brings us an essential text that 
when published in 1967 made an i 1 the field of ecology. While the parameters of the 
assembly of discrete biotic communities is a en the innovation here was the authors breaking 
away from the mold to explore other phenomena like demography and competition. In the 30 years 
since initial publication, The Theory of Island Biogeography’s impact has resonated primarily in the 
disciplines of conservation biology and biogeography. This is an introductory text in the truest sense. 
In hindsight, Wilson notes in the preface: “the flaws in this book lie in its oversimplification and 
incompleteness, which are endemic to most early efforts at theory and synthesis. Large number of 
experiments supported 0 our i hypothesis ones did not. Also, we were satisfied to account for the ef - 

r utcome of varying populacion size and fluctua- 
llisland i ll lati arge ones 


and the effect is exacerbated wher ue amplitude of populauen! fluctuation is eat Later, oth- 


ers were quick to point out that populz eis = see m the vee wee Tae area a effect owes a lot 
to siosicaidensraias (iw Ne heless, thi { island bio- 
geography. 

Th g | he followi ics: the importance of islands, area and num- 
ber of species, explanations of the area eadiversity pattern invisibility and the variable niche, biotic 
exchange, and evolutionary chang The book begins with the area-diversity 


curve and throughout an extensive theory of the equilibrium of species is developed. The authors 
then go on to consider the influence of life-table parameters of individual organisms on the immi- 
gration and extinction rates of populations and generalize about the evolutionary strategies species 
must adopt in order to be good colonizers. The role of stepping stones in dis pore and secu of 


species and the consequences of island biogeography theory theory 
closing the book. The book is oe detailed in the mathematical formulas that describe and 
support their theory and al useful glossary of terms and general subject index. 


In summary, | would recommend i hae for island biogeography, conservation biology, or 
history of science classes. It serves as a snapshot of the early theoretical development of biogeogra- 
phy by which students can gain an appreciation for the disciplines evolution in light of the far more 
detailed work that has preceded it. The text is concise, affordable and the product of two men whose 
contributions to biology have been immeasurable.—Kevin D. Janni, Botanical Research Institute of 
Texas, Fort Worth, TX 76102-4060, U.S.A, kjanni@britorg. 


SIDA 19(3): 744, 2001 


RUPERT C. BARNEBY 
(October 6, 1911-December 5, 2000) 
A MEMOIR 


Richard Spellenberg 
Department of Biology 
New Mexico State University 
Las Cruces, NM 88003-8001, U.S.A. 

“Dear Rich: Thanks so much for your letter of July 13, for the good news in it, 
and for the little loco from near St. George, Utah, which brought a special joy 
(your no. 3182). I feel sure that you have run it correctly to A. musimonum—it’s 
simply not possible to squeeze a flower so short, a calyx so distinctly campanu- 
late, into A.amphioxys.... What gives me a particular boot about your plants is 
this: in 1942 I collected on limestone in Mokiak Pass s of St. George an astraga- 
lus at the time only in young flower which seemed to me almost certainly A. 
musimonum. ... This collection (my no. 4321) was cited provisionally in the 
protolog ..., but later I was never able to locate the material for comparison. ... 
[Ylou can imagine the satisfaction that your collection brings to me, the loss of 
my own having stuck in my mind like a splinter in the foot, healed over but 
still and always a cryptic nag” (Aug. 2, 1973). | received an essay, essentially, 
about a single collection we had made in Mokiak Pass on a circuitous return 
from a collecting trip with two graduate students to Death Valley. The collec- 
tion was not particularly significant, but still one that elicited a letter so grati- 
fying toa young botanist, a letter that simultaneously provided a confirmation 
on identification of a little-known species from a world’s expert on this im- 
mense genus and provided images of Rupert Barneby and Dwight Ripley three 
decades earlier, during a great world war, isolated in southwestern Utah sniff- 
ing out locoweeds and other marvelous flora in a very open West. As do others, 
I treasure my letters and memories from Rupert, always so positive with regard 
to the material at hand, so encouraging, and so expressive. The specimens were 
“run,” of course, in his at-that-time recently published “Atlas of North Ameri- 
can Astragalus” (1964). 

Rupert Barneby was born on October 6, 1911, at Trewyn, Monmouthshire, a 
17" century house in England nearly on the Wales border. He died at 89 years 
old at 5:10 PM, Tuesday, December 5, 2000, at the Jewish Home for the Aged in 
New York, where he resided subsequent to a mild stroke a few months earlier. 
Until the stroke, he was in his office nearly every day, continuing to work with 
his botany, at the last “primarily identifying the gazillions of specimens sent to 
him for det.” Jackie Kallunki). The world has lost a tremendous botanical tax- 
onomist and grand human being. 


SIDA 19(3): 745 — 751. 2001 


746 BRIT.ORG/SIDA 19(3) 


Early in his childhood Rupert was fascinated with plants and insects and 
fossils, and two aunts gave books to him that encouraged his interest. At 14 
Rupert excelled in producing a herbarium collection ina local naturalist’s club 
competition. Some of his identifications where challenged. Rupert knew he was 
right, and came to the realization that he had an independent intellect. Not only 
from books does one gain knowledge, but real learning could come from ob- 
serving the natural world. In public school, at Harrow, at age 14, he met Dwight 
Ripley, two years older, who had a knowledge of plant scientific names. This 
deeply impressed Rupert, and from there a life-long friendship grew. Officials 
were scandalized, Rupert enjoyed telling, by the close relationship that devel- 
oped, not so because of a schoolboy romance, but because Mr. Ripley was Ameri- 
can. After boarding school Rupert went to Grenoble University in France, and 
to Cambridge (Trinity College), where he finished a B.A. in History and Mod- 
ern Languages before he was 21. Mr. Ripley attended Oxford. While at univer- 
sity they went on joint plant-hunting trips to Spain, the Mediterranean, and 
northern Africa, bringing back plants to grow in the rock garden at the Ripley 
estate at Sussex, a garden that ultimately grew to contain 1138 species. As was 
the case for other great biologists of the past few centuries, Rupert's father re- 
sisted his study of botany; it was unsuitable for a young man. Suitable occupa- 
tions were the army, navy, or church, or as they encouraged, a career in diplo- 
macy. Rupert relates, “I was unsuitable for the army or navy and I hated the 
church. That’s really why I came to America.” Rupert was disinherited; Mr. 
Ripley’s personal fortune paid the bills. 

The relationship between Rupert Barneby and Dwight Ripley was ex- 
pressed largely in the development and appreciation of their garden. In 1939 
they published together a catalogue of the plants growing at their house, The 
Spinney, in Waldron, Sussex, England. Carlina barnebiana Burtt & PH. Davis, 
a thistle from Crete, dates from this period of their explorations. Overnight 
guests at the Ripley/Barneby house might very well find a bud vase on their 
headboard placed by Rupert, with a small bouquet of plants in their families of 
interest, taken from the garden. After 50 years of sharing their enthusiasm for 
botany and the beauty of plants, Rupert lost his life-long friend; Mr. Ripley died 
of complications arising from alcoholism. 

Rupert arrived in the United States in 1937, first living in Hollywood, later 
moving to New York. He established permanent residency in 1941. He and Ripley 
continued their plant hunting in the American West. In addition to collecting 
living plants for their rock garden, Rupert also prepared herbarium specimens, 
many of them representing undescribed taxa from that still poorly known re- 
gion. Among his several newly made friends in western botany, Alice Eastwood 
and John Thomas Howell encouraged Rupert to publish his first new species in 
1941, from Yucca Flat in Nevada, Cymopterus ripleyi. The article appeared in 
Leaflets of Western Botany, a journal that they supported financially for many 


SPELLENBERG, RUPERT C. BARNEBY, A MEMOIR 747 


years. In the same issue Eastwood described Castilleja barnebyana in honor of 
her friend who had deposited many collections at CAS. From that beginning 
he, often with others, named more than 1160 plant taxa new to science (Mi- 
mosa 217; Astragalus 118; Cassia 112; Senna 98; Dalea 61, Chamaecrista 50; etc. 
[NYBG database]) in 147 publications. In all western states, plant taxonomists 
and conservationists deal with Barneby names daily. In New Mexico, for ex- 
ample, he authored 26 plant taxa (23 Astragalus), and made new combinations 
in names for 44 more (K. Allred, NMCR database). A search of Index Kewensis 
on CD-ROM in 1997 listed 2045 taxa with Rupert as publishing author. Many 
of the drawings in his publications are by his own hand. 

Rupert arrived at the New York Botanical Garden in the early 1950's as a 
visiting scholar to consult the herbarium. He soon became an honorary cura- 
tor in Western botany. In 1959 he was given an official position to help Howard 
Irwin proceed with his studies of Cassia. In 1989 he was named curator of sys- 
tematic botany, the first and only paying job he ever held. He consulted for 
Brittonia, particularly to vet Latin descriptions for new taxa, and to critically 
read manuscripts. His special interests were xerophytic floras, taxonomy of the 
Leguminosae, Neotropical Menispermaceae. His extensive knowledge of the 
Fabaceae resulted in thousands of specimens being sent to him for determina- 
tion. These gifts have pushed the legume collection at NYBG beyond 270,000 
specimens. 

Rupert's ability to discern differences and recall detail, and to deal with 
cards and sheets of notes in extensive files while working on a typewriter, was 
matched by his truly astounding ability to synthesize. “Rupert Barneby was a 
great student of plants in the style of George Bentham and the other encyclope- 
dic workers of the nineteenth century, who would tirelessly analyze all we knew 
about enormous groups of plants and reduce that knowledge to lucid prose, 
working day after day, month after month, and year after year” (Peter Raven, as 
cited in The New York Times). So true; chatting over tea on one of my visits to 
NYBG I learned that he spent the best part of six months dealing with the varia- 
tion and synonomy of Lupinus argenteus Pursh for the Intermountain Flora 
(Fabales, Vol. 3,B, 1989). His ability to organize and synthesize massive amounts 
of detail, unrelentingly moving great projects toward completion, have given 
us magnificent taxonomic syntheses of Oxtytropis (1952), Astragalus (1964), 
Cassia (1977, 1978,) and Cassiinae (1982), Dalea and allied genera (1977), 
Mimosaceae (1996, 1998). Of his 147 botanical publications, 111 are in the 
Fabaceae. In all, they comprise more than 6,600 printed pages. 

lam grateful to Stan Welsh for the following passage, from his Jan. 16, 2001, 
presentation at NYBG at Rupert’s memorial service. It reflects upon Rupert's 
love of the West, of the field, of plants; on his humor, on his valued involvement 
with colleagues, and his view of himself relative to the “real” world. Writes Stan, 
“Field work was part of Rupert’s Psyche. He spent a huge amount of time col- 


748 BRIT.ORG/SIDA 19(3) 


lecting plants, becoming acquainted with them in nature. And, he had an al- 
most mystical quality of being able to ferret out novelties over vast expanses of 
the American West. His field experiences were sometimes interrupted by real 
stupidities, as when he was accosted by police in Arkansas in 1963.‘A suspected 
murderer was known to be in flight, in a Jeep, and naturally anyone in an out 
of state Jeep [Rupert's favorite field vehicle] was it. lt was a nasty experience 
being forced out of the car at gun point by a porcine state cop of the lowest 
(almost Hollywood) style—huge belly, flabby cheeks, cigar-butt clamped into a 
tiny red mouth—and then have all my possessions pawed over. Only botanists 
believe in anything so other-worldly as a botanist, and | do sort of sympathize 
with the Law faced by Linnaeus; one cannot fairly expect comprehension or 
sympathy. The quote is from a letter to Isely (16 July 1965), and was in response 
to Duane’s having been held in jail for some hours by Colorado police, him look- 
ing suspiciously like a bank robber. Botanists are a suspicious bunch.” 

Rupert understood and used Latin well, consulting for Brittionia and, upon 
request, helping those less skilled in the idiom to prepare proper descriptions 
and diagnoses. He could also be relied upon to provide opinions on the use of 
Latin in botany. For example, in New Mexico a number of us were preparing a 
review of plant species of conservation concern in the state, and we encoun- 
tered a specific epithet spelled in two ways in the literature. Solution: ask Rupert 
— and we received: “Dear Rich: there’s so much wrong with the epithet mesa- 
verdae that it would be best put on an index expurgatorius, but as we don’t 
have a method for this it is best left exactly as originally written. Latinized Mesa 
Verde would be mensa viridis, giving a genitive mensae-viridis: simply putting 
a Latin genitive ending on one or both parts of the Spanish place name is not at 
all that same thing. If any tinkering were to [be] attempted it would be best to 
think of mesaverde as one word and make a genitive mesaverdei. In any case 
mesae-verdae is even more grotesque than the original monstrosity, which | 
would recommend you leave unaltered, as a warning to those who assume that 
they have mastered Chopin yet are at page one of Czerny’s exercises” (14 March, 
1988). lam so glad I asked. 

The botanical community showed its appreciation for Rupert’s magnifi- 
cent contributions again and again by bestowing upon him prestigious awards. 
Rupertalways struck me asa modest and unassuming individual, appreciative 
of the work of others, truly interested in their progress. Awards were not his 
cup of tea. In response during an interview regarding his receipt of the 
Millenium Botany Award at the International Botanical Congress in 1999, given 
for his lifetime of contribution to botany, he said, “I’m conscious of the prestige 
of the medallion, but hideously aware that it’s an award for survival rather than 
for merit. It’s part of the dismal cult of personality that started in Holly wood 
and now has infected the entire planet.” Among other prestigious awards are 
the Distinguished Service Award, NYBG, 1965; the Henry Allan Gleason Award, 


SPELLENBERG, RUPERT C. BARNEBY, A MEMOIR 749 


an annual award from NYBG for an outstanding recent publication in plant 
taxonomy, ecology, or geography, in1980; the Asa Gray Award, American Soci- 
ety of Plant Taxonomists, for his contribution to systematic botany, in 1989; 
and the Engler Silver Medal, International Association of Plant Taxonomists 
highest honor for publications, for his monographic work in Mimosa, in 1993. 
He also was awarded an honorary doctor degree in 1979 from The City Univer- 
sity of New York. In 1991, NYBG established the Rupert C. Barneby Fund for 
Research in Legume Systematics, a fund that continues to support research in 
this large, important family. 

Rupert was an unofficial mentor and valued colleague to many. Duane Isely 
spent a sabbatical period with him at NYBG. Many, among them, Jim Grimes, 
Melissa Luckow, and André de Carvalho, recently have credited him as an in- 
spiration in their lives. Ghillean Prance shares that “he will be remembered by 
thousands of colleagues for his uncommon generosity in sharing his inexhaust- 
ible knowledge and precise editorial skills.” Stan Welsh writes to me, “.. [he] was 
a master of words, works, and wonders....| miss him already.” Cronquist wrote 
(Brittonia 33:263. 1981), “Rupert isa gentleman and our resident classical scholar. 
If we need to know something about Latin, or Greek, or the niceties of English 
construction, we turn to him. He is kind, considerate, and learned. No polemi- 
cist, he can come up with the piquant mot juste when he chooses. We love him.” 
The last speaks for so many. He had a delightful sense of humor, and loved a 
twist of the word. Loften used a heading on my letters that involved a pun, “From 
the Herbarium, where...” In response I received a letter from Rupert, “From the 
New York Botanical Garden, where Brittonia waives the rules.” 

Volume 33 of Brittonia was dedicated to Rupert C. Barneby on the occasion 
of this 70 birthday, “in recognition of his devotion and intellectual commit- 
ment to plant systematics and his extraordinary depth and breadth of scholar- 
ship.” In issue number three of that volume (33:263-274. 1981) is a series of let- 
ters from friends and colleagues from several nations. Each letter lauds his 
intellect, his accomplishments, and to a one, each expresses deep appreciation 
for the warmth of this extraordinary person. Joseph Kirkbride, who received 
his doctoral training at NYBG, brings back memories for many in one para- 
graph of his letter, “In his office, he had prepared a pot of tea and opened a pack- 
age of biscuits. The tea was Jackson of Piccadilly, and the blend was ‘Earl Grey’s, 
his favorite blend and brand of tea. That first ‘tea time’ was a marvelous experi- 
ence. He put me at ease and kept the conversation going as he introduced him- 
self. It isa landmark in my life.” In a modern e-mail message to Pat Holmgren, 
Stan Welsh writes, “My trips to the garden were always highlighted by the morn- 
ing and afternoon teas in his office.” In my own trips to the Northeast, for pro- 
fessional reasons or otherwise, | would always make a special effort to arrive at 
Rupert's office to spend some time visiting with him over a cup of tea, each 
time a very special moment, the memories of which I now value so much. 


750 BRIT.ORG/SIDA 19(3) 


In the same vein, in a communication from Noel Holmgren (Dec. 8, 2000), 
| learn that he also considers Rupert an important mentor in his development. 
He expressed what so many of us have felt in our interactions with Rupert. 
“There was no formal structure to the lessons, they were just part of relaxed, 
enjoyable conversation. He loved the same plants that 1 was becoming ac- 
quainted with. Each species of plant had a special character, be it the place it 
grew, its appearance, or its relationships to other species. He always gave the 
feeling that there was a spirit residing in each plant. You, Rich, have had these 
enlightening conversations with Rupert and so have so many others. So many, 
many others. I know this after years of being right across the hall from his of- 
fice. In some ways his hearing loss was sometimes my gain. | could listen to his 
tea-time conversations with people, whenever I chose. His wonderful and sud- 
denly explosive laughter. Pll really miss him.” As will I], and so many others 
who Rupert so generously touched in his long and productive life. 

Author’s note.—A number of individuals responded enthusiastically and 
helpfully when | asked for information regarding Rupert Barneby’s great life. 

Clearly he was extremely important to, and well-liked by, them. I know there 
are dozens of others who have had exchanges with Rupert that they would have 
been happy to share. lam particularly grateful to Pat Holmgren, who forwarded 
numerous very valuable sources of facts and perspectives on his private and 
professional life. Particularly helpful were New York Times, 10 Dec 2000, NYBG 
Herbarium Sheet #254, 15 Oct 1997, an NYBG press release, Dec 8, 2000, and 
Rupert Barneby’s vitae. 


SPELLENBERG, RUPERT C. BARNEBY, A MEMOIR 751 


R f by hot by N | Hol , who writes (31 Jan 2001),“TI pt g pt {R [ t tal M y 30th 
1978.1 he last full day in the field y he flew back to N or had pesayltig t0 get 
acandid photo of him daily for the preceding 10 11 could Il it off. | thougt i 

if he were aware of what | was trying to do, so finally, out of frustration | asked him to pose. | couldn't believe how 
natural and photegenlc he was. The backdrop is on the San Rafael Swell in Emery County, Utah, south of Interstate 
Highway 70. 


752 BRIT.ORG/SIDA 19(3) 


Book REVIEWS 


RICHARD S. FeiGer. 2000. Flora of the Gran Desierto and Rio Colorado Delta. (ISBN 
0-8165-2044-5, hbk.). University of Arizona Press, 355 S. Euclid Ave., Suite 
103, Tucson, AZ 85719, U.S.A. (Orders: www.uapress.arizona.edu, 520-621- 
1441, 520-621-8899 fax). $75.00, 700 pp, 440 illustrations, 8 1/2" x 11". 


| led , Abbreviations. Part I. The Envi d Human 


Contents. 
cluding Paleoclimate, Pent Cinere Major Habitats, History and Human Influences, Growth Forms, 
and Bortanical History). Part H. The Flora (the systematic portion, of 521 pages). Gazeteer (by Bill 
Broyles and Richard Felger). Appendices (including A. Growth Forms and Distribution of Plants in 
Northwestern Sonora; B. Distribution of Plants in Sykes Crater; C. Commonly Cultivated Trees and 
ants in Northwestern Sonora; E. Systematic Ar- 


Eo 


Shrubs in Northwestern Sonora; D. Non-native P 
rangement and Relative Abundance of the Grasses in a obese Sonora; F. ne aie Distribu- 


tions of Grasses in Northwestern Sonora) Literature Cited lt 
“Excuses, corrections, and ae oe to the book are Lae on the University of Arizona 
Herbarium (ARIZ) website <I I lu/HERB/ tools/gran_des.htmls. 


The flora area is in the others corner of the Mexican state of Sonora, an area of approxi- 
mately 15,000 square km (5790 square n mi, about 5% the size of the mia state of Arizona) and one 
of the most arid regions of North America. “It isa substantial portion of the extremely arid center, or 
the Sonoran Desert. Within ae eee there are expansive dune fields, maritime strands 


S,a 


heart, of 
small river, a once-great river and its delta, tidal wetlands, desert plains, steep granitic mountains, 
desert oases, and an enormous black and red volcanic field featuring its own mountain, lava flows 
cinder cones, and formidable craters. Also included is the Quitobaquito oasis, along the teal 
border but mostly on the Arizona side.” Average annual precipitation varies from 40 to 195 mm, 
depending on locality, and fearon is extreme - months or years may pass without significant 
rainfall, or much of the year’s precipitation may occur during a few hours. “Average laverage!] maxi- 
mum daily sneer ees 38°C (OOF) during June, July, and August, and temperatures ex- 
ceeding 38°C are common from late April to early October.” 
Within this cooker of an area, F ee: cocuments tne ocdurrence af 575 species of vascular plants 
| hese are North Ameri- 


a 


of t 


(by my count from Appendix A)i g | 


can natives). 


Comprising the bulk of the book are 
* original keys: 
iad but tusetul jectutes) desenpdone of Specs: an int maspeciic taxa, cad ‘onl yon plants and 


d’ and 


VLALEY 
emphasizing oo that seem important to dade mieadiae the variation ad: je of 
ronment: 


nd distribution, nativity, weediness, palatability, uses, and comparative notes 


pee in this envi 


* comments on habitat a 
on how to distinguish species from similar ones—this commentary often expansively developed, 
proweing a vehicle for Felger’s correspondingly expansive personal knowledge of these plants: 

* collection citations for each cane Bs great ae of these Felger’s own collections; and 


* illustrations—apparently | | by a line drawing, often with details, these gath- 
ered from a variety of sources, with s sriginals 


Ira Wiggins's 1964 “Flora of the Sonoran Desert” (which covers the Gran Desierto region) is cited 
only twice—once in the ‘Botanical History’ section (p. 38) in connection with mention of early col- 
lections made by Wiggins and once (p. 39) in connection with synonyms in FGDRCD not appearing 


SIDA 19(3): 752.2001 


BOOK REVIEWS 753 


in the Wiggins flora. This latter work, however, has more to go before outliving its usefulness, al- 
though, as one would expect after the last 35 years, the nomenclature is outdated in many areas and 
5 7 } } d ihed d Jac a] : a pe | > Bae 


connect from the earlier treatment probably reflects a true discontinuity, because the Flora of the 
Gran Desierto is truly an original, with trace of debt only to be inferred. And not only is F@DRCD 
packed with information, it is nicely organized and easy to read, good reading. Good price. For biolo- 
gists and conservationists with an interest in American desert floras or simply in the flora of west- 
ern North America, this book is a must-have.—Guy L. Nesom, Botanical Research Institute of Texas, 
509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A 


Karen L. Witson and Davin A. Morrison (eds). 2000. Monocots: Systematics and 
Evolution. (ISBN 0 643 06437 0, hbk.). CSIRO Publishing, PO. Box 1139 (150 
Oxford Street), Collingwood VIC 3066, Australia. $175.00, i-xiv, 1-738 pp, 
b/w ca ae 81/4" « ll a 


M S | lting f heS d Internationa 


= 


Conference on the Comparative Biclooy of the Monnesiyledons held in Sydney, Australia, during 
the week beginning 28 September 1998. A total of 280 individuals from 31 countries ata ieee in 
the conference, whose aim was to “increase our scientific understanding of the relationships, class 
fication and functional biology of the monocots...” According to the preface, the 72 paper. ee 
in this volume “are based on presentations given at the conference, but many have been updated or 
extended to take into account new information.” Given the rate at which much of the field of sys- 


rogressing, such updating is an i ideration. In addition, all the 
papers were peer-reviewed. A look at the authors isa good adiaeen of the quality of the conference 
and volume—they include many of the most recognized and aac authorities worldwide on 
monocots, and systematics in general, working today. Such well-kno ames as Barrett, Bernhardt, 

Briges, Chase, aaah Se Soldblat ss en, Merrow, Rudall, Soltis ae Tomlinson (to mention a 
volume. The ieee s have been oie o three major sec- 


tions (General Comparative Biclesy of the Monocots, Sys of the Lil Ce Systematics of the 
Commmelinoids), with each of these divided into a num = i eta For example, the section 
D:.] lL k; | FAs i | +4 ] - DI ] Tiz 1 ¥ ] 


on Comfy ig pay g gi geography & Fossils, 
Development & Organization, Chemotaxonomy & Cytology, Micromorphology, Anatomy & Em- 
ee and Reproductive Biolo 

If size (weight) is any indication of amount and qua lity of content, then this hefty six pound 
book is ce ene worthwhile. Indeed, the overall quality of the numerous papers is quite high, and 
the volume is extremely useful for anyone wishing to see a wide-angle snapshot of the current un- 
derstanding of monocot biology. In particular, the conference ene and proceedings editors 
should be congratulated on ane Rope act a conterence/proceedings that pene ee 
froma wide sings us lisciy anatomy, biogeography 
mental cane molecular systematics, [ a ieenniee cuneaaee biology). While means in oe 


ation is currently providing some of the most profound and epee insights in the field today, 


other cipine are also Sa major contributions, as shown so well in this volume. 
s said, I personally (with a taxonomist’s bias) oan ee of the articles focusing on mo- 
i. pec meme interesting. For example, the first paper of the volume, “Higher-level 
systematics of the Monocotyledons: An assessment of current knowledge and a new classification” 
M.W. Chase et al.), clearly shows how molecular systematics has developed and matured. It is a 


SIDA 19(3): 753. 2001 


754 BRIT.ORG/SIDA 19(3) 


well-presented and exciting look at current understanding of the relationships of the various mono 
cot groups based on a combined analysis of DNA sequence data from three genes. The consensus tree 
presented shows Acoraceae as the sister group to the rest of the monocots, followed by Alismatales 
(including Araceae and Tofieldiaceae) as the next diverging order, followed by Pandanales, 
Dioscoreales, Liliales, and finally Asparagales-commelinoids. Such an eee of monocot 


relationships would only have been dreamed about several decades ago. Another particularly inter- 
esting paper was “Phylogenetic studies of Asparagales based on four plastid DNA regions” (by M 

Fay et al.). In this case, a combined analysis of four plastid DNA regions was used to examine ae 
tionships among members of the Asparagales. This paper likewise is well-presented, has a visually 


easily understandable tree, and proviges a sasemanne eee at this ange oe important order. While 
many of the most important i the-less an impor 
tant Ani: of the molecular pylons ges of ae order. Understanding that the Amaryllidaceae, 


ragaceae, Iridaceae, and Orchidaceae (for ewe ie oe in the same order would have 


- unthinkable just a generation ane Here we see ad a and explanation of their 
eens While some of the d ge as further molecular work is done, studies such 
as these are important t d histi li gofm monocot phylog- 
eny. Valuable conteibadions from the other sections eciadine titles such as “Stem vasculature in 
climbing monocoty ledons: A comparative approach,” “Mating strategies in monocotyledons,” “Ins 
and outs of orchid ier or “Fire response and conservation biology of Western ene 
species of Restionaceae” could be reviewed here in detail. However, suffice it to say that overall this is 


a volume of diverse and exceptionally lly high h quality papers. 
If anything negative can be said, it would be that as expected from a proceedings volume, and 


from the size of the monocots (ca. 25% of the world’s flowering plant species), the papers do not pro- 
vide even coverage of all monocots groups. By necessity, it reflects the participants’ interests and 
expertise, and many groups have not been included (in Laven the grasses have been excluded 
but are covered in a well done companion volume). However, there is quite broad coverage, ranging 


8 
from such well known groups as Asparagales, ee Liliales, Orchidaceae, Cotmenelinacene 
and Cyperaceae, to lesser known but none-the-less interesting groups such as the Restionaceae 

In summary, this is a valuable addition to our mae of monocot ale and provides an 
ent overview of the current state of understanding. It will undoubtedly be one of the most 


— 


exce 
useful and broadly cited peicteness on monocot Seat ane evolution cau many years to come. 


Further, tl the diverse and dynamic 
nature of a enies and evolution at the beginning of the 21“ century.—George M. Diggs, Jr, Dept. of 
Biology, Austin College, Sherman, TX 75090, and Botanical Research Institute of Texas, Fort Worth, TX 


76102, U.S.A. gdiggs@a pees u. 


Aba GRAHAM and FRANK GRAHAM, JR. 1995. Kate Furbish and the Flora of Maine. 
(ISBN 0-88448-175-1, hbk, ISBN 0-88448-176-X, pbk). Tilbury House Pub- 
lishers, 132 Water Street, Gardiner, MA 04345, U.S.A. (Orders: 800-582-1899). 
$55.00 (hbk), $30.00 (pbk). 162 pp, 43 full color illustrations, 100 b/w il- 
lustrations, 6 b/w photographs, and a b/w map of Maine, 8" x 10". 

This is the oF of a aspirit driven vat a fire to collect and paint the unique and diverse flora that 

was Maine. Inz ntury (1834-1931) Catherine (Kate) Furbish would ‘paint 

a reputation as field Bolanict and botanical illustrator by creating more than 1,300 botanical water- 


colors and sketches. Self-taught in both fields of art and science, Kate had a no-nonsense approach to 


SIDA 19(3): 754. 2001 


BOOK REVIEWS 755 


her painting. She wasa serious naturalist with a botanist'’s eye. “Ido not claim Artistic merit, but merely 
atruthful representation of what I saw in the plants, free from all decorative effects.” 
Born the only girl amongst five brothers, she held a strong reserve inherited from her father. 
er work is not fussed over or decorative, but true botanical illustrations and yet the artistry does 
come through. Her illustration of Bloodroot, ee canadensis, created between 1870-1880 
n she was at her most productive, is maste n both composition and form. At 36 years old, 
unmarried and still ae an her parents, - was a woman of post Civil War Victorian conven- 
mee holding within an unrelenting personal quest; a body of scientific and artistic work that 
out to spring kort like no yorhen period of her life. Photographs taken of her at the time show a 


eet direct g nae earlier ploHees DORIS: 
ld ] Rene 


Kate was not daunted by arduous ‘solo’ f 
One of these expeditions in 1880 would lead her to ca the Furbish’s Lousewort, Pedicularis 
aes a wild snapdragon found nowhere in the world except along a 130-mile stretch of the 
Saint John River in Northern Maine. Its oy) ai eas her name endure. She was also not 
deterred by gender prejudices and lack of proper degree. Ina letter to a colleague she ex- 
presses her frustration with another botanist: “I think he is one ofthese men, who if I was young and 
the bloom was on the peach, ere more interested in helping me. I tried to show my appreciation 
by sending him my best work [a painting of trilliums] but I’m not going to wail over it all. For my part I 
help everyone whom I can and put my self out to do it too.” 

S as admired and respected by noted botanists of the day including her one time prodigy, 
Merritt Ree Fernald who was to become a noted field botanist and systematist at the Harvard 
Herbarium and editor of the New England Botanical Club's journal Rhodora. In his Second Edition of 
the ae ay Catalogue for Maine Plants he speaks o ae ve * Aster cordifolius that she had 
discovered in Aroostook County. “Dedicated to Mai tist- botanist, the “posy-lady 
of the Madawaska Acadians, ena herundaunted iat vii brush, has done more than 
any other to an known the wonderful flora nes 

It was not until the 19th century that | 
States and Asa Gray was one of the driving forces. He helped ae the popular interest in identifica- 
tion ane collection e wild plants. Thus, Kate's favorite and often referenced books were Asa Gray’s 
How Plant Manual of the Botany of the Northern United States. Unlike the rule of thumb 
that in biology diversity of ges diminished from the equator toward the poles, Maine held many 
ane? flora sist waiti Het be d d by advent botanists like Kate. Maine was also varied 

ds and li ing from beaches to salt hes to rocky headlands, 


y a } NT | United 


sandy plains, dense forests, river valleys, mountaintops, thousands of lakes and ponds, old fields, and 
cold sphagnum bogs. Maine also lay i rtion of its flora is made up of Southern 
plants in their northern most limits, and Northern ae plants in their southern most limits. 


Maine was ripe for exploration! 
At 49 years old, in the span of one decade Kate collected more than 1000 species of plants and 
of these eee 850. mies 1983 the complete checklist of vascular plants in Maine totals 2,137 known 


species.) “It h mplished by means of hard work and persistent effort, and without regard to 
fatigue,” wrote eee a have wants a a most ae on ae nue UG and inthe hedges, on 
foot, in hay ricks,in country vised rafts,(equipped 


with hammer, saw, nails, nile ripper ieee vasculum, ne in ue on sie rane o on ee 
and knees on the surface of bogs, and backing out, when I dare not walk, in order to pr d 
treasure. Called ‘ crazy, a deals this isthe way i: work as veer done, the Flowers being my ce soci- 


etyand the Manuals, t the only literature fo I happy y hours!” 
I like the way the authors have yaar Kate's ‘time-line’ of paintings along with the ‘time-line’ 
of the botanical history of Maine. 1 the design and layout of the book, the illustrations are 


SIDA 19(3): 755. 2001 


756 BRIT.ORG/SIDA 19(3) 


interspersed throughout the text in a very pertinent and pleasing manner. It is evident that much 
thought has been given to the layout and design of this fine biography.—Linny Heagy, Linny/ 
Designer, Illustrator, Arlington, TX, U.S.A., a0005835@airmail.net. 


Danie. W. Gabe. 1999. Nature and Culture in the Andes. (ISBN 0-299-16124-2, 
pbk.). University of Wisconsin Press, 2537 Daniels St., Madison, WI, 53718, 
US.A. $18.95, 298 pp, 46 figures, 6" x 9" 

The thrust of this volume is what the author refers to as the nature/c 

this work he proves that this is more than just an attempt at neologism:; it is, in fact, a reality of the 

Andean region. The nature/culture gestalt is “a mutually interactive skein of human and nonhuman 

components, rather than opposing polarities or separate entities”: Pg. 5. The Western eeachaal of cat- 


] 1 


And throughout 


8 
these two seemingly unrelated phenomena, whereby the interrelatedness 
Bly 


egorization t 
is lost and questions are soul partially answered. Cultural geographer and professor emeritus at the 
University of Vermont, Daniel W. Gade delivers a groundbreaking volume for the annals of Andean 
history, ecology, and ethnobiology. Nature and Culture in the Andes is a book with a holistic vision 
that attempts to broaden the perspective achieved solely by mene ewe sea of inquiry. 
self 


The ten chapters include an introduction and conclusion laced w 
tary on the author's observations of Andean culture throughout on years of fieldwork. The main 
chapters themselves are free of the author's self-reflexive voice, but are framed within the concepts 
that are a result of his own self-reflection. In the introductory chapter, references are made to every- 
one from Goethe to Nietzsche and the author provides an autobiographical discussion that frames 
his insights and perspectives on the geographically and culturally diverse region of Andean South 
America. The second chapter, “Andean Definitions and the Meaning of lo Andino,” covers the vari- 
ous meanings behind the term the Andes, geographically and culturally, and seeks to revise these 
definitions with his own. As the author states, “Over the past 400 y the definiti 
has shifted twice; et a nonregion to a region, and from a physic ale tity toa seine area”: Pg. 41. 

The third chapter, “Deforestation and Reforestation of the Central Andean Highlands,’ is a chap- 


n of the Andes 


ter that stands out for several reasons. The common oo tac of the Andes is one of a vast treeless 
lly a result of the economic demand 
role of anthropogenic fire and 

its role in environmental management. Fire was also used tone the Pacific Northwest regions of 
North America (Boyd 1999); ane European colonists there frowned on its use, and the result 


region. Contrary to popular belief, the treeless An 


| 3 


for wood and subsequent deforestation. The author 


being what many today See as ve pores pimieval when in actuality the current tree popula- 


tion is relatively new. In the es management was encouraged by Euro- 
pean settlers, the result ae the treeless Andes as we know it. Various species of Eucalyptus were 
introduced to the region and replaced to some extent the species lost. The author goes on to offer 42 
plant species, ‘iain! at the genus level, that once covered the Andean highlands of Bolivia, Ecua- 
dor, and Peru. Tree removal was also a result of opening up more land for agriculture. As a result of 
population pressure, more and more trees were removed and this eventually led to environmental 
acer ts However, without more land settled life would be limited in most regions 

fourth ne Malana aud Settlement LL aaa in Miz ee Bolivi ia, naibeusces the 


disease ee of malaria in | | by Euro- 
pean settlement. The author's case study provides a context for discussion see by one of the 


argest 
and most virulent malaria histories in the region of western South America is brought to light through 
he Andes as a Dairyless 


— 


migration, population density, and racial/ethnic change. Chapter five, “T 


SIDA 19(3): 756. 2001 


BOOK REVIEWS 757 


Civilization: Llamas and Alpacas as Unmilked Animals,” continues in the spirit of debunking the 
traditional anthropological assumption that milk bearing animals are always milked. Chapter six, 
“Epilepsy, Magic, and the epee in Andean America,” discusses the relationship of controlling epilep- 


tic seizures by wearing the nail of tapir around ones neck. Chapter seven, “Valle in of Mystery on the 


panne a Margin and the Inca Coca Connection,” discusses how the popu 
the Incas did not cultivat a has not properly been challenged. Chapter zee oa as Rat 
City,” a. the inundation of brown and black rats in this port city in Ecuador. Arriving as a 


result of overseas travelers, rats have played an re role in the health and aed oad 
terns of sah aes In the early 1900's the bubonic plague wreaked havoc 


due in no small part to the co-existence of rats and humans and was squelched by a majoe aati 
sanitation ee ee rats remain in the urban Guayaquil scavenging on trash left by hu- 
mans. o ing ond tion is a metaphorical Sleniiicanee Lats py aee me port func- 


Z 4 ] 


sent local poverty”: Pg. 183. eae nine, “Carl Sauer and the Andean Nexus in New World Diversity,” 
covers the life work of Carl Sauer’s stuaie>) in ma Wore ae domestication and biodiversity. 
ot only to its diversity, but al I bok wens = a valuable part of 
] i botany, 


any course in Andean Ethnobiology. G pp 
] 1 1l¢-l this stext 7 toa 1 P| lt hd 7 { 


eridite acme Moreover this is lively and interesting reading that dismisses ideas long held 


geoora phy 


about the pre-Columbian Andean environment and I relationships and seeks 
to replace them wala ideas cnet euule only be the result of rigorous research and a long career in the 
field. Perfect for RECORMAD hy Nanirear Culture in une Andes 


fo) 


Latin American studies. Daniel Gade nial in sane hie rature/culture estalt 


theory in central a ca and sets a lev el of by current and future 
pee of Ethnobiology.— Kevin D.Janni,B IR h Institut if Texas, Fort Worth, TX 76102- 
OU.S.A, kjanni@britorg. 


is bold, scho ial and i eae I would highl gin 
and Andea 


ye | iz Amal AA 
lOlaLl 


REFERENCE 
Boyp,R.(ed.) 1999. Indians, fire, & the land in the Pacific Northwest.Oregon StateUniversity 
Press. Corvallis, Oregon. 


EstTeLLeE LEveTIN and KAREN MCMAuon. 1999. Plants and Society. Second Edition. 
(ISBN 0-697-34552-1, pbk.). WCB/McGraw-Hill, Dubuque, IA, U.S.A. (Or- 
ders: www.mbhe.com). $55.60, +77 pp, b&w photos, 81/2" x 11". 

Plants and Society is not unlike other ee scaoeyercuaie level texts ae have come out in the last 

two years and have attempted to present bot life, rather than an 

esoteric nde discipline Ps ‘ittle applicability Homers ie: texts have ae heavily on 


the cultural aspects of plant use, where this text uses historical examples to punctuate its scientific 
discussions 

T ok comes in twenty-five chapters grouped in seven units. This firs 
the basic botany found in introductory courses, the rest of the book either looks at the ae as- 
pects of botany or the historical impact of the relationship between people and plants. Unit one, “the 
botanical connection to our lives,” covers flowering plants, fungi, an overview of the plant sciences 
and a general discussion of phytochemistry. Unit two is made up of eight chapters of introductory 


SIDA 19(3): 757. 2001 


758 BRIT.ORG/SIDA 19(3) 


botany covering: the pa on body, ang i paysiclogy, the plant life cycle (f1 fruits, and seeds) 
genetics of plant life. This portion of the book looks like 
most any other general botany textbook, with the exception of the boxes ee ed throughout with 
examples of applied and/or historical aspects of the given chapters subject m 

Beginning with unit three the text focuses more on the relationship of slats and society. Unit 
three is six chapters covering plants as a source of food. Human nutrition and the origins of agricul- 
ture set up the following four chapters covering the grasses, legumes, starchy staples, and the case of 


worldwide hunger. Unit four he commercial ale nies from pants in hii ae 
ters. These cate are aevoted to Poca pevelage es, h ind 3a- 
per meen Un | i the rol [planes in human health, specifi 

I psycl ive plants, and poi lallerg is aes ein 
of fungi in the natural envi tas | d food, and fungi health. Unit 


g 
seven closes the book with a chapter on sesleay that focuses on plants and the environme 

Each chapter follows the typical textbook design with key words in bold font, tables a boxes 
with case studies and/or ee ae me quizzes throughout the body of the chapter, a chapter 
summary, review questions and a list of further reading. Following the main chapters are an appen- 
dix on the metric Pee an happend on paat classification, a glossary, and an inde 


= 


I have a few design for this otherwise well-put-together text. oo the unit on fungi 
(chapters 22-24) would pproy follow chapter nine, the diversity of plant life. Secondly, 
units four and five could have b | | her. The ae devoted to psychoactive plants 
is superfluous and the same subject matter could bate been incorporated into the chapters on herbs 


and spices, medicinal plants, and poisonous 4 ants. | have two substantive criticisms for the text. 
Chapter eight, “Plant Systematics and Evolution,” is essentially a discussion on the oo of 
scientific and mes, a discussion of aa nomenclature, Se of Darwi 
ee on the H.M.S. ace In ae ena there is one paragraph that ti | its 
t to identify evolutionary | With the title eae carries | 
would like to see a more in-depth discussion of p balogeny and its implications for analyzing evolu- 
tionary eae My errand substantive criticism is that this book is essentially about plants 
with reference to | i nutrition, and civilization. In other words, there isn’t as much 
balance ee plants and sce as the title indicates. lam quite satisfied with the first half of the 
book and its introductory discussion of botany. At unit three the book takes a closer look at plants 
and people, but is still mostly about plants. Perhaps if each chapter had an individual case study of 
an ethnic group and their relationship with plants would correct this 
In summary, while this book is not as balanced as it could be it is nevertheless appropriate for 
nee penoe level units on pene and civilization and ces be se peace with ue texts io 


| } ] 
A better Ddalance 


Lh | id bly versatile and 


ore toa wider audience. Pla nts and Society is representative 
of the current endl in curriculum era that is shifting the focus of traditional botany courses by 
making the subject matter al and relevant to daily life and cease ies so.—Kevin 
D.Janni, Botanical Research fastihuee of a Fort Worth, TX 76102-4060 U.S.A, it 


TLOPK. 


SIDA 19(3): 758. 2001 


BOOK REVIEWS 


Bery_ B. Stmpson and Mo_ty C. OcorzALy. Economic Botany. Plants in our World, 
Third Edition. 2001. (ISBN 0-07-290938-2, hbk.). McGraw-Hill Higher 
Education, McGraw-Hill Companies, Inc., 1221 Avenue of the Americas, New 
York, NY 10020, US.A. (Orders: wwwimhhe.com). $70.50, 544 pp, illustrated, 
81/2" x 11". 


The distinction between economic botany and eee is Sica Say) For manly people, eco- 
h Amazonia 


nomic botany isan inventory plants used for projec 
in search of the cure toa modern disease. Neither o ae perspectives is particularly accurate. While 
melded in this text are a nium per of et thnobotanical examples, this book, as its title indicates, is 

botany is Mucus seus ie ne influenced.) 


This third edition of Economic Botany outlines the breadth of th ne and its impact on our 
lives, past and present. 
The book is made up of 19 chapters that have been signifi l dated si I Sa 


of the second Scion. eae one and two from the first two editions have been condensed an 
one, as on Their ne enipereo by People.” This in 
traditional methods of plant ma- 


ter discusses plants, 
nipulation, variation aad selection in leans plants, the nature of plant species, the naming of 
plants, and determining the relationships among plants. Chapter two reviews current ideas on the 
adoption of agriculture, as well as the origins of particular crops. 

The bulk of the book is made up of chapters three through 17 that focus on important an- 
giosperm and gymnosperm crop species. The first five of these chapters group food plants by the 
parts of the plants (ie - fruits, leaves, stems) harvested for food. Chapters eight through 16 cover prod- 
ucts that are Dananly extracted from plant parts. Substances such as volatile oils, alkaloids, latexes, 
g to their use: spices and perfumes, textile fibers, and bi e compounds. For 


are groupe 
each group of plant products discussed, the cial occurrences, chemistries, a ae within 
the plants in which they occur are outlined. Chapter three discusses fruits and nuts of temperate 
regions, focusing on ae s and their relatives and other oe Chapter four covers fruits and nuts 
from warm regions, like citrus fruits, tomatoes, peppers and eggy 
and forage grasses are covered in chapter five, which reviews the major grain crops and grass plants 
and forage grasses. Chapters six is devoted to legumes: pulses, tamarind, and carob. Chapter seven 
covers foods from leaves, stems, and roots and is one of the larger chapters in the book. Discussed in 
chapter seven are the structure and function of stems, leaves, and roots, biennial and annual crops, 
vegetables from bulbs, starchy root crops, and sweets from stems and roots 
Chapter eight covers the chemistry and ecology of spices, herbs, and perfumes. Chapter nine 

discusses the composition of seed oils like polyunsaturated, unsaturated, and moderately saturated 
oils and vegetable fats. Chapter 10 discusses hydrogels, elastic latexes, and resins. Chapter 11 focuses 
on medicinal plants, and in this edition include updated information on the most commonly us 
herbal remedies. The chapter covers the history of medicinal plants, the ene of plant ze 
medicines, and dietary supplements. Chapter 12 is devoted to psyct 

includes information about the chemistry and pharmacology of p 
of drug use use. Chapters 13 and 14 cover stimulating me Se parece Chapter 15 
covers fibers we and tannins, followed by chapter 16 covering wood, cork, and bamboo. Chapter 17 
covers ornamental plants, specifically, nursery crops, florist crops, and houseplants. ee 18 on 
Algae is expanded from the previous editions to include new information on bioactive algae that 
produce newly discovered toxins and research on the nae potential of algal compounds. The 
final chapter discusses the uses of plants in pine ee and is also revised. The book closes with a list 
nd an index 


=) 
h ti dri 
gs, and a history 


of additional readings, a glossary (new 


SIDA 19(3): 759, 2001 


760 BRIT.ORG/SIDA 19(3) 


In this updated edition the authors have made an effort to make the book more accessible in 
several ways. First, the material has been condensed into 500 pages that appear in two-column for- 
mat to save page space. Color photos have been excluded, but there exist a number of new photos 
that are up to date and more globally inclusive. Another new design feature are the boxes that high- 
light interesting case-study material about the plants included in different chapters. The concentra- 
tion in this edition is on species of ae economic Enpentante, in a Western world, rather than 
trying to be encyclopedic. TI I 


of plants includ- 
ing aspects of history, morphology, chemistry, sad moneva usage. At first glance this is a very text- 
book-like edition, however there are no summaries or review questions at the end of each chapter, 
suggesting that the target audience for this edition are graduate students. 

In summary, this updated and revised edition of Simpson and Ogorzaly’s original Economic 
Botany is an excellent text that should be the foundation for graduate level economic botany classes. 
The updated setae on ee: plans, erenabotany, genetics and ee and 

ustainability and cons book relevant and practical to students and instr 
tors of economic uae | would highly recommend this book to anyone wanting a thorough global 
view of economic botany for a class or personal use.—Kevin D. Janni, Botanical Research Institute of 
Texas, Fort Worth, TX 76102-4060 U.S.A, kjanni@britore. 


SIDA 19(3): 760. 2001 


BOOKS RECEIVED 761 


Books RECEIVED 
Book reviews will be published in upcoming issues of Sida. 


ral ° IMA 1 fama | Tae | 


B. NorpenstaM, G. EL-Guaza_y, and M. Kassas (eds.). 2000. Plant Systematics for 
the 21st Century. (ISBN 1-85578-135-2, hbk.). Portland Press Ltd, 59 Port- 
land Place, London W1B 1QW, UK. (Orders in North America: Princeton 
University Press, +1 William Street, Princeton, NJ 08540, 609-883-1759, 609- 
883-7413 fax). $120.00, 366 pp, figures. 


Rosert W. Kicer and Duncan M. Porter. 2001. Categorical Glossary for the Flora 
of North America Project. (ISBN 0-913196-70-3, pbk.). Hunt Institute for 
Botanical Documentation, Carnegie Mellon University, 5000 Forbes Avenue, 
Pittsburgh, PA 15213-3890). $5.00 (shipping $4.00), 165 pp. 


Marco LAMBertINI (Translated by JoHN VENERELLA). 2000. A Naturalist’s Guide to 
the Tropics. (ISBN 0-226-46828-3, pbk.). The University of Chicago Press, 
110305. Langley, Chicago, IL 60628, wwwopress.uchicago.edu, 800-621-2736). 
$25.00, 312 pp, illustrated, color. 


S.HJ.V. RApANARIVO, J.J. LAVRANOoS, A.J.M. LEEUWENBERG, and W. Roos it. 1999. 
Pachypodium (Apocynaceae). Taxonomy, Ecology and Cultivation. (ISBN 
90-5410-485-6, hbk.). A.A. Balkema Publishers, Old Post Road, Brookfield, 
VT 05036-9704, www.balkema.nl, ball @ball nl, 802-276-3837 fax, 
31-10-4135947). $48.00, 128 pp, 90 color photos. 


N.M. Nayar and T.A. More. 1998. Cucurbits. (ISBN 1-57808-003-7, hbk. ). Science 
Publishers, Inc., P.O. Box 699, May Street, Enfield, NH 03748, 603-632-7377, 
603-632-5611 fax). $75.00, 340 pp. 


EpwarD A. Cope. 2001. Muenscher’s Keys to Woody Plants: An Expanded Guide to 
Native and Cultivated Species. (ISBN 0-8014-8702-1, pbk.). Cornell Univer- 
sity Press, Sage House, 512 E. State Street, Ithaca, NY 14850). $22.95, 377 pp. 


Pat HAtuipay. 2001. The Ilustrated Rhododendron: Their Classification Portrayed 
Through the Artwork of Curtis’s Botanical Magazine. (ISBN 0-88192-510- 
1, hbk.). Timber Press, 133 SW Second Ave., Suite 450, Portland, OR 97204- 
3527, U.S.A. (Orders: www.timberpress.com, 800-327-5680, 503-227-2878, 
503-227-3070 fax). $69.95, 268 pp., illustrations, color. 

Ropert H. MOHLENBROCK. 2001. The Illustrated Flora of Illinois. Grasses. Pani- 
cum to Danthonia, Second Edition. (ISBN 0-8093-2360-5, hbk.). Southern 
Illinois University Press, P.O. Box 3697, Carbondale, IL 62902-3697, 
danseit@siu.edu, 618-459-6633, 618-453-1221 fax). $50.00, 455 pp, 306 line 
drawings, maps. 


SIDA 19(3): 761. 2001 


762 BRIT.ORG/SIDA 19(3) 


Ropert H. MOHLENBROCK. 2001. The Hlustrated Flora of Illinois. Sedges. Cyperus 
to Scleria, Second Edition. (ISBN 0-8093-2358-3, hbk.). Southern Illinois 
University Press, PO. Box 3697, Carbondale, IL 62902-3697, danseit@siuedu, 
618-459-6633, 618-453-1221 fax). $59.95, 223 pp, 128 line drawings, maps. 

ROBERT H. MOHLENBROCK. 2001. The Illustrated Flora of Illinois. Pokeweeds, Four- 
O-Clocks, Carpetweeds, Cacti, Purslanes, Goosefoots, Pigweeds, and Pinks. 
(ISBN 0-8093-2380-x, hbk.). Southern Ilinois University Press, RO. Box 3697, 
Carbondale, IL 62902-3697, danseit@siu.edu, 618-459-6633, 618-453-1221 
fax). $39.95, 277 pp, 139 line drawings, maps. 


Donatp Watts. 2000. Elsevier's Dictionary of Plant Names and Their Origin. (ISBN 
0-444-50356-0, hbk.). Elsevier Science, PO. Box 211, 1000 AE Amsterdam, 
The NETHERLANDS) (North American orders: Elsevier Science Inc., PO. 
Box 945, Madison Square Station, New York, NY 10160-0757, wwwelseviercom, 
31-20-485-2603, 20-485-3533 fax). $209.00, euro 181.51, NLG 400, 1032 pp. 


ARMANDO T. HUNZIKER. 2001. Genera Solanacearum. The Genera of Solanaceae 
Illustrated, arranged according to a New System. (ISBN 3-904144-77-4, 
hbk.). Koeltz Scientific Books, Herrnwaldstr. 6, D-61462 Konigstein, GER- 
MANY, wwwkoeltz.com, koeltz@t-online.de, 49-6174-93720, 6174-937240 
fax). $120.00, 240.00 DM, 500 pp, 136 line drawings. 


WARREN L. WAGNER, DERRAL R. Herest, and S.H. SoHMER. 1999. Manual of the Flowering 
of Hawai ‘i, Revised Edition, (2 volumes). (ISBN 0-8248-2166-1, hbk.). Published 
in association with Bishop Museum Press, University of Hawai'i, Press, 2840 
Kolowalu Street, Honolulu, HI 96822-1888, uhpbooks@hawaii.edu, 808-956- 
8697). $95.00 (2 vols.), 1919 pp, illustrated. 


B. LERoy Davipson. 2001. Lewisias. 2000. ISBN 0-88192-447-4, hbk.). Timber Press, 
133 SW Second Ave., Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: 
www.timberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). 
$34.95, hbk. 238 pp., 68 color photos, 20 color plates, 13 line drawings. 


Frep WaAMPLER (Text) and MARYROSE WAMPLER (Artist). 2000. Trees of Indiana. ISBN 
0-253-32885-3, hbk.). Indiana University Press, 601 North Morton Street, 
Bloomington, IN 47404-3796, iupress@indiana.edu, 812-855-6804, 812-855- 
7931). $49.95, 192 pp, 160 color photos (paintings). 


Eel } 


. a oF yee 


l/Useful Plants 

FREDERICK J. SIMOONS. 1998. Plants of Life, Plants of Death. ISBN 0-299-15904-3, 
pbk.). The University of Wisconsin Press, 2537 Daniels Street, Madison, WI 
53718). $34.95, 568 pp, illustrated. 


SIDA 19(3): 762. 2001 


BOOKS RECEIVED 763 


Paut Minnis (ed.). 2000. Ethnobotany, a Reader. (ISBN 0-8061-3180-2, pbk.). University 
of Oklahoma Press, 4100 28th Avenue N.W, Norman, OK 73069-8218, 
www.ou.edu/oupress, 405-364-5978 fax). $18.95, 384 pp, illustrated. 


AMANDA McQuabrE CraAwrorD. 1997. Herbal Remedies for Women: Discover Nature’s 
Wonderful Secrets just for Women. (ISBN 0-7615-0980-1, pbk.). Prima Pub- 
lishing, P.O. Box 1260BK, Rocklin, CA 95677, www.primahealth.com, 800- 
632-8676, 916-632-4400). $18.00, 291 pp. 


KATHRYN BERNICK. 1998. Basketry and Cordage from Hesquiat Harbour. (ISBN 0- 
7718-9525-9, pbk.). Royal British Columbia Museum, P.O. Box 9815 Stn Prov. 
Govt, 675 Belleville Street, Victoria, British Columbia, V8W 9W2,CANADA). 
$14.95, 152 pp, illustrated. 

Piers ViTessky. 2001. Shamanism. (ISBN 0-8061-3328-7, pbk.). University of Oklahoma 
Press, 4100 28" Avenue N.W, Norman, OK 73069-8218, wwwou.edu/oupress, 
405-364-5978 fax). $12.95, 128 pp, 250 color and b/w illustrations. 

AnpbrEO. BAREL, MArc Paye,and Howarp |. Marsacu (eds.). 2001. Handbook of Cosmetic 
Science and Technology. (ISBN 0-8247-0292-1, hbk.) Marcel Dekker, Inc. 
Cimarron Road, PO. Box 5005, Monticello, NY 12701-5185, www.dekkercom, 
212-696-9000, 212-685-4540 fax). $235.00, 886 pp. 


Botany/Molecular 


Bryan G. Bowes. 2000. A Color Atlas of Plant Structure (paperback edition). (ISBN 
0-8138-2693-4, pbk.). lowa State University Press, 2121 South State Street, 
Ames, [A 50014-8300, www.isupress.com, 800-862-6657, 515-292-0155, 515- 
292-3348 fax). $49.95, 192 pp, 380 + figures, color. 


Kincsey R. STERN. 2000. Introductory Plant Biology, Edition Eight. (ISBN 0-07- 
012205-9, hbk.). McGraw-Hill Companies, wwwimhhe.com, wwwmhhe.com/ 
botany, 800-262-4729). $67.00, 589 pp, illustrated. 


GorDON UNo, RICHARD Storey, and RANDY Moore. 2001. Principles of Botany. 
(ISBN 0-07-228592-3, pbk.). McGraw-Hill Companies, www.mbhe.com, 
www.mhhe.com/botany, 800-262-4729). $67.00, 589 pp, illustrated, CD- 
ROM included. 

Davip H. Benzinc. 2000. Bromeliaceae: Profile of an Adaptive Radiation. (ISBN 
0-521-43031-3, hbk.). Cambridge University Press, The Edinburgh Building, 
Cambridge CB2 2RU, UK, www.cu p.cam ac.uk). $120.00, 690 pp, illustrated. 


THomas J. GivnisH and KENNETH J. SytsMa (eds.). 1997. Molecular Evolution and Adaptive 
Radiation. (ISBN 0-521-57329-7, hbk.). Cambridge University Press, The 
Edinburgh Building, Cambridge CB2 2RU, UK, 40 West 20" Street, New 
York, NY 10011-4211, www.cup.cam.ac.uk). $39.95, 621 pp, illustrated. 


SIDA 19(3): 763. 2001 


764 BRIT.ORG/SIDA 19(3) 


CHRISTOPHER P. DuNN (ed.). 2000. The Elms: Breeding, Conservation, and Disease 
Management. (ISBN 0-7923-7724-9, hbk.). Kluwer Academic Publishers, 
PO. Box 989, 3300 AZ Dordrecht, The NETHERLANDS. $125.00, 361 pp, 
illustrated. 


WILLIAM Dupin. 2001. Soils. (SBN 0505-09148-4, pbk.). lowa State University Press, 
2121 South State Street, Ames, [A 50014-8300, www.isupress.com, 800-862- 
6057, 515-292-0155, 515-292-3348 fax). $19.95, 110 pp, illustrated. 

David P. Mappison and Wayne P. Mappison. 2000. MacClade 4. Analysis of Phy- 
logeny and Character Evolution. Software and Manual on CD-ROM. (ISBN 
0-87893-470-7,CD-ROM). Sinauer Associates, Inc., PO. Box 407, Sunderland, 
MA 01375-0407, 413-549-4300, 413-549-1118 fax). $125.00 Uncludes Class- 
room Version and all prior versions’ manuals). 


GERALD C. NELSON. 2001. Genetically Modified Organisms in Agriculture: Economics 
and Politics. ISBN Academic Press, 525 B Street, Suite 1900, San Diego, CA 
92101-4495, wwwacademicpress.com). Price not given, 344 pp. 


Ecology/Conservation/Landscape Ecology 


PETER FE Frotiiott, Luis A. BojyRQUEZ-T APIA, and MARIANO HERNANDEZ-NARVAEZ. 2001. 
Natural Resources Management Practices: A Primer. (ISBN 0-8138-2541- 
5, hbk.). lowa State University Press, 2121 South State Street, Ames, [A 50014- 
8300, www.isupress.com, 800-862-6657, 515-292-0155, 515-292-3348 fax). 
$52.95, 250 pp, illustrated, color. 

D.M,J.S. Bowman. 2000. Australian Rainforests: Islands of Green in a Land of Fire. 
(ISBN 0-521-46568-0, hbk.). Cambridge University Press, The Edinburgh 
Building, Cambridge CB2 2RU, UK, wwweup.cam.ac.uk). $85.00, 345 pp, 
illustrated. 


REED FE Noss (ed.). 2000. The Redwood Forest: History, Ecology, and the Conser- 
vation of the Coast Redwoods. (ISBN 1-55963-726-9, pbk.). Island Press, 1718 
Connecticut Avenue, N.W, Suite 300, Washington, DC 20009. (Orders: Is- 
land Press, P.O. Box 7, Dept. 2PR, Covelo,CA 95428, 800-828-1302). $30.00, 
352 pp, illustrated. 


BERNARD K. MALoneY (ed.). 1997. Human Activities and the Tropical Rainforest, 
Past, Present and Possible Future. (ISBN 0-7923-4858-3, hbk.). Kluwer 
Academic Publishers, PO. Box 989, 3300 AZ Dordrecht, The NETHERLANDS. 
$142.00, 206 pp, illustrated. 


Bruce A. STEIN, LYNN S. KUTNER, and JONATHAN S. ADAms (eds.). 2000. Precious Heri- 
tage: The Status of Biodiversity in the United States. (ISBN 0-19-512519-e, 


SIDA 19(3): 764. 2001 


BOOKS RECEIVED 765 


hbk.). Oxford University Press, 2001 Evans Road, Cary, NC 27513, wwwoup- 
usa.org, 800-451-7556). $45.00, 399 pp, illustrated, color. 


Grapy L. WessTER and ConraD J. BAHRE. 2001. Changing Plant Life of La Frontera. 
Observations on Vegetation in the United States/Mexico Borderlands. (ISBN 
0-8263-2239-5, hbk.). University of New Mexico Press, 3721 Spirit Drive 
S.E., Albuquerque, NM 87106-5631, 800-249-7737, 505-277-9270 fax). $60.00, 
272 pp, 41 halftones, 6 maps. 


RoLAND H. Wauer. 2001. Naturally ... South Texas. Nature Notes from the Coastal 
Bend. (ISBN 0-292-79139-9, pbk.). University of Texas Press, P.O. Box 7819, 
Austin, TX 78713-7819, 512-232-7634). $22.95, 256 pp, 29 line drawings. 

Ropney W. Bovey. 2001. Woody Plants and Woody Plant Management: Ecology, 
Safety, and Environmental Impact. (ISBN 0-8247-0438-X, hbk. (Marcel Dekker, 
Inc. Cimarron Road, P.O. Box 5005, Monticello, NY 12701-5185, 
wwwdekker.com, 212-696-9000, 212-685-4540 fax). $195.00, 564 pp. 


Horticulture/Gardening 


Mary F Iris. 2000. Gardening in the Desert. A Guide to Plant Selection and Care. 
(ISBN 0-8165-2057-7, pbk.). The University of Arizona Press, 355 S Euclid 
Ave., Suite 103, Tucson, AZ 85719, www.uapress.arizona.edu, 520-621-1441, 
520-621-8899 fax). $17.95, 210 pp, illustrated. 


Tommy Cairns, Marity YOUNG, JOLENE ADAs, and Bos Epserc (eds.). 2000. Modern 
Roses XI. The World Encyclopedia of Roses. (ISBN O-12-155053-2, hbk.). 
Academic Press, 6277 Sea Harbor Drive, Orlando, FL 32821-9816, 
www.academicpress.com, 800-894-3434 x6403, 619-699-6380 fax). $99.95, 
638 pp. 

PETER THOMPSON. 2001. The Looking-Glass Garden. Plants and Gardens of the Southern 
Hemisphere. (ISBN 0-88192-499-7, hbk.). Timber Press, 133 SW Second Ave., 
Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: wwwtim ber press.com, 
800-327-5680, 503-227-2878, 503-227-3070 fax). $39.95, hbk. 451 pp., 51 
color photos. 

THAD M. Howarp. 2001. Bulbs for Warm Climates. (ISBN 0-292-73125-6, hbk.). 
University of Texas Press, PO. Box 7819, Austin, TX 78713-7819, 512-232- 
7634). $60.00, 288 pp, 179 color photos. 

ALAN Toocoop (ed.) and Peter ANDERSON (Photography). 1999. Plant Propagation. 
(ISBN 0-7894-4116-0, hbk.). DK Publishing, Inc., 95 Madison Avenue, New 
York, NY 10016, www.dk.com). $34.95, 320 pp, illustrated. 


SIDA 19(3): 765. 2001 


766 BRIT.ORG/SIDA 19 


=a 


3) 


Jit Nokes. 2001. How to Grow Native Plants of Texas and the Southwest. Revised 
and Updated Edition. (ISBN 0-292-75574-0, hbk.). University of Texas Press, 
PO. Box 7819, Austin, TX 78713-7819, 512-232-7634). $60.00, 632 pp, 30 color 
photos, 20 b/w illustrations. 


RICHARD L. Duse. 2001. Turfgrasses: Their Management and Use in the Southern 
Zone, Second Edition. (ISBN 1-58544-161-9, pbk.). Texas AGM University, 
4354 TAMU, College Station, TX 77843-4354, www.tamu.edu/upress, 979- 
458-3982, 979-847-8752 fax). $27.95, 336 pp, 51 color, 15 b/w photos, 68 line 
drawings. 


Juces JANick (ed.). 1999, Horticultural Reviews, Volume 23. (ISBN 0-471-25445-2, 
hbk., ISSN 0163-7851). John Wiley & Sons, Inc., One Wiley, Drive, Somerset, 
NJ 08875). Price not given, 366 pp. 

Biographical 

CHRISTOPHER KELLy (ed.). 2000. Jean-Jacques Rousseau. The Reveries of the Solitary 
Walker, Botanical Writings, and Letter to Franquiéres. Translated and 
Annotated by Charles E. Butterworth, Alexandra Cook, and Terence E. 
Marshall. ISBN 1-58465-007-9, hbk.). Dartmouth College, University Press 
of New England, Hanover, NH 03755, 603-643-7100. (Orders: Plymbridge 
Distributors Ltd., Estover Plymouth, PL6 &P2, UK). £47.00 (about $60.00), 
349 pp. 


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ISSN 0036-1488 


TABLE OF CONTENTS 


TWO NEW SPECIES OF LIATRIS SERIES PUNCTATAE (AS =) CENTERED IN NORTH CENTRAL TEXAS 


G.L. NESOM AND RJ, O'KENNON—767 


LAENNECIA TURNERORUM (ASTERACEAE: ASTEREAE), A NEW SPECIES FROM TRANS-PECOS TEXAS 
G.L. NEsomM—789 


TWO NEW SPECIES OF IMPATIENS (BALSAMINACEAE) FROM INDIA 
MUKTESH KUMAR AND STEPHEN SEQUIERA—795 


ANOTHER NEW SPECIES OF TRIPOGON (POACEAE) FROM INDIA 
C.N. SUNIL AND A.K. PRabEEP—803 


IPOMOEA SORORIA (CONVOLVULACEAE), A NEW SPECIES FROM YUCATAN, MEXICO 
DANiet F. AUSTIN AND Jose Luis TAPIA MUNO2—807 


TAXONOMIC REVIEW OF CHE NUM (ASTERACEAE: HELIANTHEAE) 
G.L.N 11 


TAXONOMY OF THE DICHOTOMA GROUP OF DICHANTHELIUM (POACEAE) 
RICHARD J. LEBLOND—-821 


THE GENUS HESPEROYUCCA (AGAVACEAE) IN THE WESTERN UNITED STATES AND MEXICO: NEW NOMENCLATURAL COMBINATIONS 
KAREN H. CLARY —839 


NEW COMBINATIONS IN CHIONOLAENA (ASTERACEAE: GNAPHALIEAE) 
G.L. Nesom—849 


SYSTEMATICS OF THE CAREX JAMESII COMPLEX (CYPERACEAE: SECT. PHYLLOSTACHYAE) 
Ropert F.C. NACZI AND Bruce A. FoRD—853 


Ae: 


GENETIC DIVERSITY IN THE CAREX JAMESII COMPLEX (CYPERACEAE: SECT, PHYLLOSTACHYAE 
WITH INSIGHTS INTO THE EVOLUTION AND ORIGIN OF THE NEWLY DESCRIBED SPECIES CAREX TIMIDA 
Bruce A. FORD AND ROBERT F.C. NACZI—885 


STENOTIS (RUBIACEAE), A NEW SEGREGATE GENUS FROM BAJA CALIFORNIA, MEXICO 
EDWARD E. TERRELL—899 


TAXONOMIC REVIEW OF HOUSTONIA ACEROSA AND H. PALMERI, WITH NOTES ON HEDYOTIS AND OLDENLANDIA (RUBIACEAE) 
Epwarb E. TeRRELL—-913 


NOMENCLATURAL CHANGE IN THE DIGITARIA COGNATA COMPLEX (POACEAE: PANICEAE) 
JOSEPH K. WipFF—923 


NEOTYPIFICATION OF ENSLENIA ALBIDA AND A NEW COMBINATION IN AMPELAMUS FOR CYNANCHUM LAEVE 
(APOCYNACEAE: ASCLEPIADOIDEAE) 
ALEXANDER KRINGS—925 


PRELIMINARY RESULTS TOWARD A REVISION OF THE AMARANTHUS HYBRIDUS SPECIES COMPLEX (AMARANTHACEAE) 
MIHAI CosTEA, ANDREW SANDERS, AND GILES WAINES—931 


NOTES ON SOME LITTLE KNOWN AMARANTHUS TAXA (AMARANTHACEAE) IN THE UNITED STATES 
MIHAI CosTea, ANDREW SANDERS, AND GILES WaAINES—975 


CAREX CUMBERLANDENSIS, A NEW SPECIES OF SECTION CAREYANAE (CYPERACEAE) 
FROM THE EASTERN UNITED STATES OF AMERICA 

Ropert F.C. Naczi, RoBerT KRAL, AND CHARLES T. BRYSON—993 

FLorisTics 


A SYNOPSIS OF THE FERNS AND FERN ALLIES OF NEBRASKA, WITH MAPS OF THEIR DISTRIBUTION 
STEVEN B. ROLFSMEIER, ROBERT B. KAUL, AND David M. SUTHERLAND—1015 


IN THE SAN Marcos River, TEXAS 


NS 


EXPANSION OF THE EXOTIC AQUATIC PLANT CRYPTOCORYNE BECKETTII (ARACEAE 
Rosert D. DoYLE—1027 


VASCULAR PLANT TYPES IN THE ARIZONA STATE UNIVERSITY HERBARIUM 
STEFANIE M. ICKERT-BOND AND DONALD J. PINKAVA—1039 


PLANT SPECIES-AREA RELATIONSHIPS IN TEN NORTH CENTRAL TEXAS PROTECTED NATURAL AREAS 
MONICA GRANADOS, RoserT J. O’KENNON, AND Bruce F. BeNz7—1061 


STATUS OF QUERCUS X LEANA AND QUERCUS X RUNCINATA (FAGACEAE) IN ILLINOIS 
GORDON C. TUCKER AND JOHN E, EBINGER—1073 


THE VASCULAR FLORA OF MADISON COUNTY, TEXAS 
AMANDA K. NEILL AND HUGH D. WiLson—1083 


2 SIDERACIONES SOBRE EL ORIGEN DE LA FLORA ARVENSE Y RUDERAL DEL ESTADO DE QuUERETARO, MEXICO 
SE AURELIO COLMENERO ROBLES, CONCEPCION RODRIGUEZ JIMENEZ, AND FERNANDEZ Nava RAFAEL —1123 


VINES OF A TEMPERATE STATE: STILL UNDERCOLLECTED? 
ALEXANDER KRINGS—1 147 


A NEW SYNONYM FOR ERAGROSTIS PILGERI (POACEAE: ERAGROSTIDEAE) 
J. Jost ALEGRIA OLVERA AND ARTURO GRANDA PAUCAR—1157 


SCLERIA LACUSTRIS (CYPERACEAE), AN AQUATIC AND WETLAND SEDGE INTRODUCED TO FLORIDA 
LETTE C. JACONO—1163 


SOUTH AMERICAN SKULLCAP (SCUTELLARIA RACEMOSA: LAMIACEAE) IN THE SOUTHEASTERN UNITED STATES 
ALEXANDER KRINGS AND JOSEPH C. NEAL—1171 


ee BONDUC (FABACEAE) NEW TO LOUISIANA 
LUAM G. VERMILLION—1181 


HOUSTONIA LONGIFOLIA (RUBIACEAE): NEWLY DOCUMENTED FOR THE FLORA OF TEXAS 
JASON R. SINGHURST AND W.C. HOLMeS—1183 


NEW RECORDS IN PSEUDOGNAPHALIUM (ASTERACEAE: GNAPHALIEAE) FOR THE UNITED STATES 
G.L. NESoMm—1185 


ARTHRAXON (POACEAE: ANDROPOGONEAE) NEW TO SOUTH AMERICA 
LJ. Dorr AND S. MicueL NiINO—1191 


ANAGALLIS ARVENSIS SUBSP. FOEMINA (PRIMULACEAE) NEW TO LOUISIANA 
CHARLES M. ALLEN, SARA THAMES, PHILLIP PAUL, AND SELENA DAWN NewmMaN—1195 


ALTERNANTHERA SESSILIS (AMARANTHACEAE) NEW TO ARKANSAS 
CHRISTOPHER S. REID AND DAVID X WILLIAMS—1197 


VASCULAR PLANTS NEW TO KENTUCKY 
J. RICHARD ABBOTT, RALPH L. THOMPSON, AND Ruby A. GELIS—1199 


BOOK REVIEWS AND NOTICES 788, 794, 802, 838, 848, 898 912,930, 1060, 1082, 1122, 1146, 1156, 1162, 1180 
1194 1198, 1203-1210 


SIDA, CONTRIBUTIONS TO BOTANY UPDATE —1211 
REVIEWERS FOR VOLUME 19, 2000-2001—1212 


INDEX TO VOLUME 19, 2000-2001—1213 
TITLES OF ARTICLES WITH AUTHORS—1213 
AUTHOR—1218 
BOTANICAL NAMES—1220 
SUBJECTI—1226 
NEW NAMES AND NEW COMBINATIONS—1228 


SIDA GUIDELINES FOR CONTRIBUTORS— 1231 


INDEX TO NEW NAMES AND NEW COMBINATIONS IN SIDA 19(4), 2001. 


AMARANTHUS BLITUM VAR. PSEUDOGRACILIS ( THELL.) COSTEA, COMB. ET STAT. NOV.—981 
AMARANTHUS BLITUM SUBSP. OLERACEUS (L.) COSTEA, COMB. NOV.— 984 

AMARANTHUS HYBRIDUS SUBSP. QUITENSIS (KUNTH) COSTEA & CARRETERO, COMB. NOV.—955 
AMARANTHUS POWELLII SUBSP. BOUCHONII (T HELL.) COSTEA & CARRETERO, COMB. NOV.—964 
AMPELAMUS LAEVIS (MICHX.) KRINGS, COMB. NOV.—927 

CAREX CUMBERLANDENSIS NACZI, KRAL, & BRYSON, SP. NOV.—994 

CAREX TIMIDA NACZ! & B.A. FORD, SP. NOV.—879 

CHIONOLAENA COSTARICENSIS (NESOM) NESOM, COMB. NOV.—850 

CHIONOLAENA CRYPTOCEPHALA (Nesom) Nesom, COMB. NOV.—850 

CHIONOLAENA DURANGENSIS (Nesom) Nesom, COMB. NOv.— 850 

CHIONOLAENA MACDONALDII (Nesom) Nesom, ComB. NOV.— 850 

CHIONOLAENA SALICIFOLIA (BERTOL.) NESOM, COMB. NOV.—850 

CHRYSOGONUM VIRGINIANUM VAR. BREVISTOLON NESOM, VAR. NOV.—817 

DICHANTHELIUM ANNULUM (ASHE) RJ. LEBLOND, COMB. NOV.—826 

DICHANTHELIUM DICHOTOMUM (L.) GOULD VAR. NITIDUM (LAM.) RJ. LEBLOND, COMB. NOV.—829 
DICHANTHELIUM DICHOTOMUM (L.) GOULD VAR. RAMULOSUM ( Torr.) RJ. LEBLOND, COMB. NOV.—830 
DICHANTHELIUM DICHOTOMUM (L.) GOULD VAR. ROANOKENSE (ASHE) RJ. LEBLOND, COMB. NOV,—83 1 
DICHANTHELIUM LUCIDUM (ASHE) R.J. LEBLOND, COMB. NOV,—83 1 

DICHANTHELIUM SPHAGNICOLA (NASH) R.J. LEBLOND, COMB. NOV,— 834 

DIGITARIA PUBIFLORA (VASEY) WIPFF, COMB. ET STAT. NOV.—923 

HESPEROYUCCA NEWBERRY! (MCKELVEY) CLARY, COMB. NOV.—845 

HESPEROYUCCA PENINSULARIS (MCKELVEY) CLARY, COMB. NOV.—8 45 

HOUSTONIA ACEROSA (A. GRAY) BENTHAM & HOOKER F. VAR. POLYPREMOIDES (A. GRAY) TERRELL, COMB. NOV.—916 
HOUSTONIA ACEROSA (A. GRAY) BENTHAM & HOOKER F. VAR. TAMAULIPANA (B.L. TURNER) TERRELL, COMB. NOV.—916 
HOUSTONIA PALMERI VAR. MUZQUIZANA (B.L. TURNER) TERRELL, COMB. NOV.—919 

IMPATIENS SHOLAYARENSIS M. KUMAR & SEQUIERA, SP. NOV.— 

IMPATIENS VIOLACEA M. KUMAR & SEQUIERA, SP. NOV.—798 

IPOMOEA SORORIA D.F. AUSTIN & J.L. TAPIA, SP. NOV.—807 

LAENNECIA TURNERORUM NESOM, SP. NOV.— 789 

LIATRIS AESTIVALIS NESOM & O'KENNON, SP. NOV.—768 

LIATRIS GLANDULOSA NESOM & O’KENNON, SP. NOV.—778 

STENOTIS TERRELL, GEN. NOV.—901 

STENOTIS ARENARIA (ROSE) TERRELL, COMB.NOV.—902 

STENOTIS ASPERULOIDES (BENTH.) TERRELL, COMB. NOV.—903 

STENOTIS . BRANDEGEANA (ROSE) TERRELL, COMB. NOV.—904 

STENOTIS AUSTRALIS (1.M. JOHNST.) TERRELL, COMB. NOV.—906 

STENOTIS BREVIPES (ROSE) TERRELL, COMB. NOV,—907 

S (1.M. JOHNST.) TERRELL, COMB. NOV.—908 

STENOTIS MUCRONATA (BENTH.) TERRELL, COMB. NOV.—909 

STENOTIS PENINSULARIS (BRANDEGEE) TERRELL, COMB. NOV.—910 

TRIPOGON RAVIANUS SUNIL & PRADEEP, SP. NOV.— 


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TWO NEW SPECIES OF LIATRIS SERIES PUNCTATAE 
(ASTERACEAE: EUPATORIEAE) CENTERED IN 
NORTH CENTRAL TEXAS 
G.L.Nesom and RJ.O’Kennon 
Botanical Research Institute of Texas 
509 Pecan Street 
Fort Worth, TX 76102-4060, U.S.A. 

ABSTRACT 


Liatris aestivalis, sp. nov. is recognized from Texas and OMahome as a species separate from L. 
mucronata DC. and L. punctata Hook., although it | 


with both of the latter as L. 


angustifolia (Bush) Gaiser. The type of L. angustifolia isa pl best identified as L. mucronata, perhaps 
showing some paEMEnCe of Eonar avon with L. aest ee Liatris aestivalis prod SEPEIObos 
corms and has mostly | in foli dcapi features 


as well as phenology. The new species is known from 16 counties primarily along a nor rile south band 
from central counties of Texas into south-central Oklahoma; it is sympatric with L. mucronata and 
the two sometimes grow in close ¢ association, but anes are come) ee to Pay ov sae 
pnge in Rowers oe P 


their Liatris glandul sp. nov. has | 1 d f five localities i pale 


County but is known to ~ extant at only a amale site pags it ale has been cena in three other 
Texas counties Soulard a is te arto tol | 
weakl 


] 
UL iangular, 


yg I all species of the genus in its vestiture e of SuDMAle glan- 
dular hairs. Liatrisglandul lsoi ly-flowering pletely fr 


ele 


L. mucronata, which grows in close sympatry at the Dallas Caan ae A description: istration 
distribution map, and key contrast with L. mucronata are provided here for each of the new species. 


RESUMEN 


Liatris aestivalis, sp. nov.,se reconoce de Texas y Oklah dade L. mucronata 
D nctata Hook., aunque? na sido Suomi aaea | con estas dos diltimas como L. angustifolia 
(Bush) ee El tipo de L L. mucronata, quizas 
muestre algo de influencia de iibedacion con L. aestivalis. Liatris destivalis produce bulbos 
soup Eey: yi 


a mayoria de las veces como L. mucronata ee difiere de ee ues 
foliares y del capitulo asi como en la ee — nueva 16 
condados situados a lo largo de una banda norte-sur desde los trales de Texas hasta el 
sur-centro de Oklahoma; es simpatrica con L. mucronata y ambas viven a menudo en intima 
asociacion, pero estan completamente separadas o solo ligeramente solapadas en el periodo de 
floracion. Se han identificado intermedios putativos en varios condados de la regi6n donde su 
lee sgt ea ] Se hath, ay cn ea | 4 ] Bin A ] ee eee ] pee 


L o 
de Dall Slo p llien una tnica alocelidad: aie bg ico colectada en 
otros tres condadosde T hacia el Es similar a 1 I g filarios 


it Paes are | 


género por 
su ineuments de pels espace: glandulares ene glanaulose tambien plone antes, casi 
L.mucronata, q I la localidad 


del condadode Dallas Se ofrecen seu descripcion il id pa de distribucion, y una 


clave que contrasta L. mucronata con las nuevas specs 


SIDA 19(4): 767 — 787, 2001 


768 BRIT.ORG/SIDA 19(4) 


Field evidence and review of herbarium collections have revealed the presence 
of two previously undescribed species of Liatris, both of which are geographi- 
cally centered in north central Texas. For the first, field observations led to re- 
evaluation of species definitions in the field and herbarium. For the second, 
review of earlier herbarium collections led to recognition of a distinctive spe- 
cies and field confirmation that it is still extant. 

Contemporary botanists in north central Texas have observed the occur- 
rence of early-flowering plants similar to Liatris mucronata DC. but distinct 
from it. Gaiser (1946, 1950) recognized these ‘non-mucronata plants as L. 
angustifolia (Bush) Gaiser, but her morphological and geographic definition of 
the taxon was diffuse. In contrast, L. angustifolia was treated by Shinners (1951), 
Menhusen (1963), and in the Flora of Texas (Correll @ Johnston 1970) as a syn- 
onym of L. mucronata and ina survey of the Oklahoma flora (Waterfall 1969) 
as asynonym of L. punctata var. nebraskensis Gaiser. The name L. angustifolia 
has been sporadically applied by original identification and annotation to a 
few of the collections cited in the present manuscript, but the name never gained 
currency as representative of a valid species. In other recent floras and check- 
lists of Texas and Oklahoma (Hatch et al. 1990; Johnston 1990; Taylor & Taylor 
1994; Jones et al. 1997; Diggs et al. 1999), these plants are identified as L. 
mucronata and L. angustifolia has not been included even asa synonym. Kartesz 
(1999) treated L. angustifolia asa synonym of L. densispicata (Bush) Gaiser ‘var. 
interrupta Gaiser (the correct disposition of ‘var. interrupta is as L.mucronata 
var. interrupta Gaiser). 

With the benefit of field observations of phenologically distinct entities 
around Fort Worth and other north central Texas localities, we sorted collec- 
tions of Liatris mucronata sensu lato from BAYLU, SMU/BRIT, TEX/LL, MO, 
OKL, and OKLA~arriving at a reasonable definition of two morphological 
groups, witha third group of possible intermediates. We examined types of taxa 
that held the possibility of representing the generall 1 (or diffusely 
recognized) entity from north central Texas and found that aname has not been 
correctly applied to it. In this report we provide a name for the species as well as 
means of identification and documentation of its geographical extent. 


Liatris aestivalis Nesom & O’Kennon, sp. nov. (Figs. 1, 2, 6). TyPr: U.S.A. TEXAS. 
TARRANT Co. W side of city of Fort Worth, undeveloped area (advertised for sale) with rock 


oer near corner of Green Oaks Drive and Genoa Read. NE corner of Ridgmar Mall area, 


il Glen Rose (C jor associates Yucca pallida, Silphium 
albiflo orum, Hedyotis nigricans, Ceanothus herbaceus, Sorsiactann nutans, Schizachyrium 
scoparium, Polygala alba, Salvia texana, Stenosiphon linifolius, Grindelia lanceolata var. 
texana, Phyllanthus sedi Paronychia virginica, Oenothera macrocarpa, Vernonia 
lindheimeri, Scutellaria wrightii, Dalea hallii, with Liatris mucronata common but gener- 
ally in deeper soil, iG in early re Liatris aestivalis abundant on flats and exposed a. 
ah little competition, plants in early to mid-flower, 8 Jul 2001, Guy L. Nesom FWIS, Robert]. 
O’Kennon, and George M. Diggs (HOLOTYPE: BRIT; IsoTyPES: BAYLU, COLO, F, FLAS, GA, GH, 


NESOM AND O’KENNON, NEW SPECIES OF LIATRIS FROM NORTH CENTRAL TEXAS 769 


ISC, KANU, KSC, MIN, MO, NCU, NLU, NY, OKL, RM, SBSC, TAES, TENN, TEX, UARK, UC, 
UNM, US-to be distributed). 

Differta L. mucronata DC. foliis inferis | ioril hyllarii ioril ioribus leniter 2-3- 

seriatis, et florescentia praecici. 


Plants 20-65 cm tall (avg. ca. 30 cm), with 1-2(-5) erect stems arising from a 
globose or subglobose corm 13-30 mm in diameter; stems, leaves, and phyllar- 
ies densely punctate with imbedded, clear-resinous glands, otherwise mostly 
glabrous. Leaves crowded, spreading-ascending, 0.6-2.5 mm wide, the lower 7- 
15 cm long, shortened upward, usually ca. 1/3 as long at midstem and half as 
wide, usually reduced to short, spreading bracts in the lowermost part of the 
capitulescence, margins smooth to minutely papillate or sparsely short-ciliate, 
the narrower not punctate, the broader inconspicuously punctate, otherwise 
glabrous. Capitulescence 4-15 cm long, ca. 13-30 mm wide, capitula densely 
arranged, upper cauline leaves sometimes continuing upward as capitular 
bracts not wider than the capitulescence. Capitula turbinate-cylindric, 9-12 
long, 3.5-5 wide; phyllaries 6-11 per head, in 2-3 series weakly graduate in 
length, the outermost ca. 1/2-2/3 the length of the inner, inner 11-12 mm long, 
oblong-lanceolate to lanceolate-triangular, acute to acute-attenuate at the apex, 
not abruptly mucronate or cuspidate but sometimes with a long, narrow tip, 
commonly becoming dark purple in anthesis, glabrous or the outer sparsely 
ciliate. Florets 3-4(-5) per head; corollas pink-purple, sessile-glandular, 6-10 
mm long, tube 4-7 mm long, glabrate within, lobes lanceolate, 1.5-3 mm long; 
filaments sparsely puberulent-pilose. Cypselae 4.5-6 mm long, oblong-oblan- 
ceolate in outline, compressed to distinctly 3-angled, with 9-11 longitudinal 
nerves, densely hairy on nerves and interstices with duplex hairs (0.2-)0.4-0.6 
mm long, sometimes minutely sessile-glandular, pappus of plumose bristles 
5.5-8 mm long, ca. equal the corolla length. Chromosome number unknown. 

Etymology.—The epithet alludes to the summer flowering of the species, a 
distinctive feature compared to L. mucronata. 


Additional collections examined: U.S.A. OKLAHOMA. Atoka Co.: 3.0 mi N of Caddo, rocky hillside, 11 
Jul 1967, Johnson 172 (OKL). Bryan Co.: 2 mi W, 1.5 mi N of Matoy, pine-oak woods, 23 Jul 1962, C. & J. 
Taylor 916 (OKL). Johnston Co.: 2.5 mi W and | mi Nof Mannsville, on Baum Limestone outcrop, Jun 
1960, Taylor s.n. (OKL). Love Co.: 6.5 mi NE of jet OK 77 on OK 32 (ca.8 mi E of Marietta), limestone 
outcrop, eroded sandy slope, post oak-juniper woods, 270 m elev. 14 Aug 1979, Wagner a sgeues 


4070 te cee Marshall Co.: 1 mi SE of Madill, prairie, 21 Jul 1959, Goodman 6921 (OKL, SMU). 
airie near Sulphur, 5 Aug 1939, Broad ie 598 (OKL); Platt National * Cold 
ae near ed 20 Jun 1935 [early flower], Merrill 703 (MO), Platt National Park, Flower Park, 


near oa 30 Sep 1935 [past flower], Merrill 15. te TEXAS. Anderson Co.: ca. 350 m S of jet Co. 
Ro n Co. Road 2202, N 3173987, W 95.73025, limestone outcrop, 2 Jun 1999, Holmes 10140 
ne : mucronata eallened at the same site on 25 Oct 1997, Singhurst 6132a (BAYLU). Cooke 
Co.: near Tyler Bluff, W edge of county, blackland prairies, 2 Jul 1946, Correll and Correll 12961(SMU), 
NE side of St. Jo, Co. Road 414—Brushy Mound Road, W of FM 2382, calcareous soil, 7 Aug 1995, Lusk 
s.n. (BRIT); Don Vogel Ranch (Elm Fork of the Trinity River), Co. Road 302, 2 mi E of Montague Co. 
line, 13 Aug 1995, Lusk s.n. (BRIT); Cooke Co.: County Road 343 0.5 mi S of FM 1630, Freemound 


770 BRIT.ORG/SIDA 19(4) 


Community, growing in close proximity to L. mucronata (also see 22 Jul collection of putative inter- 
mediate L. mucronata x L. aestivalis made from nearby locality), 14 Jul 2001, Lusk s.n. (BRIT); 3 miS 
of Eraon FM 51, ranch of J. David Thomas, M.D., 16 Jul 2001, Lusk s.n. (BRIT); Marysville Community, 
Ll mi Sof Co. Road oe on E side of Co. Road 417 (0.7 mi S of N. Fish Creek bridge), 2 Aug 2001, Lusk 
sn. (BRIT). Coryell Co.: 5 mi W of Gatesville, prairie, 23 Aug 1945, Muller 8732 (LL, MIN, SMU). Den- 
ton Co.: 15.5 mi W of ben gravelly (limestone) shoulder above stream, 9 Oct 1949 [end of fruit- 


ing], Shinners 11892 (SMU); between Sanger and Denton on 1-35, along service road, 0.7 mi N of FM 
3163 (Milam Road), S mucronata growing nearby, (local 2 found by Jeff Quayle), 17 Jul 2001, Lusk 
s.n. (BRIT). Hays Co.: 2 mi SE of Wimberly, across road from entrance to Hidden Valley Ranch, com- 


mon locally on shallow limestone slopes, 30 Aug 1948 very end of f aosarc ine 383 (TEX). 
Montague Co.: Hugh Garnett Ranch, ca. 4 mi SW of Dye Mound on FM 3206, 7 Aug 1995, Lusk s.n. 
(BRIT); Don Vogel’s Hardy Ranch, FM 1630 at FM 677, in Hardy community, SE corner of the county, 
L. sane growing nearby, 14 Jul 2001, Lusk s.n. (BRIT); Don Vogel's L er Ranch, a ‘ield 
Road at Gladys Community, E of FM 677, growing with L. squarrosa, 14 Jul 2001, Lusk s.n. (BRIT); 
Jane D. Lucas Ranch, FM Road 677, 5 mi S of jct with US Hwy 82 in center : aun. 14 Jul 2001, 
Lusk s.n. (BRIT); Garnett property, ca. 4 mi S of Dye Mound on Dye Mound Road, ca. 8 air mi SW of 
Saint Jo, aad soil over limestone, L. aestivalis in early flower and common on rocky slopes, L. 
mucronata abundant but none yet in flower, 10 Jul 2001, Nesom FW16 with Garnett, Lusk, Miller. 
O’Kennon, and ome (BRIT, TEX); Nored Road, between Farm pee 455 and Dye Mound Road, ca. 
lL air mi SSW of Saint Jo, ae along road, exposed li llow sand 1, L. aestivalis 
abundant, inearly flower, 10 Jul 2001, Nesom F W390 with Garnett, Lusk, Miller, O’ fedeer Ss Quayle 
(BRIT); site of old Dye Community, Farm Road 3206 at jet with Dye Mound Road, ca. 7 mi SW of 
Saint Jo on FR 3206; steep, oe limestone slopes, L. aestivalis in early flower ae ee on 
open slopes, L. mucronata common, not yet in flower, 10 Jul 2001, Nesom FW44 with Garnett, Lusk, 
Miller, O’Kennon, and Quayle (BRIT, TEX); Clear Creek Loop (road), between Farm Road 3206 and 
677, ca. 0.6 mi SSE of jct with Farm Road 3206, ca. Sair mi S o nant Jo, open roadside with exposed 


> 


hy 


limestone outcrop and shallow soil, nearly clay-sand ide, L. destivalis relatively com- 
mon, barely into flower, 10 Jul 2001, Nesom FW46 with Garnett Lusk, Miller, O’Kennon, and Quayle 

(BRIT, KSC); Clear Creek Loop (road), between Farm Road 3206 and 677, ca. 1 mi SW of jet with Farm 
Road 677, ca. 2.5 air mi Sol paint Jo, opt poadlas with exposed limestone outcrop and shallow soil, 


L. aestivalis relatively c ) r,1OJul ae Nesom FW47 with Garnett, Lusk, Miller, 
O’Kennon, Bs Quayle (BRIT, TEX) Parker Co.: Ranch House Road 200 meters north of Vista Road 
near Lake Weatherford, deep sand, L. mucronata also in immediate area but not in flower, 6 Aug 


Ol, om 15007 and Jetf Quayle (BRIT, TEX). Tarrant Co.: [no other locality data], | Aug 1926, 
Killian 6577 (LL), WSW part of city of Fort Worth, between Cumberland St. and Vickery Blvd, just N 
of Vickery, E of Ridglea oe ees eae Lae owned land ee Hills Park), open 


area of limestone slopes, slopes and not yet in 1 flower, 
12 Jul 2001, Nesom FW48 and O’Kennon (BRIT, sll SW part st au of Fort Worth, N side 
arene Blvd., 0.7 mi NE of jet with Hwy 377, just N of jct Hwy 377 and 1H 20, open ae 
liv N side of road, a Ree a at edge of public land (Z. Boaz South 
aa ae 2001, Nesom F Wsi and O’Kennon (BRIT, KSC); SW part of city of Fort Worth, Lakeview 
Road just W of Benbrook Lake, 0.3-0.4 mi E of jct with 377, open limestone outcrop, e 
slope, this habitat hee continous for ca. 0.3 mi eastward along Lakeview Road, L. mucronata 
abundant, long before flowering, 12 Jul 2001, Nesom FW53 and O’Kennon (BRIT), W portion of City 
of Fort Worth, Westridge ei ca. 2 blocks N 2 a ie see Blvd., undeveloped lot with open, 
steep slopes of Glen R | silty soil, 22 Jul 2001, Nesom FW55(BAYLU, 
BRIT, KSC, TEX); Tandy Hills Municipal Parke ca. 2 air mi E of head n Fort Worth, N of Hwy 30, 
limestone slopes with shallow to deeper soil on flats and slope bottoms, L. mucronata abundant on 
ower slopes in deeper soil but not yet in flower, 22 Jul 2001, Nesom FW56 and O’Kennon (BRIT, KSC, 
., TEX, US); Fe. Worth, 26 Jul 1929, Whitehouse s.n. (TEX). Travis Co.: rocky hillsides and summits 


NESOM AND O’KENNON, NEW SPECIES OF LIATRIS FROM NORTH CENTRAL TEXAS 771 


around Austin, | Aug 1926, Tharp s.n.(SMU). Wise Co.: dry gravelly prairies on steep sandstone slopes, 
on E ee of co. road E of Big Sandy Creek, ca. 4 air mi NE of Alvord and ca. 3.8 air mi ESE of Park 
Springs, Post Oak Ridge, LBJ Natl. Grassland, 850-930 ft, Antlers Sand Formation, 1 Aug 1989, Orzell 
and stm 11265 (TEX); Te N sie a of Bald Knob Cemetery, ca. | air mi S of LBJ Natl. Grassland 
field headquarters, ca. 3.3 a of Alvord and 10 air mi NNW of Decatur, Bald Knob - lL BJ 
Natl. Grassland, shallow soil eer prairies/outcrops, 1100-1180 ft, 1 Aug 1989, Orzell lges 
1125/ (BRIT, TEA). 


Gaiser (1946, p. 370) noted that “confusing intermediates” apparently are formed 
between Liatris “angustifolia” and L. mucronata, especially as “the centre of the 
range for both of these species lies in central Texas.” A group of plants puta- 
tively intermediate between L. aestivalis and L. mucronata is recognized here: 
most of these plants have foliar features similar to L. mucronata and a large num- 
ber of phyllaries ((11-)13-18) strongly graduate in length, like those of L. 
mucronata; the flowering period of these populations ranges from July through 
mid-October (the flowering dates of these collections are not included in Table 
1). In contrast, these plants have relatively long phyllaries, at the upper range 
for L. mucronata, the inner with longer, more attenuate apices than typical for 
L. mucronata, more similar to those of L. aestivalis. The geographic coherence 
of the putative ‘intermediates’ (as cited below) suggests that they may have a 
similar evolutionary origin. Asa tentative identification, these collections have 
been annotated as “L. mucronata possibly > L. aestivalis,” although their distri- 
bution ranges somewhat outside of the known range of L. aestivalis (compare 
Figs. 5 and 6). The type of L. angustifolia (provenance uncertain, see comments 
below) is similar to plants of this group. 

Nesom FW57 (intermediate, from Tandy Hills in Fort Worth) was flower- 
ing at the same time as typical Liatris aestivalis (Nesom FW56) at the same site 
but considerably before L. mucronata Nesom Pye) a this locality, L. 
mucronata and the putative intermedi hed in the field) grew 
in relatively deeper soil and were both more abundant than the typical Ee 
aestivalis, which occurred only ona laine small area of limestone outcrop. 
Only three major phenotypes id mong the Tandy Hill plants—those 
putatively intermediate were obvious but ae of introgression was not. 
Lusk s.n. (Cooke Co., Tex.) also was flowering at the same time as L. aestivalis at 
the same site (Lusk s.n.,above) but well before L. mucronata (not collected). 


POSSIBLE INTERMEDIATES BET WEEN L. AESTIVALIS and L. ae OKLAHOMA. Love Co.: 
ni E of Marietta, roadside cut W of Hickory Creek, 1 Aug 1958 13 (OKL). TEXAS. Bosque 
174, 3.6 mi SW of Walnut Springs, 25 Sep 1950, Cory 58025 (SMU); Clay Co.: 5.2 mi E of 
a Hwy 82,6 Oct 1945, ies 10848 (SMU). Cooke Co.: Cla Hamilton Freemound Ranch, 
1.8 mi S of FM 1630, 0.8 mi E of gate on County Road 343, these plants appa ly intermediate be- 
tween L. mucronata (typical plants nearby) and L. aestivalis, 22 Jul ae Lusk s.n. (BRIT); J. & M. 
pera Ranch, Co. Rd. 414, 0.1 mi W of Brushy Mound Cemetery on S side, NE of Saint Jo, 28 Jul 2001, 
ce s.n. (BRIT). Dallas Co.: S side of Tenison Park, 9 Sep 1950, Cory 57909 ae Dallas, 10 Jul 1925, 

Hynes s.n.(TEX); off Northwest Highway, E of Vickery, 7 Sep 1940, Longnecker 79 (SMU). Fannin Co.; 
ca. 3 mi SE of Bonham, among grasses in open, scrub oak-cedar hills of Bonham State Park, 16 Sep 


BRIT.ORG/SIDA 19(4) 


772 


Wek Lae 
AGRA 
PSG = me 
r& Ske =. — > = a ha 
= —— = gta, oe 


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NESOM AND O’KENNON, NEW SPECIES OF LIATRIS FROM NORTH CENTRAL TEXAS 773 


Fic. 2. Habit and details of Liatris aestivalis. 


774 BRIT.ORG/SIDA 19(4) 


1960, Correll 23498 (LL). Grayson Co.: 6 mi SE of center of Sherman, remnant prairie vegetation, 
Chamberlain’s Ridge substation, 23 Sep 1993, Nee 43947 (MO). Hood Co.: Hwy 171, 2.8 mi NW of 
Cresson, 30 Sep 1950, Cory 58481 (SMU). Montague Co.: 10 mi E of St. Joe, 3 Oct 1975, Lipscomb 554 
(SMU). Navarro Co.: 15.7 mi SE of Corsicana on Hwy 75, 2 Oct 1949, Shinners 11809 (SMU). Tarrant 
Co. 3 mi W of Ft. Worth, 23 Sep 1936, McCart s.n. (SMU); Tandy Hills Municipal Park, ca. 2 air mi E of 
downtown Fort Worth, N side of Hwy 30, limestone slopes with shallow to deeper soil on flats and 
slope bottoms, 22 Jul 2001, Nesom FW57 with O’Kennon (BRIT, KSC, OKL, TEX, US): dry, s y ground, 
2 Sep 1929, Ruth 78(SMU), SE of Fort Worth near Village Creek, 4 Oct 1946, Whitehouse 17295 (SMU). 


Taxa of Series Punctatae 
The taxa involved in the present study are members of sect. Liatris series 
Punctatae Gaiser (1946), which is characterized by relatively few-flowered ca- 
pitula, phyllaries with non-petaloid tips, and a plumose pappus. Of the five 
species recognized by Gaiser in series Punctatae, L. punctata Hook. and L. 
densispicata (Bush) Gaiser are distinct in their elongate rootstock-like corms 
(vs. globose to subglobose corms). Liatris densispicata was regarded by Gaiser 
as endemic to Minnesota; it has more recently been treated as a synonym of L. 
punctata (Ownbey & Morley 1991). Liatris bracteata Gaiser is a Texas endemic 
distinct in its large, many-flowered (10-14 florets) capitula on relatively long 
peduncles. Liatris mucronata, L. aestivalis, and the second new species (de- 
scribed below) are similar in their relatively few-flowered capitula and globose 
to subglobose corms. 

Infraspecific variants have been described in both Liatris mucronata and 
L. punctata, but in most of Texas and Oklahoma L. mucronata is closely similar 
to L. punctata, except for the difference in corm morphology. They intergrade 
in parts of their range, as noted by Gaiser (1946) and Menhusen (1963). Fig. 5 
(present manuscript) maps L. mucronata as well as L. punctata and indicates 
that they have a nearly contiguous range in Texas. Plants with intermediate 
morphology are relatively common along the zone where their ranges meet. In 
Texas, the morphological difference between L. aestivalis and L. mucronata is 
easier to ascertain than than of L. punctata and L. mucronata, since the critical 
features of distinction are not underground parts. Within series Punctatae, we 
focus on comparisons of L. aestivalis with L. mucronata because of their simi- 
larity in corm morphology and their recent conceptual confusion. 


Distinction of Liatris aestivalis 
The original circumscription of Liatris(Lacinaria) angustifolia by Bush is not 
adequate to distinguish it from L. mucronata and the specimens of L. angustifolia 
cited by him include collections of L. mucronata. In Gaiser’s morphological de- 
scription of L. angustifolia, she fairly clearly was observing the core of the same 
taxon recognized here as L. aestivalis, but she also cited various out-of-range 
Texas collections (here regarded as L. mucronata), apparently including plants 
with atypically long but otherwise mucronata-type phyllaries. 

Also among Gaiser’s citations for Liatris angustifolia are collections from 


NESOM AND O’KENNON, NEW SPECIES OF LIATRIS FROM NORTH CENTRAL TEXAS 775 


southern Missouri, eastern Kansas, and Nebraska. Our examination of collec- 
tions from these states, as well as northern Arkansas, confirms their similarity 
to L. aestivalis, but this ‘northern segment is distinct in a number of features 
(mostly ‘gigas’ compared to L. aestivalis) and geographically disjunct. While 
evidence for the distinction of L. mucronata and L. aestivalis in Texas and Okla- 
homa is clear, variation patterns in the overall range of both L. mucronata and 
L. punctata need to be restudied. 

Morphological differences between Liatris aestivalisand L. mucronata are 
conspicuous in features of the capitula. The long lower cauline leaves, abruptly 
reduced upward, also provide a useful feature of differentiation. Plants of L. 
aestivalis also are generally shorter, thinner-stemmed, an e smaller corms 
producing a smaller number of stems, but these differences are hard to quan- 
tify on the basis of herbarium specimens, as collectors avoid the larger diffi- 
cult-to-press plants. 

The flowering period of L. destivalis is earlier than that of L. mucronata 
(Table 1). At most localities, the two apparently are complete separate in flow- 
ering period—if overlap occurs, it is only for a short time around the end of 
August. In fact, the flowering period of a given population appears to bea fairly 
reliable indicator of its identity (L. aestivalis vs. L. mucronata). As identified 
here, L. aestivalis also has a relatively discrete geographic distribution (Fig. 6), 
although it is imbedded within that of L. mucronata. Details of morphological 
and phenological differences between the two taxa are given in the following 
couplet. 


aa 


. Lower cauline leaves usually 2-3.5 times longer than the midcauline and 1.5-2 

times wider; phyllaries 6-11 per head, in 2-3 series weakly graduate in length, the 

outermost ca. 1/2-2/3 the length of the inner, inner 11-12 mm long, oblong-lan- 

ceolate to lanceolate-triangular, acute to acute-attenuate at the apex, not mucr- 

onate or cuspidate but sometimes with a long, narrow tip, lateral veins usually evi- 

dent above the middle and continuing to bas the apex; florets 3—4(-5) per head; 

flowering mostly mid-July through mid-August Liatris aestivalis 
. Lower cauline leaves similar to the midcauline | in length and width; phyllaries 11- 

15(-18) per head, in (4-)5-6 series strongly graduate in length, the outermost 1/3- 

1/5 the length of the inner, inner 7-9 mm long (-12 mm in putative esata 

mostly oblong-obovate and obtuse to abruptly truncate-rounded at the apex, 

tipped by a thick, often spinulose mucro or cusp, lateral veins ee not evident 

above the middle or at least on the distal third; florets (3—)4—5(—6) per head; flower- 

ing mostly mid-September through mid-October Liatris mucronata 


—~ 


Habitat and plant associates 

Populations of Liatris aestivalis in Tarrant County apparently are restricted to 
sites with shallow soil over outcrops of Glen Rose Limestone, on slopes or on 
flatter areas at the slope bases. The exposed limestone at other L. aestivalis sites 
in Texas and Oklahoma is closely similar in origin and characteristics to the 
Glen Rose formation. The species apparently is true to such sites, except for the 


776 BRIT.ORG/SIDA 19(4) 


Parker County plants (O’Kennon 15007), which grow in deep sand. The latter, 
however, are otherwise inseparable from typical L. aestivalis, which also grew 
in a typical habitat about 50 feet away. Corms of the new species tend to be 
shallow (1-5 cm below the surface, or even partly exposed) in relatively loose 
soil, but some occur in rockier substrate. 

Associated plant species at these limestone outcrops in Tarrant County are 
remarkably predictable: Artemisia ludoviciana var. mexicana, Ceanothus 
herbaceus, Chamaesyce fendleri, Dalea hallii, Dyssodiopsis tagetoides, 
Echinacea angustifolia, Eryngium leavenworthii, Grindelia lanceolata var. 
texana, Hedyotis nigricans, Heliotropium tenellum, Marshallia caespitosa, Opun- 
tia humifusa, Paronychia virginica, Penstemon cobaea, Phyllanthus polygonoides, 
Polygala alba, Salvia texana, Schizachyrium scoparium, Scutellaria wrightii, 
Silphium albiflorum, Sorghastrum nutans, Stenosiphon linifolius, Styllingia 
texana, Thelesperma filifolium, Vernonia lindheimeri, and Yucca pallida. In 
Tarrant County, Yucca pallida is a conspicuous and reliable predictor of the 
presence of L. aestivalis, although the Liatris is even more simply located by 
looking for exposed slopes with the white limestone. In Montague County, Yucca 
arkansana replaces Y. pallida asa conspicuous associate of L. aestivalis and other 
local variations in species composition are evident. 

Liatris mucronata also occurs at most sites where we have found L. 
aestivalis, providing evidence that the two taxa are distinct. While there is over- 
lap in microhabitat, L. mucronata occurs in deeper soil, compared to the rocky, 
shallow soil to which L. aestivalis is restricted. Presumably, the combination of 
phenological separation and distinction in microhabitat provides genetic iso- 
lation between the two taxa. The broader ecological range of Liatris mucronata 
corresponds with a geographic range wider than that of L. aestivalis (compare 
Figs. 5 and 6). 

Chromosome numbers for taxa of series Punctatae have been reported by 
Gaiser (1950) and Menhusen (1963). Counts for L. mucronata (two populations— 
Comal and Nueces cos., Tex.) were diploid (2n = 20). Counts for “L. angustifolia” 
from eight localities were diploid; a single population of this taxon, from Wichita 
Co., Tex., was reported by Gaiser to have plants at diploid and tetraploid levels. 
It is not clear that sampling by Gaiser included plants identified here as L. 
aestivalis, and Menhusen did not include Texas plants in her study. 


— 


Types examined 
Lacinaria angustifolia Bush, Amer. Midl. Naturalist 12:315. 1931. Liatris angustifolia 
(Bush) Gaiser, Rhodora 48:368. 1946. Type: TEXAS. “Cultivated specimen in green house from 
seed from Texas plants, TYPE, type locality unknown, sheet No. 211347” (as cited by Bush 
1931, p. 315); the specimens label has “Cult. Green House fr. Texas,” (HOLOTYPE: MINI). Date ol 
° } = j 


1 not indicated. 


Gaiser (1946, p. 369) gave the type locality of Lacinaria angustifolia with more 
precision—“Tarrant Co. dry soil, Fort Worth, Sept. 5, 1912, A. Ruth 78 (M, 211347),” 


NESOM AND O’KENNON, NEW SPECIES OF LIATRIS FROM NORTH CENTRAL TEXAS 777 


but in this she apparently took label data from some other specimen giving the 
collection number as “Ruth 78,” perhaps even confusing it with the type of 
Lacinaria ruthii (see below). Ruth, however, used the number “78” on labels of 
Liatris of various collections from different localities and different dates, al- 
though all of them apparently were made from Tarrant County or nearby lo- 
calities. Bush received specimens from Albert Ruth for use in his study, but if 
Ruth collected the seeds from which the type of L. angustifolia was grown, it is 
curious that Bush was not able to provide more precise collection data. 

This plant is similar to those cited above as perhaps indicating gene flow 
between Liatris aestivalis and L. mucronata. As in L. mucronata, it has leaves 
nearly equal in length and width from base to midstem, and the 15 phyllaries 
are strongly graduate in length, forming a distinctly cylindric involucre, and 
with densely ciliate margins. The phyllaries, however, are more elongate than 
typical for L. mucronata and the apices vary from obtuse-mucronate to ovate- 
lanceolate. We have annotated this collection as “Liatris mucronata perhaps 
with genetic influence of L. aestivalis.” 

a Ic eae Bush, Amer. Midl. Naturalist 12:314. 1931. Type: TEXAS. 
r Co. on poor, rocky, sterile ground, 10 Jun 1923, A. Ruth 24 (HOLOTYPE: ISC). 


— 


This was treated by Gaiser (1946) asa synonym of Liatris mucronata, and that 

identity is confirmed here. The collection is a single plant with six flowering 

stems from a globose corm. The capitulescence is narrow (hence the epithet) 

and the capitula have mostly three florets each, but the capitula have ca. 15 

phyllaries and the shape and apex are similar to those of L. mucronata. Phyllar- 

ies are conspicuously tomentose on the lamina and have finely long-ciliate 
margins. The collection date isanomalous and probably is incorrect—the spikes 
are in full flower or slightly past (no unopened florets remaining)—a flowering 

date anytime in June is considerably earlier than any known collection of L. 

mucronata and most collections of L. aestivalis (see Table 1). 

Lacinaria ruthii Bush, Amer. Midl. Naturalist 12:316.1931 (non Lacinaria ruthii 
Alexander 1933 = Liatris squarrulosa Michx.). Type: TEXAS. TARRANT Co: Poly- 
technic, 5 Sep 1912, A. Ruth 78 (HOLOTYPE: MINI). 

This was regarded by Gaiser (1946, p. 370) to be among intermediates between L. 

angustifolia and L. mucronata, but the type specimen belongs with nearly typical 

L. mucronata. Flowering in early September is early for L. mucronata, late for L. 

aestivalis, but all morphological features of this plant are L. mucronata: phyllaries 

18 in number, strongly graduate in length, with an abruptly rounded, cuspidate 

apex, lightly tomentose on the surface, with ciliate margins; florets 4 per head. 


A STIPITATE-GLANDULAR SPECIES OF LIATRIS 


In examination of collections identified as Liatris mucronata, a small set of 
plants with stipitate-glandular vestiture was separated. These collections were 


BRIT.ORG/SIDA 19(4) 

1. Flowering phenology of plants of Liatris aestivalis and L. mucronata in Texas and southern 
Kasaaes Lower tabular data show number of collections scored at mid-flower for each date (L. 
aestivalis: 21 total collections; L. mucronata: 136 total collections; for each collection scored, an esti- 
mate was made of when the plant/plants was/were in mid-flower). Mae tabular data show per- 
centage of the total number of collections scored at mid-flower.Graph show 


iec 


spec mpar d by 
the percentage of the total number of collections scored at mid-flower. At most sites where the 
two species co-occur, overlap in flowering apparently occurs rarely or not at all. 


L. aestivalis 
L. mucronata 


Oi-Jun 15-Jun 01-Jul 
0 5 14 


15-Jul 01-Aug 15-Aug 01-Sep 15-Sep 01-Oct 15-Oct 01-Nov 
24 28 24 5 0 
0 


9 44 36 11 0 
| —L. aestivalis ~~ L. mucronata 
wm 50 
5 : 
% 40 = — 
a / ‘ 
Oo 30 - a 
oO Paes f \ 
= ‘. 
S 20 : SS 
& wee. \ 
8 10 + Sa = “~ 
(0) me 
Oo. 0 Tv Ss tT ~ 7 — _ : ae aes ~ a... T ~ 
S S$ > > we) RS) Q g ce om 3 
» » Ss ys ¥ o g S 
ee oe °° NCI x oo a Oo ia ad a 
Flowering date (mid-flower) 
L. aestivalis 


O1-Jun 15Jun O1-Jul 15-Jul O1-Aug 15-Aug O1-Sep 15-Sep 01-Oct 15-Oct O1-Nov 
1 3 5 6 5 1 0 
L. mucronata 


0 12 60 49 15 


0 
made in the 1930s and early 1940s from within Dallas County, before dense 
residential, commercial, and transportation development covered most of the 
natural habitat. A collection of this entity was made in 1936 by L.O. Gaiser, who 
also obtained chromosome counts from five plants of the population, and the 
SMU sheets were examined by Cory and Shinners, but none of these research- 
ers apparently found taxonomic significance in the distinctive features of these 
plants. In the present perspective, however, the vestiture of these plants is unique 
in the genus, and other features set it apart from L. mucronata. The discovery of 
a large, multi-segmented population in a nature preserve, growing sympatri 
cally with L. mucronata but obviously distinct from it in morphology and phe- 
nology, allows us unhesitatingly to describe it at specific rank 


Liatris glandulosa Nesom & O’Kennon, sp. nov. (Figs. 3, 4, 6). Type: US.A. TEXAS 
Dallas Co.: City of Garland, Dallas metro area, 


ca. 16 air mi NW of downtown Dallas, Spring 
Creek Forest Preserve, between Interstate Hwy 190 and Arapaho bea vicinity of Holford 
Read ae habitats on open relatively flat, li 


Chalk formation) 
ry fill, 21 Jul 2001, Guy L. Nesom FW62 with Rol be O’Kennon and Thomas 
M. Fre sitions OTYPE: BRIT, ISOTYPES: GH, KSC, MO, NY, OKL, TEX, US 


S—to be distributed). 


Differt a L. mucronata DC. vestimento stipitato- ae foliis latioribus laxioribus, phyllariis 
paucioribus leniter 3-4-seriatis, et florescentia praecic 


NESOM AND O’KENNON, NEW SPECIES OF LIATRIS FROM NORTH CENTRAL TEXAS 779 


Plants 3-6 cm tall with 1-2(-4) erect stems arising from a subglobose to de- 
pressed-globose corm 2-4 cm in diameter; stems, leaves, and phyllaries densely 
punctate with imbedded, clear-resinous glands, stems and leaves also densely 
stipitate-glandular with trichomes 0.1-0.2(-0.3) mm high, the whole plant 
sticky to the touch, very sparsely arachnoid, especially along the margins. Leaves 
crowded, spreading-ascending or the lower slightly deflexed, rigid to lax, the 
lower 6-11 cm long, 1.5-4(-4.5) mm wide, usually 1/2-1/3 as long at midstem 
and ca. 1/2 as wide, usually reduced to short, spreading bracts in the lower part 
of the capitulescence. Capitulescence 8-18(-28) cm long, mostly 1.5-2.5cm wide, 
of densely arranged capitula. Capitula turbinate-cylindric, 8-12 mm long, 3-5 
mm wide; phyllaries 7-10(-12) per head, in 3-4 series weakly graduate in length, 
the outermost 1/2-2/3 the length of the inner, inner 9-11(-12) mm long, ob- 
long-lanceolate to lanceolate-triangular, broadly to narrowly acute or acumi- 
nate at the apex, sometimes with an indurate mucro, green, rarely becoming 
purplish, very densely glandular with a mixture of sessile-sunken glands and 
slightly elevated ones, margins narrowly hyaline below the apex. Florets 3-4(- 
5) per head; corollas pink-purple, punctate-glandular, 7-8 mm long, tube 4-5 
mm long, glabrous to glabrate within, lobes linear-lanceolate, ca. 3 mm long; 
filaments puberulent-pilose. Cypselae 5-6 mm long, oblong-oblanceolate in 
outline, brown to nearly black, compressed to 3-angled, with 8-10 nerves, 
densely hairy on nerves and interstices with duplex hairs 0.2-0.6 mm long; 
pappus of plumose bristles 6-7.5 mm long, ca. equal the corolla length. Chro- 
mosome number, 2n = 20 (Gaiser 1950). 

Etymology—The epithet alludes to the stipitate-glandular vestiture of the 
species, a distinctive feature compared to L. mucronata and all other species of 
the genus. 


Additional collections examined: TEXAS. Bosque Co.: ca. 4 mi S of Clifton along Hwy 6, | Sep 1946, 
Pudig and York 46334 (LL, MO); gravel road E of Tx Hwy 6 and ca. 2.5 NW of Clifton, N of Meridian 
Creek K, dry, Srey limestone hill, 800 ft, Somancne as limestone, community dominated by 
J Aug 1989, O Bridges 11401 (TEX). Dallas Co.: 11 mi W of 
Dallas on US Has 80, challey limestone (Austin Chalk) at fault on road to Ft. Worth, voucher for 
chromosome count of n = 10, 27 Aug 1936, atl 122 (OKL),; N of eek Park, prairie, 2 Aug 
a Longnecker 6 (LL, SMU-2 sheets); [city of D asl, off Hillcrest Road [in current vicinity of South- 
ern Methodist University], on hineone outcrop, 21 Aug 1942, Lundell 115610 L, SMU); Duncanville, 
Dallas metro area SW of downtown Dallas, junction of Clark Road and Camp Wisdom Road, ca. 2 
acre lot on NE corner, currently advertised for sale, shallow, gravelly limestone soil, common species 
aie nutans, sank undulatum, Centaurea solstitialis, Mentzelia oligosperma, Diplotaxis 
muralis, Solid iss otis nigricans, ca. 10 plants of Liatris glandulosa in mid flower, 25 
Jul 2001, Nesom FWI5 and OKennon (BAYLU, BRIT, COLO, GH, ISC, KSC, MIN, MO, NCU, NLU, NY, 
OKL, RM, TEX, TAES, UC, US); Coomb’s Branch 19 Aug[without year], Reverchons.n.(MO). McLennan 
Co.: W of Hillcrest Hospital, limest ilon creek bank, 9 Aug 1947, Smith 1007 (TEX). Travis Co.: 
“pocket” in rocks on Bee Creek, 6 Sep 1901 [past flr, frting heads shattering], Ferguson 451 (TEX). 


We found plants of Liatris glandulosa in two Dallas County sites where collec- 
tions had not been previously made. The Duncanville site is about 6 miles south 


780 BRIT.ORG/SIDA 19(4) 


NESOM AND O’KENNON, NEW SPECIES OF LIATRIS FROM NORTH CENTRAL TEXAS 781 


Fig. 4. Habit of Liatris glandulosa. 


782 BRIT.ORG/SIDA 19(4) 


© Liatris punctata 
@ Liatris mucronata 


@ L. mucronata <> L. punctata 


Fic. 5. Distributi f Liatri 1 L. punctata in Texas i 
taxa. Distinctions between varieties in L. peneN (var. es Gaiser ane var. mexicana Gaiser) and between 


varieties in L. mucronata 


r / F 


southeast of the 1936 collection site by Gaiser (her collection 122)—this isa flat 
(probably artificially so), open field of about two acres with exposed limestone 
in some places. Numerous invasive plant species are abundant, and a large “for- 
sale” sign signals the close approach of ‘development of this lot and extirpa- 
tion of the Liatris population. We collected most of these plants, taking corms 
for cultivation. 

The Spring Creek site is a nature preserve that now protects the only known 
population of Liatris glandulosa in Dallas County. Here, the species grows in 
apparently natural habitats, and it also occurs in abundance in a ‘reconstituted’ 
substrate of mixed crushed limestone, gravel, and asphalt that filled a quarry 
and gravel pit site now located ina central portion of the preserve. This portion 
of the quarry was active until around 1988 (when acquired by Dallas County), 


© LOE Ee es 
CT oe ee 
tae eae as nice AL 


| 

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oN 
a 
se 


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= 
ae 
ae 

\ 


784 BRIT.ORG/SIDA 19(4) 


so the revegetation (including Liatris recolonization) apparently has occurred 
within the last 12 years, lending optimism that L. glandulosa may be success- 
fully sustained at Spring Creek and propagated elsewhere. 

In the previously quarried area at Spring Creek (E side of Holford Road), 
with the plants of Liatris glandulosa, these species were common: Yucca 
drkansana, Asclepias asperula, Hedeoma reverchonii, Heliotropium tenellum, 
Hedyotis nigricans, Paronychia virginica, Dalea multiflora, Pedomelum 
linearifolium, Eryngium leavenworthii, Mentzelia oligosperma, Grindelia 
lanceolata var. lanceolata, Hymenopappus scabiosaeus, Iva angustifolia, and 
Thelesperma filifolium. Liatris mucronata (Nesom FW68) also is relatively com- 
mon on the quarry fill, as well as other sites on deeper soil, but it was not yet in 
flower on 25 July, when the L. glandulosa was collected. Natural habitat adja- 
cent to the old quarry site, encroached by Juniperus virginiana, Cercis 
canadensis, Rhamnus caroliniana, Rhus trilobata, harbors L. glandulosa and 
herbaceous species in addition to those above: Schizachyrium scoparium, 
Sporobolus compositus var. drummondii, Krameria lanceolata, Eriogonum 
longifolium, Echinacea angustifolia, and Amphiachyris sp. On a hilltop with 
exposed limestone (W side of Holford Road, about 1600 feet from the quarry 
site noted above), L. glandulosa grows with a similar set of species: Yucca 
arkansana, Hedyotis nigricans, Heliotropium tenellum, Hedeoma reverchonii, 
Paronychia virginica, Eriogonum longifolium, Phyllanthus polygonoides, 
Styllingia texana, Toxicodendron radicans, Krameria lanceolata, Vernonia 
lindheimeri, Thelesperma filifolium, Echinacea angustifolia,and Amphiachyris 
sp. Liatris mucronata also is relatively common in deeper soil in this area (to 
the periphery of the L. glandulosa plants) but was not yet in flower on 25 July. 

Spring Creek Forest Preserve is close to the northern boundary of Dallas 
County, suggesting that similar sites may still exist in adjacent Collin County. 
Habitats for Liatris glandulosa on the southwest side of Dallas County (Gaiser 
122; Nesom FW75) are associated with the White Rock Escarpment, which ex- 
tends southward. We will search for additional populations of the ‘sticky 
gayfeather’ in this area and in more southern counties and hope that Dallas- 
area naturalists will also do so. 


Distinction of Liatris glandulosa 

Plants of Liatris glandulosa have previously been identified as L. mucronata, 
but no other species of Liatris (across the whole genus) has a vestiture of stipi- 
tate-glandular hairs. Observation of these glandular plants growing sympatri- 
cally with L. mucronata and flowering far in advance of it indicates that two 
evolutionarily independent entities are present. There apparently is slight over- 
lap in flowering time with L. mucronata, and some plants have been observed 
that suggest that limited gene flow occurs. These two species are distinguished 
by the following contrasts. 


NESOM AND O’KENNON, NEW SPECIES OF LIATRIS FROM NORTH CENTRAL TEXAS 785 


— 


. Stems and leaves densely stipitate-glandular, sticky to the touch; leaves relatively 

lax, ascending to loosely spreading or the lower deflexed; phyllaries 7-10(-12) per 

head, in 3-4 weakly graduate series, the outermost 1/2-2/3 the length of the inner, 

inner mostly oblong- sanieeolaie to lanceolate-triangular, broadly to narrowly acute 

at the apex, sometimes with an tes rate mucro; florets 3-4(-5) per head; flowering 

mostly mid-July through early = mber Liatris glandulosa 
Stems, leaves,and phyllaries eerie to weakly punctate, otherwise essentially gla- 
brous or sparsely ciliate along the margins, not at all sticky; leaves stiffly soreading- 

ascending; phyllaries 11-15(-18) per head, in (4-)5-6 series strongly eae in 

length, the outermost 1/3-1/5 the length of the inner, inner mostly oblon 

vate and obtuse to abruptly truncate-rounded at the apex, tipped by a ee ae. 

spinulose mucro or cusp; florets (3—)4—5(-6) per head; flowering mostly mid- 

September through mid-October Liatris mucronata 


SS 


— 


We hypothesize that Liatris glandulosa and L. aestivalis have a sister relation- 
ship and that these two as a pair are most closely related to the northern seg- 
ment of L. mucronata. Compared to ‘southern’ L. mucronata, the new species 
are both early-flowering and grow ina similar habitat, both are relatively re- 
stricted in geographic distribution, and they produce fewer-flowered capitula 
with fewer, weakly graduate phyllaries. Their geographic ranges are nearly con- 
tiguous—the populations of L. glandulosa in southwestern Dallas County are 
disjunct from closest L. destivalis habitats in Tarrant County by an area of deep 
soil (Woodbine sand and Eagle Ford clay) without rock outcrops—or they may 
overlap southward. In addition to the difference in vestiture, plants of L. 
aestivalis differ from L. glandulosa in their narrower leaves and phyllaries that 
usually turn purple. 


Postscript.—The question arises—how did these species escape the notice of 
Lloyd Shinners, who lived, worked, and botanized in north central Texas? Ina 
commentary on Liatris mucronata, Shinners (1951) noted that “After observ- 
ing and collecting the plants for five years, I fail to recognize two species [L. 
angustifolia and L. mucronata] as distinguished by Dr. Gaiser ...” Shinners in- 
cluded L. angustifolia as a synonym of L. mucronata and observed that the lat- 
ter is “very common in north central Texas in the wild, ona variety of soils (but 
commonest on limestone and chalk outcrops and calcareous prairies).” In this 
case, with all respect to the acuity of our predecessor, reality remained elusive. 
At least in part, he may have been led in his observations by Gaiser’s diffuse 
morphological and geographical concept of L. angustifolia narrowed here to L. 
aestivalis) and its distinction from L. mucronata. Shinners collected L. 
mucronata from various habitats in 12 Texas counties, but there is only a single 
record of his collection of the earlier-flowering L. aestivalis—this from a Den- 
ton County plant far past flowering. Shinners also examined the SMU collec- 
tions of L. glandulosa and recorded (by annotation) involucre height, and V.L. 
Cory similarly recorded by annotation his observation that the plants were glan- 
ular, but neither botanist carried the investigation further. 


186 BRIT.ORG/SIDA 19(4) 


We also note that, given the broader ecological range of Liatris mucronata, 
it certainly was once much more abundant than L. aestivalis, at least in the 
Fort Worth area. The most common habitat for L. mucronata is open, flat prai- 
rie—this habitat is easily turned to commercial and residential development 
and it has been. The relatively numerous cited collections of L. aestivalis from 
within the city of Fort Worth (where it is now more “common” than L. 
mucronata) reflect the persistence of scattered little plots of steep, rocky slope 
that have not yet been built upon. A few of these sites have been preserved within 
the city park system. 

If the existence of Liatris glandulosa had been earlier recognized, it might 
or might not have prevented the apparent destruction of most of the species’ 
habitat in Dallas County. The value of preservation of natural sites is empha- 
sized by the present example, where the only known extant population of L. 
glandulosa exists in areas of a relatively small park (83 acres) preserved within the 
city of Garland as part of the Dallas County Park and Open Space Program. The 
park was created to protect the old-growth forest that occurs there—preserva- 
tion of the prairie patches harboring the rare Liatris was not a central concern. 


ACKNOWLEDGMENTS 


We thank the staffs at ISC, MIN, OKL, and OKLA for loans of pertinent speci- 
mens, the staffs of BAYLU, MO, and TEX-LL for help during visits there, Craig 
Freeman and Caleb Morse (KANU) for sending a copy of the Menhusen disser- 
tation, Bill Burk at the Couch Botanical Library (UNC-Chapel Hill) for help with 
other literature, and Mark Mayfield for comments on the manuscript. Hugh 
Garnett, Shirley Lusk, and Betty Miller showed us populations of Liatris 
aestivalis and L. mucronata in north central Texas that they have been observ- 
ing over a number of years and shared original insights about these plants, in- 
cluding their own recognition that the name L. mucronata has represented more 
than a single species. We are grateful to Tom Frey, Landscape Architect with 
the Garland Parks and Recreation Department, for guiding us within the Spring 
Creek Forest Preserve for observation of L. glandulosa. Illustrations of the new 
species are by Linny Heagy. 


REFERENCES 


Bush, B.F. 1931.Some Lacinaria species. Amer. Midl. Naturalist 12:312-318. 

Corrett, D.S. and M.C. JoHNston. 1970, Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner, IX. 

Diccs, G.M., Jr., B.L. Liescoms, and R.J. O’Kennon. 1999. Shinners and Mahler's illustrated flora 
of north central Texas. Sida, Bot. Misc. No. 16. 

Gaiser, L.O. 1946. The genus Liatris. Rhodora 48:165-183, 216-263, 273-326, 331-382, 
393-412. 


NESOM AND O’KENNON, NEW SPECIES OF LIATRIS FROM NORTH CENTRAL TEXAS 


Gaiser, L.O. 1950. Chromosome studies in Liatris. lll. Punctatae. Amer. J. Bot. 37:763-777. 

Hatcu, S.L.K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas 
Agric. Exp. Stat. MP-1655, College Station. 

JOHNSON, M.C. 1990.The vascular plants of Texas (ed. 2). Published by the author, Austin, FX. 

Jones, S.D., J.K.Wiprr, and P.M.Montcomery. 1997.Vascular plants of Texas. Univ. of Texas Press, 


Austin. 

Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the 
vascular flora of the United States, Canada, and Greenland. First Edition. In: Kartesz, J.T., 
and C.A.Meacham. Synthesis of the North American Flora, Version 1.0. North Carolina 
Botanical Garden, Chapel Hill, NC. 

MenuuseNn, B.R. 1963. Variation in the Punctatae series of the genus Liatris. Ph.D. diss., Univ. 
of Kansas, Lawrence. 

Owneey, G.B. ano T. Mortey. 1991. Vascular plants of Minnesota. Univ. of Minnesota Press, 


Minneapolis. 

SHINNERS, L.H. 1951.Notes on Texas Compositae—VIl. Field & Lab. 19:74-82. 

Taytor, R.J.and C.E.S. Tavior. 1994. An annotated list of the ferns, fern allies, gymnosperms 
and flowering plants of Oklahoma (ed. 3). Biology Dept. Herbarium, Southeastern Okla- 
homa State University, Durant. 

Warerract, U.T. et al. 1969 and 1994, issued 1994. Identification of Oklahoma plants. |. Key 
to the vascular plant families of Oklahoma, by RJ.Tyrl (coordinator) et al. (1994). Il. Keys 
to the flora of Oklahoma, ed. 4, by U.T. Waterfall (1969). Flora Oklahoma, Inc., Noble, 
Oklahoma. 


788 BRIT.ORG/SIDA 19(4) 


Book REVIEW 


THOMAS M. BONNICKSEN. 2000. America’s Ancient Forests: From the Ice Age to the 
Age of Discovery. (ISBN 0-471-13622-0, hbk.). John Wiley & Sons, Inc., One 
Wiley, Drive, Somerset, NJ 08875, U.S.A. Orders: http://www.wiley.com/ 
Corporate/ Website /Objects/Products/0,9049,38301,00.htm1). $90.00, hbk., 
608 pp, b/w figures, 6" x 9" 


What a wonderful synthesis of information from fields as widely varied as botany, ecology geology, 


archaeology, anthropology, and history! Thomas Bonnicksen has produced a work that will capture 


the imagination of anyone interested in the grandeur and beauty of the forests of North America. 
The first half of the book describes in wonderful detail how the forests of North America came 


to be what they were at the time of European settlement. The first two chapters set the stage with a 
cogent discussion of what drives ice ages, and a magnificent description of what the ice age forests 
must have looked oe The ae chapter takes the reader on a journey through time as Bonnicksen 


describes the mi iti { forests and the expansion of prairies 


oOo oO tr I 
} t} i : 1. ‘ } ] j 
J 


xt four chapters provide an 
account of the effects both bene ficial and detrimental that the ae ay Americans had on the 
en: development of the North American landscape. Major themes in these chapters include: 

effects on the fauna and mass extinctions caused by changing climates and hunting practices; 2) 
he a sia: and managing of forests for food and shelter; 3) the d tication of native species, 


1oW various native cultures utilized the natural resources found in forests; 4) an 


dic 


on how the Native Americans’ use of fire both intentionally and accidentally 
altered the eee 


The sone part of the book provides wonicenay) ist descriptions of the North American 


¢ 


uropea em. Chapter 9 gives an accounting of the forests first 
encountered by Spanish ce in the he and western rinited States. Chapter 10 provides 
well-documented historical accounts of the ecology and uses of the forests by colonists in the east. 
Chapter 11 looks at forests [rom the perspective of missionaries, sisi were often Ene first t to see the 
extensive forests of Canada and the North Central United Sta use of 


the written descriptions of these early pioneers who often ene beautifully detailed records of what 


they saw, and were often well-educated in poreny and natural history. The final chapter provides a 


view of ee not often considered in many other texts, that of the trappers and fur traders who were 
often the first Europeans to lay eyes on the vast forests of the rugged western part of the continent. 

| would recommend this book to anyone interested in the natural history of North America. It 
is packed full of wonderfully interwoven bits of information from a wide variety of disciplines, and 
Bonnickesen tells this complex story in a style that is immensely readable, and thoroughly enjoy- 
able.—Debra K. Trock, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102- 
4060, U.S.A, dtrock@brit.or 


= 


a 


SIDA 19(4): 788, 2001 


LAENNECIA TURNERORUM (ASTERACEAE: ASTEREAE), 
A NEW SPECIES FROM TRANS-PECOS TEXAS 


G.L.Nesom 
Botanical Research Institute of Texas 
509 Pecan Street 
Fort Worth, TX 76102-4060, U.S.A. 


ABSTRACT 
Laenneciaturneror p. nov., from Brewster County, es is similar to ene sae of Laennecia 
sect. Sophiifolium (L. coulteri, L. sophiifolia,and L. may hirsute-pilose 


vestiture, acuminate phyllary apices, elaminate ray florets, and small, strigose cypselae with a 
uniseriate, caducous pappus. It differs from those three species in its stems, leaves, and phyllaries 
sparsely (vs. densely) glandular, basal leaves persistent at flowering (vs. senescent and usually ab- 
sent by flowering), reduced vestiture of the phyllaries, and non-accrescent pappus (vs. accrescent). 
Plants of L. turnerorum are spring ephemerals and grow with plants of L. coulteri. 


RESUMEN 


a : a ] 
a 
Ls | 


Laenneciaturnerorum sp. nov, del condado de ee ster, Texas 


dennecia 
sect Sophiifo lium (L. coulteri, L aa l hi 


pilose apices de| ] ] eee 


y cipselas pequenas, estrigosas con 
] Pit dp] : 1] 
L 


anh hojas, y filarios esparsamente 
| 1 ] b 1 . 1 7 ‘ ] f] 


(vs id senescentes y usualmente 


ausentes en la loracien). indumento de los filarios tue y vilano no acrescente (vs. acrescente). 


Las plantas de L. turnerot I y crecen con plantas de L. coulteri. 


Recent collections in trans-Pecos Texas by Dr. B.L. Turner have brought to light 
a previously undescribed species of Laennecia Cass. Plants of the new species, 
which apparently are spring ephemerals, are currently known from about six 
plants of a single collection. Their small size, disciform capitula, and short du- 
ration probably account for their escape from detection by earlier botanists. 
Turner returned to the type locality about four weeks after the type collection 
was made and found that plants of the new species had died and dried essen- 
tially beyond recognition, while those of Laennecia coulteri (Gray) Nesom at 
the same site remained alive and in reproductive condition (B.L. Turner pers. 
comm.). 


Laennecia turnerorum Nesom, sp. nov. (Fig. 1). Tver: U.S.A. TEXAS. BREWSTER CO. Turner's 
Valentine Section, vicinity of old ‘residence, ca 58 mi S of Alpine along Hwy 118, 29 34'N, 
103° 34’ W, 3330 ft elev, intermixed with Laennecia coulteri, 21 Mar 2001, B.L. Turner 21-73 
(HOLOTYPE: TEX; ISOTYPE: BRIT 


] F | a een ea : : 384 Ie : | ] “1 


1 }] at ee | : 4 f] ] = sd] ii ] 


auriculatis 
= t c 


vel 
is, et cypselis 
? Pa 
llearii : 


£ L 


a dene) glandulosa foliis petiolatis bipinnatis, et capitulis majoribus. 


SIDA 19(4): 789 — 793. 2001 


Wi 
re 
te " 


SAlZ 


BRIT.ORG/SIDA 19(4 


Fic. 1. Habit of Laennecia turnerorum (from isotype). 


) 


NESOM, A NEW SPECIES OF LAENNECIA FROM TRANS-PECOS TEXAS 791 


Plants annual, herbaceous, from a slender taproot, producing 1-3 erect to as- 
cending stems from the base, 16-20 cm tall; stems and leaves sparsely and mi- 
nutely sessile-glandular and stipitate-glandular, also sparsely hispid-pilose 
with thick-based trichomes 0.2-1 mm long, not at all woolly. Leaves basal and 
cauline, basal and lowermost cauline persistent but sometimes senescent at 
flowering, 2-5 cm long, blade portion oblanceolate to obovate, 5-12 mm wide, 
2-pinnatifid with ovate lobes or segments, lower cauline leaves with narrow, 
petioliform, non-clasping base, upper cauline gradually reduced in size to 1-2 
cm long, becoming epetiolate and clasping. Capitula ca. 5-15 in a corymboid or 
subcorymboid capitulescence; involucres broadly campanulate, 3-4 mm high, 
5-7 mm wide; phyllaries in 3-4 series, sometimes fused at the base into a disc 
3-4 mm in diameter, the outermost 3-3.7 mm long, 0.8-1 mm wide, 3/4 to 
equally as long as the inner, completely herbaceous, without an evident midrib 
or midvein, coarsely hispid along the midportion and sparsely glandular, inner 
3.5-4 mm long, narrower, with prominent hyaline flanges on the distal half, all 
witha hyaline, purplish apex. Pistillate florets ca. 200, in 2-4 series, corolla tube 
1mm long, whitish, lamina absent. Bisexual florets 15-18, corollas tubular, 2.2 
mm long, yellow, with purple lobes 0.2-0.3 mm long. Cypselae elliptic-oblong, 
0.8-1 mm long, tan, sparsely short-strigose, sparsely sessile-glandular at the 
apex; pappus uniseriate, of 16-18 fragile, white, caducous, non-accrescent 
barbellate bristles ca. 2 mm long, slightly longer than the style branches of pis- 
tillate florets, slightly shorter than the disc corollas. 

Etymology and distribution.—The epithet commemorates the two Turners: 
Billie, who collected the plants in the course of preparing a florula of the “Val- 
entine Section” (640 acres), and his wife Gayle, who surprised Billie with a 
Valentine’s Day gift of the property. The area is in a large basin essentially sur- 
rounded by desert mountains. The plants of Laennecia turnerorum were col- 
lected in silty limestone-derived soil in a low area with desert grasses; abun- 
dant Larrea, scattered Yucca, and other shrubs occur slightly upslope. Although 
the immediate site has been slightly disturbed, plant species in the area appar- 
ently are native. 

Within the genus, Laennecia turnerorum isa member of sect. Sophiifolium 
Nesom (Nesom 1990a), the plants characterized by their annual duration, hir- 
sute-pilose vestiture, elaminate corollas in pistillate florets, and relatively small, 
strigose cypselae with very small, deciduous glands and a uniseriate pappus of 
basally caducous bristles. The other North American species of sect. 
Sophiifolium are L. coulteri (Gray) Nesom (southwestern USA through north- 
ern Mexico), L. sophiifolia (Kunth) Nesom (southwestern USA through Mexico 
and Guatemala, then apparently disjunct to northern South America), and L. 
mapimiana Nesom (northeastern Durango, southeastern Chihuahua; Nesom 
1990b). Among these, L. turnerorum is most similar to L. mapimiana in its short 
duration (spring ephemeral), relatively small stature, sparsely glandular 


792 BRIT.ORG/SIDA 19(4) 


vestititure, basal leaves persistent until flowering, leaves non-clasping below, 
subclasping to clasping above, corymboid capitulescence, and relatively large 
capitula. Laennecia turnerorum can be identified among its most closely similar 
congeners by contrasts in the key below. 


KEY TO NORTH AMERICAN SPECIES OF LAENNECIA SECT. SOPHIIFOLIUM 


. Leaves clasping to subclasping, oblong in 1 outline with a broad lamina, sharply 
toothed, the teeth cut 1/4-1/3 to the mid Laennecia coulteri 
. Leaves clasping or not, oblanceolate to son ener er or oblong in outline, 
pinnatifid to 2-pinnatifid, the segments cut 2/3-3/4 to the midrib. 
2. Plants (15—) 30-70 cm tall; leaves not clasping; cee ida paniculiform, co- 
lumnar to pyramidal; involucres 1.5-2.5 (-3.5) mm wide Laennecia sophiifolia 
2. Plants 9-20 cm tall;at least the upper leaves one capitulescence corymboid 
to subcorymboid; involucres 5—7 mm wide 
3. Plants 7-10 cm tall, stems strongly decumbent; leaves 1-2 cm long, oblong 
to obovate-oblanceolate in outline, epetiolate or with a short, weakly differ- 
entiated petioliform portion, blades 3-6 mm wide, 1(—2)-pinnatifid; pappus 
of 9-12 bristles aennecia mapimiana 
. Plants 16-20 cm tall, stems erect to basally ascending; leaves 2-5 cm long, 
spatulate in outline, petioliform portion strongly differentiated, blades 5-12 
mm wide, 2-pinnatifid; pappus of 16-18 bristles Laennecia turnerorum 


Ww 


Direct comparisons of Laennecia turnerorum to L. coulteri and L. sophiifolia 
are as follows. 

* similar to L. coulteri in its subclasping upper leaves; different in its short 
duration, sparsely glandular vestiture, smaller stature (16-20 cm tall vs. [20- 
}30-110 cm), smaller (1.5-2 cm long vs. 1.5-10 cm), petiolate, bipinnatifid cauline 
leaves, non-clasping lower leaves, basal leaves present (vs. absent) at flowering, 
corymboid capitulescence (vs. columnar-paniculate to corymboid), and larger 
capitula (5-7 mm wide vs. 3-5 mm). 

* similar to L. sophiifolia in its pinnatifid leaves and non-clasping lower 
leaves; different in its short duration, sparsely glandular vestiture, smaller stat- 
ure (16-20 cm tall vs. [15-] 30-70 cm), slightly smaller (1.5-2 cm long vs. 1-4 
cm) cauline leaves with shorter, broader segments, the upper leaves subclasping 
to clasping, basal leaves present (vs. absent) at flowering, capitulescence 
corymboid (vs. columnar- to pyramidal-paniculate), larger capitula (5-7 mm 
wide vs. 1.5-2.5[-3.5] mm), and greater number of pappus bristles (16-18 vs. 9- 
12[-15)). 

The geographic range of Laennecia turnerorum and its relationship to L. 
mapimiana should be studied by search during the early spring. The latter also 
appears to be narrowly distributed; it is separated at its closest point by about 
300 kilometers from the L. turnerorum locality. Both taxa are highly restricted 
in range compared to L. coulteri and L. sophiifolia. 

The new species brings the number of species in Laennecia to 18 (Nesom 
1990a, 1990b, 1992). Morphological and molecular data (Zardini 1981; Nesom 


NESOM, A NEW SPECIES OF LAENNECIA FROM TRANS-PECOS TEXAS 793 


2000; Noyes & Rieseberg 1999) indicate that the genus is not closely related to 
Conyza Less., where most of the species have been placed previously. Laennecia 
instead apparently is related to South American groups, possibly most closely 
to subtribe Podocominae (Nesom 1994, 2000) 


ACKNOWLEDGMENTS 


lam grateful to the staff of TEX-LL for their help during a recent visit and to 
Ted Barkley and John Strother for their comments on the manuscript. Linny 
Heagy provided the illustration. 


REFERENCES 


Nesom, G.L. 1990a. Taxonomy of the genus Laennecia (Asteraceae: Astereae). Phytologia 
68:205-228. 

Nesom, G.L. 1990b.Laenneci I (Asteraceae: Astereae),a new species from north- 
western Mexico. Phytologia 69: 348-350. 

Nesom, G.L. 1992. Laennecia spellenbergii (Asteraceae: Astereae), a new species from 
Durango, Mexico. Phytologia 73:267-269 

Nesom, G.L. 1994. Subtribal classification of the Astereae (Asteraceae). Phytologia 76: 


gos 


Nesom, G.L.2000. Generic conspectus of the tribe Astereae (Asteraceae) in North America, 
Central America, the Antilles, and Hawaii. Sida, Bot. Misc. 20:i-vili, 1-100. 

Noyes, R.D. and L.H. Rieseserc. 1999. ITS sequence data support a single origin for North 
American Astereae (Asteraceae) and reflect deep geographic division in Aster s.|.Amer. 
J. Bot. 86:398-41 2. 

Zarvini, E.M. 1981. Contribucidnes para una monografia del género Conyza Less. Il. 
Rehabilitacidn del género Laennecia Cass. Darwiniana 23:159-169 


794 BRIT.ORG/SIDA 19(4) 


Book REVIEW 


AMANDA MCQUADE CRAWFORD. 1997. Herbal Remedies for Women: Discover Nature’s 
Wonderful Secrets just for Women. (ISBN 0-7615-0980-1, pbk.). Prima Pub- 
lishing, P.O. Box 1260BK, Rocklin, CA 95677, www.primahealth.com, 
800-632-8676, 916-632-4400). $18.00, pbk., 291 pp., illustrations, 
7 3/8" x 91/4" 

There is currently much interest in herbal remedies and the proliferation of publications dealing 

ul alternative to “tradi- 

tional” medicine as well as the cae public’s increasing fascination with this topic. Most herbals 


with herbal medicine reflects the acceptance of herbal remedies as a use 


deal with a wide range of diseases and conditions, and may be overwhelming to the beginning stu- 
dent of herbal therapy. This on is designed specifically to address the oo ular health needs o 
women, and is full of information on nearly every problem from anemia to varicose veins. 

Amanda MeQuade Cian has spent a lifetime staying and nee about herbal thera- 


ray 


pies 


pS 


o women, and to the medical person- 

nel in women’s relies. about alanine eens for conditions and disorders that often have a 

profound impact on the quality _ as to women. 
= 


The first portion of thi wate ic ink ion al t plants, plant collecting, 

specific actions of useful nes and terminolo BEOCAISS ith tl f herbal remedies. 
by t 

h der of the book dez i mae menses, reproduction, meno- 


pause, sex sain transmitted diseases and’ abnormal cell growth.” Aeouak provides easily under- 


standable definitions of each di fi followed by a discussion of symptoms, causes and 
conventional medical treatments. es then provides good descriptions of alternative herbal thera- 
pies inc 


— 


uding specific herbal formulas and recommendations for proper nutrition and exercise to 
ca eas: the herbal treatments. She is also careful to point out what treatments are and are not 


Pl g pregnancy and lactatio 


The Book contains vain appendices including: sources for perce aasing herbal products, as 
1] 


with information or databases on 
herbal practitioners or suppliers of herbal information oe products; and lists of journals devoted to 
the dissemination of information on herbal medicine 

If you are a woman who is interested in or a user of ee | remedies or a medical practitioner 
who is interested in incorporating herbal tl s medical practice, this is the book 
for you.—Debra K. Trock, Botanical Research institutect Texas, 509 Pecan, Fort Worth, TX 76102-4060, 
U.S.A, dtrock@brit.org. 


SIDA 19(4): 794, 2001 


TWO NEW SPECIES OF IMPATIENS (BALSAMINACEAE) 
FROM INDIA 


Muktesh Kumar Stephen Sequiera 
Botany Division Botany Division 
Kerala Forest Research Institute Kerala Forest Research Institute 
Peechi 680 653, Thrissur, Kerala, INDIA Peechi 680 653, Thrissur, Kerala, INDIA 
muktesh@kfri.org 


ABSTRACT 
Two species of i ee viz., I. sholayarensis and I. violacea are described from Kerala, India. 


Key Bal i hol is, Impatiens violacea, Kerala, India 


£ 


RESUMEN 


Se describen dos especies de Impatiens viz., |. sholayarensis e I. violacea de Kerala, India. 


INTRODUCTION 


The genus Impatiens L. consists of about 900 species distributed in tropical 
Africa and South East Asia, with a few representatives in North America, Eu- 
rope and East Asia (Grey- Wilson 1985). Of these, 200 species were reported from 
India (Biju 1999) and are mainly distributed in the Eastern Himalayas and the 
Western Ghats—the megacenter of endemism in India (Nayar 1996). Among 
these, there are 86 species of Impatiens reported so far from the Western Ghats 
region (Augustine et al. 1999 

During our studies on the epiphytic flora of the Western Ghats, two epiphytic 
balsams were collected with striking differences. On critical examination they 
were found to be new to the science and are described and illustrated here. 


1. sa sholayarensis M. Kumar & Sequiera, sp. nov. (Figs. 1, 2). Tyee INDIA. 
A: Thrissur Dt., Sholayar, alt. 90 Om, 9 Oct 1998, Sequiera 


fe Pad ge li es ] 120 - oye: - lake; lel lal | fos ee 


oO oO © Oo 
lis dorsalibus c. 35 mm longa viridi-flava lab bell unco prominent, peas coed 


lakh 
5 


1 ery | Ts binet 
r 


, petalarum lateralium 


Epiphytic on trees, glabrous; stems thick, fleshy up to ie cm long, green. Leaves 
crowded towards apex of the stem; petioles 2.5-8 cm long, channeled, light 
greenish, with two prominent stipitate gland the base of the blade; lamina 
ovate to elliptic-lanceolate, 2.3-8 x 1.3-3.2 cm, acute at apex, shortly attenuate 
at the base, membranaceous, glabrous, light green above, silvery white below, 
margins crenulate, crenules 4-17 mm apart, crenatures short, furnished with 
short incurred bristles, lateral veins up to 4 pairs, indistinct. Inflorescence a 


SIDA 19(4):795 — 801. 2001 


796 BRIT.ORG/SIDA 19(4) 


Fic. 1. Impatiens snolayarenstaN pulley eseqillera: A. Habit. B. Single flower. C. ee : Dorsal petal. E. Lateral petal 
(outside view). F. 1 ).G, Lower sepal. H. Pistil. | Gynoecium.J.$ 


KUMAR AND SEQUIERA, NEW SPECIES OF IMPATIENS FROM INDIA 797 


F 7 ol 


Fic. 3. Impatiens violacea M. Kumar & Sequiera. 


798 BRIT.ORG/SIDA 19(4) 


2-flowered raceme, peduncles 1.3-2 cm long, pedicels 2.5-5 cm long, filiform, 
greenish-white. Flowers greenish-yellow with crimson red; lateral sepals 0.8-1 
x 0.2 cm, linear-ovate, slightly falcate, glabrous on both surfaces; lower sepal 
saccate, compressed, 1-3.5 x 0.5-1.1 cm, glabrous, greenish-yellow with crimson- 
red, mouth descended, with a prominent hook at the lower side, spur curved, tip 
obtuse, white; dorsal petal erect, 0.6-1.1 x 0.4-0.9 cm, gland dotted, glabrous; lateral 
petals united at the base, 3-lobed, 1.7-1.9 x 1-12 cm, upper lobes with a small spine 
and a pouch. Androecium 0.8-1 cm long; stamens 5. Ovary elliptic-lanceolate, 
1-12 x 0.3m, slightly falcate, glabrous, stigma straight, toothed. Capsule elliptic- 
lanceolate, up to 1.2 cm long, glabrous. Seeds pear shaped, tip ciliate. 

This species is allied to Imaptiens kulamavuensis Pandurangan & VJ. Nair, 
but differs in having moniliform stem; crowded, glabrous leaves; glabrous lateral 
sepals; 35 mm long, greenish yellow with crimson red lip with a prominent 
hook at the mouth; keeled, glabrous dorsal petal; lobules of the lateral petals 
with a spine & pouch and a enaped ciliate seeds. 

Distribution.—I is known only from its type locality. 

Ecology and conser vation status.—Impatiens sholayarensis occurs in the 
evergreen forests at an altitude of 700-800m (type collection has ‘900m’). In 
the type locality only about 20 populations exist in the wild. Due to its restricted 
distribution and less number of occurrences this taxon is considered as a criti- 
cally endangered species. Loss of habitat and destruction of host trees and hu- 
man interference are the major threats in the area. 

Etymology.—The species is named after its type locality. 


2. Impatiens violacea M. Kumar & Sequiera, sp. nov. (Figs. 3, 4). Type: INDIA. KERALA 
Idukki Dt, Munnar, Pettimudi, Way to Edamalakudy, alt. 2000m, 25 Aug 1998, Sequiera 20731 
(HOLOTYPE: KFRI; ISOTYPE: MH). 


een anadiona aumtlis peu! ) vioiaeeo, folit crenaturis, brevibus, floribus violaceis, sepalis 


,apic eunco prominenti, pet alis 
dorsalibus viridi- flavis, petals ea ‘bus 3- ete reba eee quibusque sacco minuto, 


stigmatis apice obtuso s diff 


oO 


Epiphytic on trees, glabrous, stems thick, 1-1.5cm in Soci up to lO cm high. 
Leaves crowded towards the apex of the stem; petioles up to 3.5 cm long, violet, 
channeled, with 2 prominent stipitate glands near the base of the blade; lamina 
lanceolate to ovate-lanceolate, 2-6 = 1.4-4 cm, acute at apex, truncate at base, 
margins crenulate-serrate, violet, crenules 4-6 mm apart, crenatures short, fur- 
nished with short incurved bristles, violet. Inflorescence a 1-4 flowered raceme, 
peduncles 1.5-2 cm long, pedicels 2-4 cm long, violet. Flowers violet; lateral 
sepals linear-lanceolate, glabrous, 0.8-1 x 0.2-0.3 cm, slightly keeled on upper 
side; lower sepal saccate, 1.75-2 x 0.6-1.1 cm, violet, glabrous, wrinkled, with a 
short prominent hook at the mouth; dorsal petal erect, 0.6-0.8 x 0.5-0.7 cm, 
greenish yellow, gland dotted, deeply hooded, hood dark green; lateral petals 
fleshy, 0.8-1 x 0.6-0.8 cm, 3-lobed, gland-dotted, with a small pouch in each 


— 
UO 


KUMAR AND SEQUIERA, NEW SPECIES OF IMPATIENS FROM INDIA 799 


Fic. 4. Impatiens violacea M. Kumar & Sequiera. A. Habit. B. Single flower. C. Sepal. D. Dorsal petal. E. Lateral petal (out- 
side). F. Lateral petal (inside). G. Lower sepal. H. Pistil. |. Gynoecium. J. Androecium. K. Seeds. 


Taste 1. Comparison between Impatiens kulamavuensis and Impatiens sholayarensis. 


BRIT.ORG/SIDA 19(4 


) 


Impatiens kulamavuensis 


Impatiens sholayarensis 


Stems less than 1. cm 
Stoloniferous 
Leaves alternate 


Stems up to5 cm 
Stem moniliform 
me fascicled 


Leaves glanduliferous ves glabro 
Lateral sepal hairy out side eae sepal a aie on both the surfaces 
Lower sepal up to 15 mm long Lower sepal up to 35 mm lon 
Lower sepal without hook Lower sepal with a hook at i mouth 
Lower sepal greenish-yellow Lower sepal greenish-yellow with crimson red 
Dorsal petal glabrous Dorsal petal gland dotted 

d 


Lobules oh lateral petals without spine Lobules fe lateral petals with a spine 
and pouc 


and 
Stigma ane with a silky pubescent apex Stigma on with toothed apex 
Ovary ellipsoid Ovary elliptic-lanceolate, slightly falcate 
Seeds smooth Seeds ciliate 


4st 


TABLE 2.cComparison between Impatiens violacea and Impatiens viridiflora. 


|. violacea |. viridiflora 

Petiole violet Petiole green 

Leaves green with violet tinge, nerves violet Leaves dark green, nerves green 

Crenatures short, violet Crenatures long, green 

Peduncles violet bearing 1-4 flowers duncles green bearing 1-3 flowers 

Pedicel violet Pedicel green 

Flowers violet Flowers green 

Sepals large, dark green, linear-lanceolate, not Sepals large, dark green, linear-oblong, 
deflexed or reflexed, slightly keeled on deflexed or reflexed, slightly not keeled. 
upper side. 

Lower sepal saccate, wrinkled, violet, Lower sepal saccate, wrinkled, green, spur 
spur laterally compresse laterally Sees 

Dorsal petal 2-lobed, veliouee Dorsal, 2-lobed, gree 

Lateral petals 3-lobed, yellow a small pouch Lateral petals 2- aaa green without small 
in each lateral lobules pouch in each lateral lobules 

Capsule elliptic ovate Capsule ellipsoid 


KUMAR AND SEQUIERA, NEW SPECIES OF IMPATIENS FROM INDIA 801 


lateral lobules. Androecium 0.5-0.6 cm long, stamens 5 in number. Ovary ellip- 
tic-ovate, glabrous, 0.4-0.5 x 0.15-0.2 cm; style short; stigma with an obtuse silky 
pubescent tip. Capsule elliptic-ovate, l-1.1 x 0.3-0.4 mm, glabrous. Seeds almost 
tetragonal, smooth. 

This species is similar to Impatiens viridiflora Wight in its structure and 
shape but differs in having violet colored petiole, short crenatures; violet col- 
ored flowers, lateral sepals linear lanceolate, glabrous, lower sepal violet col- 
ored and wrinkled with a prominent hook at the apex; dorsal petal greenish 
yellow, lateral petals 3 lobed with small pouch in each lateral lobules; stigma 
with obtuse silky pubescent tip and tetragonal, smooth seeds. 

Distribution.—Impatiens violacea is known from Pettimudi area of Munnar, 
Idukki Dt., Kerala, India. 

Ecology and conservation status.—This species occurs only in the montane 
shola forests at an altitude of 1800-2200 m. Only 50 individuals exist in the 
wild habitat. Cultivated in KFRI green house for in situ conservation. Due to its 
restricted distribution and less number of individuals occurring in the wild 
the species is considered as critically endangered. Human interference and 
habitat destruction are the major threats for this balsam. 

Etymology.—The species is named after its violet color of the lateral sepal. 


ACKNOWLEDGMENTS 


We are thankful to the Chief Conservator of Forests (Wildlife), Kerala Forest 
Department, Kerala for the financial support and J.K. Sharma, Director, KFRI, 
for the facilities and encouragements. Thanks are also due to J.F Veldkamp, 
Leiden University, The Netherlands, for the Latin diagnoses. 


REFERENCES 


Auaustine, J., N. SASIDHARAN, and M. Sivapasan. 1999. Balsams of Periyar Tiger Reserve, South- 
ern Western Ghats, Kerala. In: M. Sivadasan and Philip Mathew, eds. Biodiversity, tax- 
onomy and conservation of flowering plants, Mentor Books, Calicut. Pp 275-292. 

Buu, S.D. 1999. Chronicles of discoveries—The pursuit of plants. In: T.M.Manoharan, S.D 
Biju, T.S. Nayar and PS. Easa, eds. Silent Valley—Whispers of Reason, Kerala Forest De- 
partment, Kerala. Pp. 145-174. 

Grey-WiLson, C. 1985. The family Balsaminaceae. In: M.D. Dassanayake and F.R. Fosberg eds. 
A revised handbook to the flora of Ceylon. Amerind Publishing Co., New Delhi. 

Nayar, M.P. 1996. Hot spots of endemic plants of India, Nepal and Bhutan. Tropical Botanic 
Garden & Research Institute, Thiruvananthapuram. 


802 BRIT.ORG/SIDA 19(4) 


Book REVIEW 


LAURENCE PARENT (photographs) and Jor Nick Parosk! (text). 2001. Texas Mountains. 
(ISBN 0-292-76592-4, hbk.). University of Texas Press, PO. Box 7819, Austin, 
TX 78713-7819, U.S.A. (Orders: http://www.utexas.edu/utpress, 512-471- 
4032). $39.95 hbk., 156 pp., 120 color photos, 10" x 11". 


As a child, this reviewer was captivated by the high FuppeO T mountains of _Colbrade and WOH: 
Asan adult, a trip to Big Bend National Park it career change, 
many subsequent trips to the area, and a love for all aepetis a the natural history a wild places. This 
beautiful book with photographs by Laurence Parent and text by Nick Patoski, takes the reader on an 
incredible journey to the pieainialine region of the Trans-Pecos in far west Texas. The com seen 


o photographs and prose describe places that are both rugged and beautiful, and much more entic- 


han the over-exploited areas of the intermountain west. | looked at the details in every picture 
and found myself delighted every time [came upon a particular view and could say, “Wow! I’ve been 


ga 


there.” 

Parent’s photographs capture the mountains and the vegetation of the Trans-Pecos in all of 
their glory through the seasons. The vibrant reds, oranges and yellows of a desert sunset or sunrise 
are contrasted with the rich shades of Sogn of the rivers, canyons and high mountain forests. The 


breathtaking agoraphobic vi phot phs of rocky pinnacles, Ponderosa pine for- 


ests and the myriad of strange and austal deca plant ts. 
-atoski’s text provides a glimpse into the lives of the rugged individuals who inhabit these 
beautiful and remote parts of Texas. Interviews with people who are relative newcomers to 


gion as well as those who have made the Trans-Pecos their home for generations, ppvicema into 


oe | remote 


what draws people to live and work in places that are considered by some to be 
A love of the land is evident in all of the people Patoski came to know through this work. 

If you love mountains, if you love the west, if you love wide open spaces and the concepts of 
self-reliance and individual freedom you will love this book.—Debra K. Trock, Botanical Research In- 
stitute of Texas, 509 Pecan, Fort Worth, TX 76102-4060, U.S.A., dtrock@brit.org 


SIDA 19(4): 802. 2001 


ANOTHER NEW SPECIES OF TRIPOGON (POACEAE) 


FROM INDIA 
C.N. Sunil A.K. Pradeep 
Department of Botany Department of Botany 
S.N.M. College, Maliankara University of Calicut 
Kerala 683 516, INDIA Kerala 673 635, INDIA 
ABSTRACT 


A new species of Poaceae, Tripogon ravianus Sunil & Pradeep from the Western Ghats of Tamil Nadu, 
India is described and illustrated 


RESUMEN 


ie de Po , Tripogon ravianus Sunil & Pradeep de los Western 


Ghats de Tamil Nadu, India. 


During previous floristic investigations on the Western Ghats of Kerala, India, 
two new species of Tripogon Roem. & Schult. were described (Pradeep & Sunil 
1999). Since then, yet another remarkable undescribed species of Tripogon from 
Pykara on the Western Ghats of Tamil Nadu, South India has been discovered. 
It is described here as new. 

Tripogon ravianus Sunil & Pradeep, sp. nov. (Fig. 1). Type: INDIA.TAMIL NADU. Niiairi 


Dt. Pykara near Udagamandalam, 1990 m, 3 Dec 2000, Sunil 2176 (HOLOTYPE: BRIT, ISOTYPEs: 
CALI, kK, L, NY). 


Tripogoni anantaswaniano affinis, T. ravianus racemis 15-40 cm longis, spiculis 10-25 mm longis et 
flosculis 6-14 instructis, glumis inferioris profunde 2-lobatis, lemmatis apicaliter 4-lobata facile 
distinguendus. 

Tufted perennial. Culms 24-70 cm tall; nodes glabrous. Leaf blades 5-35 cm 
long, 2-4 mm wide, the margins slightly scabridulous or scabrid especially to- 
wards the base, the upper surface pubescent with short white, papillose hairs, 
glabrous abaxially. Leaf-sheaths 3-9 cm long, glabrous, apex pubescent witha 
tuft of 2-3 mm long hairs; ligule a fine membrane. Racemes terminal, 15-40 
cm long, with 15-48 spikelets; rachis smooth, glabrous. Spikelets 1-2.5 cm long, 
linear, secund, dark gray, 6-14-f lowered, distant, up to 7 mm apart. Lower glume 
2-4 mm long, 1-2 mm wide, coriaceous, broadly ovate, unequally 2-lobed, 1- 
veined, the inner margin of smaller lobe slightly serrulate or entire, apex of 
both lobes acuminate. Upper glume 5-6 mm long, 1.5-2 mm wide, lanceolate, 
notched at apex and awned, coriaceous, l-veined; awn 1.5-2 mm long, scabrid, 
the lateral lobes ca. 1 mm long, lanceolate, acute. Lemma 3-4 mm long exclud- 
ing awns, 2-3 mm wide, coriaceous, dark gray, 3-veined, 4-lobed at the apex; 
awns 3, scabridulous, the central awn 4-5 mm long, the lateral awns 3-3.5 mm 


SIDA 19(4): 803 — 806, 2001 


— 


BRIT.ORG/SIDA 19(4) 


Fic. 1. Trii i ¢ il & Pradaan Hahit. bh Sheath 


f lamina. c. Portion of lamina eaunaee: d. 


Spikelet. e. ‘Lower glume. f. ance a g. Lemma. h. Palea. i. St 


j Lodicule k Caryop 


SUNIL 


long; lobes between awns ca. 2 mm long, acuminate; callus tufted; hairs up to 1 
mm long. Palea 2.5-3.5 mm long, 1-1.5 mm wide, obovate, hyaline, 2-keeled, the 
keels ciliate, apex shortly 2-lobed, the lobes acuminate. Lodicules 2, 0.5-0.75 
mm long, wedge-shaped. Stamens 3; anthers 1.5-2 mm long, oblong, creamy 
yellow; filaments 1-15 mm long. Ovary 0.75-1 mm long, ca. 0.5 mm broad, obo- 
vate; styles 2,ca.1mm long; stigma 1-1.5mm long, feathery, creamy white. Cary- 
opses 1.8-2.3 mm long, 0.4-0.7 mm wide, oblong -cylindric. Chromosome num- 
ber unknown. 

Distribution.—Tripogon ravianus is endemic to Tamil Nadu and is known 
only from the type locality, Pykara near Udagamandalam in Nilgiri District of 
Tamil Nadu. 

Ecology and phenology.—This species grows from 1500-2200 m elevation 
on road cuts and wet rocky hillsides; flowering October to late December. 

Etymology.—The specific epithet honors Prof. N. Ravi, teacher of the first 
author and a grass specialist now with the Tropical Botanic Garden and Re- 
search Institute, Trivandrum 

Tripogon ravianus is closely allied to T. anataswamianus Sreek., VJ. Nair & 
N.C. Nair,a species endemic to Kerala (Sreekumar & Nair 1991). It can be distin- 
guished by its 15-40 cm long racemes, 10-25 mm long spikelets with 6-14 flo- 
rets, deeply 2-lobed lower glumes and apically 4-lobed lemma. 


KEY TO THE SPECIES OF TRIPOGON IN SOUTH INDIA 
1. Culms thickened below by the persistent leaf-sheaths; leaves equitant, rigid, 
T. pungens 


pungent 
. Culms not tas below by the persistent leaf-sheaths; leaves not equitant, not 
ch or pungen 

. Lemmas ce at apex into 2 lobes, awned in the cleft; lobes awned or not. 

3. Central awn of the lemma flexuous, capillary, several times as long as lemma 
T. capillatus 


3. Central awn of the lemma straight or curved, not more than twice as long as 
he lemma 
4. Spikelets 10-40 mm long; paleas broadly ee T. wightii 
4. Spikelets 4-17 mm long; paleas not broadly win 
5. Culms less than 20 cm tall; central awn aie strongly twisted 
below 


T. pauperculus 
5. ae greater than 20 cm tall; central awn not geniculate or twisted. 

Perennials with wiry roots forming tufts; leaves glabrous 

7. Leaves and culms glaucous, involute; lower elaine lobed on 


e-side T. jacquemontii 
7. Leaves and culm not glaucous, flat; lower glume symmetrical 


T. vellarianus 
6. Perennials with fibrous roots, forming a close turf; leaves villous on 

the upper surface 

8. Spikelets 5-8-flowered; lower glume 2 mm, notched on one-side 


T. sivarajanii 


BRIT.ORG/SIDA 19(4) 


8. Spikelets 1-2-flowered; lower glume 0.7 mm, symmetrical 


T. roxburghianus 

2. Lemmas cleft at apex into 4 or 6 lobes or with a definite lobe between each 
lateral awn and the central; outer lobes if present awned o 

9. nee nally ligules i Heelers keels of the paleas nee 

9. 


T. bromoides 
ilign 


iles conspicuous, thin, membranous; keels of the 


te Allee 

10. Leaves glabrous; lemmas cleft at apex into 6 lobes 

10. si pubescent; lemmas cleit at apex ne 4 lope 
. Culms 10-20 cm tall:| | ly lot 


T. anantaswamianus 


lobed on one-side above 
the middle; central lobes of the lemmas ovate-acute T. narayanae 
. Culms 24-70 cm tall; lower glume deeply lobed on one-side below 
the middle; central lobes of the lemmas lanceolate 


T. ravianus 


ACKNOWLEDGMENTS 
We would like to thank Thomas A. Cope, Royal Botanic Gardens, Kew for his 
valuable comments on our specimens and P. Manimohan, Calicut University 
for correcting the Latin diagnosis. The illustration was prepared by VB. Sajeev. 
North Paravur. 
REFERENCES 


PraveeP, A.K. and CN, Suni. 1999. Two new species of Tripagon (Poaceae) from India. Sida 
18:809-8 14. 


Sreekumar, PV.and V.J.Nair. 1991. Flora of Kerala-grasses. Botanical Survey of India. Calcutta. 


IPOMOEA SORORIA (CONVOLVULACEAE), 
A NEW SPECIES FROM YUCATAN, MEXICO 


Daniel F. Austin Jose Luis Tapia Munoz 
Arizona-Sonora Desert Centro de Investigacion Cientifica 
Museum C CCI 
2021 N. vee Road Calle 43 No. 130, Colonia 
Tucson, AZ 85743, U.S.A. Chuburnd de Hidalgo, 97200 Mérida 
Yucatan, MEXICO 
ABSTRACT 


Ipomoea sororia, a new species in Ipomoea section Leptocallis is described from the Yucatan Penin- 
sula of Mexico. This Species, apparenty endemic’ toa small region of low deciduous forest in the NW 

of the e pen ninsula, is similar to the more lL. ternifolia and I. costellata. The new spe- 
cies differs from those in having cordate seca ee and three rows of largely reflexed excresences 
on the outer sepals. 


RESUMEN 


Se describe alee sororia, una nueva especie ae Ipomoea seccion Lepiocails ee la Peninsula de 
Yucatan, México. Esta especie, ay baja decidua 
en la zona NW de la peninsula, es similar a las wae ies de mas ampli distribucién, I ternifoliae I. 
costellata de la costa pacifica. La nueva esp a por presentar los sépalos con las bases 
cordadas y tres hileras de crestas reflejas en los pee exteriores. 


During preparation of the treatment of the Convolvulaceae for Flora 
Mesoamericana (Austin et al, in prep.), and collaboration on the Flora Ilustrada 
de la Peninsula de Yucatan (Austin and Tapia, in press), collections were found 
that match no known species. These specimens are similar to those in sect. 
Leptocallis (McDonald 1995), with which they are thought to be allied. Com- 
parison with the keys in McDonald's paper, and with available material in CICY 
and MEXU make it clear that the plants do not fall within the known variabil- 
ity of species from Mexico and Mesoamerica. Therefore, we describe the new 
species. 


Ipomoea sororia D.F Austin ce Pei sp. nov. (Fig. 1). TYPE: MEXICO. YUCA ATAN: 
Mpio. Dzemul, al S del entronque a las ruinas Xtampu. Selva baja caudicifolia con ceas 
columnares, 9 Nov 2000, Tapia & eee 1120 (HOLOTYPE: CICY, IsOTYPES: F FAU, K eres 
MO, NY, UCAM, XAL). 


Ipomoea ternifoliae et L. costellatae pri Isp ile, sed sepali is seriel il 
rugarum vel bullarum vel fere papillarum et praesertim cordatis basibus distinguit 
Twining herbs, annuals with fibrous root-systems, st climbing or prostrate, 


toca 2 m, glabrous or glabrescent. Leaves pedately 5-7-lobed, more or less reni- 
form to orbicular in outline, 2.5-5 cm long, 3-7 cm wide, the lobes entire, or the 


SIDA 19(4): 807 — 810. 2001 


808 BRIT.ORG/SIDA 19(4) 


Fic. 1./ ia D.F. Austin & J.L.Ta apia sp.nov. A General habit, B fl juri hesis, C Outer sepal in 
lateral view. Based on J.L. ae G. Carnevali 1120 (CICY, F, FAU, K, MEXU, MO, NY, UCAM, ‘XAU). Drawing by Rossana 
Marrufo (CICY). 


AUSTIN AND TAPIA, A NEW SPECIES OF IPOMOEA FROM YUCATAN, MEXICO 809 


basal lobes with 1-3 lobes, glabrous, the base cordate, the apex broadly obtuse 
to rounded, the petioles 2-3 cm long. Inflorescences cymose but mostly soli- 
tary on peduncles 0.5-3.5 cm long, the pedicels 4-5 mm long, the bracteoles 
lanceolate, 0.75-2 mm long. Flowers 1(-2); sepals more or less equal or the outer 
somewhat shorter, 5-8 mm long, the outer ovate, basally cordate, with a lan- 
ceolate terminal acumen 1-2 mm long, reflexed, glabrous, mostly with 3 rows 
of rugae or bullae to almost papillae, the inner 6-8 mm, ovate, long acuminate, 
glabrous, borders hyaline, the acumen 3-4 mm long, glabrous; corolla 2-2.5cm, 
funnelform, the limb white, the limb faintly 5-lobed, the throat lavender; the 
stamens included, unequal, 8-12 mm long, white, the filaments basally glan- 
dular-pubescent, inserted 4 mm above the base of the corolla tube, anthers erect, 
dithecal; the ovary lobate, 1 mm diameter, glabrous, the style white, glabrous, 
6-8 mm long, the stigma in contact with stamens, capitate, bilobate, papillose 
surface. Fruits capsular, ovoid, 5 mm long, 4 mm wide, 4-valvate, the 2-locules 
well-delimited and somewhat 2-lobed, tan to brown, the valves membrana- 
ceous, glabrous; seeds not seen. 


Paratypes: MEXICO. ames A PaTApecne 2 km Noroeste de China, 10°47°15"N, 90°31'50"W. 
Selva b ia, suelo negro, inundable, 23 Nov 1998, C. Gutierrez 
B. 6056 (CICY, MEXU, UCAM): Mpio. Calkini, 10 km al O de Tankuche, sobre el camino a Punta Are 
nas. Zona transicional entre selva baja y manglar con Hematoxyllum, 1 Dec 1988, Cabrera & Cabrera 
15304 (MEXU). 


Etymology—the epithet is based on Latin soror, -is, meaning sister. This is in- 
tended to draw attention to the similarity between this species and Ipomoea 
ternifolia. 

Distribution.—México. Campeche, Yucatan. 8-10 m. Flowering November- 
December. Endemic to southern Mexico. 

This species is noted as growing with mangroves and Hematoxyllum in 
one site, and in others with Bursera simaruba, Metopium brownei, 
Gymnopodium floribundum, Croton spp., Pterocereus gaumeri, Cienfugosia 
yucatanensis, Bakeridesia gaumeri, Myrmecophila christinae and Tillandsia 
fasciculata. Apparently, the species is tolerant of the drier deciduous forest and 
inundation where this habitat approaches mangroves. Soils are recorded as 
litosols mixed with rendsins, shallow and with outcrops of calcareous rocks, 
flooded during the rainy season (suelo negro somero muy ped regoso, suelo negro 
inundable). 

Ipomoea sororia is similar to not only I. ternifolia, but also other species in 
sect. Leptocallis (G. Don) J.A. McDonald (1995). The cordate basal portion and 
ornamentation on the sepals clearly distinguish I. sororia from I. ternifolia. Habi- 
tat will distinguish I. sororia from all related species except I. ternifolia and L. 
costellata, which share the lowland deciduous communities 

In McDonald’s (1995) key these plants lead to couplet No. 6, where the co- 
rollas are shorter than either choice, but longer than | cm. Indeed, the variation 


810 BRIT.ORG/SIDA 19(4) 


within L. plummerae makes it similar, but I. sororia is prostrate, not erect. Then, 
these plants fall between the traits at couplet No. 7, which leads on one hand to 
I. perpartita and on the other to L plummerae and I. ternifolia. Ipomoea sororia 
has the pedatisect segments, but the corolla tube is darker than the limb of 1 
perpartita. Moreover, the leaves are not twice dissected as in I. perpartita. 


REFERENCES 


Austin, D.F., JA. McDonato, and G. Murcuid S. (in preparation). Convolvulaceae. In: Davidse, 
G., M. Sousa S.and A.O. Chater, eds. Flora Mesoamericana 

Austin, D.F.and J.L. Taria M.(in press)./pomoea. In:Flora llustrada de la Peninsula de Yucatan. 

McDonato, J.A. 1995. Revision of Ipomoea section Leptocallis (Convolvulaceae). Harvard 
Pap. Bot. 6:97-122. 


TAXONOMIC REVIEW OF CHRYSOGONUM 
(ASTERACEAE: HELIANTHEAE) 


Botanical Research Institute of Texas 
09 Pecan Stre 
Fort Worth, TX 76102-4060, U.S.A. 


ABSTRACT 


Three allopatric varieties are erecopmizee wae c st sega yueami ene monner var. 


virginianum, with non leafy (Gulf coastal 
plain) var. australe, with flagellate stolons and only leafless aerial stems; and the geographically 
intermediate var. brevistolon, var. nov. with stolons of mieHmeGiale Bee and mostly Healy aerial 


stems. These three taxa are ees by other minor 
parently occur, but uncommonly, DEEWER var. veeaanin ands var. hrevistalon Variety australe is 
more strongly separated in geograr ge and more distinct in morphology. 


RESUMEN 


Cc . ] ] ] 


Chrysogonum virginianum: la nortena var. virginianum, 
con ejemplares no estoloniferos y la mayors es ie tallos aéreos con Holes la surenia (Llanura costera 
el Golfo) var. austr in hojas; y la intermedia 


y 
geograficamente var. brevistolon, var. nov., con estolones de tongitie intermedia y con la aus de 
Se 


los tallos aéreos con hojas. Estos tres taxa se separan por 
encuentran intermedios aparentemente, pero raramente, entre var. vi rginianum . Var. pevetolon: 
La variedad australe se separa mas en el rango geog y tiene una morfologia mas distinta. 


Chrysogonum L. isa monotypic genus represented by C. virginianum L. “Green- 
and-gold”) of the eastern United States, ranging from Mississippi, Alabama, and 
Florida north to Ohio and Pennsylvania. It has mainly been treated asa single 
species with two varieties (e.g., Ahles 1968; Stuessy 1977; Cronquist 1980). 
Chrysogonum australe Alexander ex Small was originally published as a sepa- 
rate species (Small 1933) but reduced in rank by Ahles (1964), without com- 
ment, to C. virginianum var. australe (Alexander ex Small) Ahles. In an unpub- 
lished treatment of the flora of “the Carolinas and Virginia,” Weakley (in 
manuscript) recognizes C. australe as a separate species but provides no ratio- 
nale for the taxonomic judgement. 

Stuessy (1977) distinguished two varieties (var. virginianum vs. var. australe) 
based on plant height (15-35 cm vs. 3-14 cm), stem orientation (erect vs. often 
decumbent), and disposal of leaves (primarily cauline vs. nearly all basal). 
Stuessy mapped intermediates, not only in the area of North Carolina and South 
Carolina where the ranges of the two taxa (in his concept) meet, but also nearly 
throughout the geographic range of the whole complex. Ahles’ (1968) distinc- 
tion of the two taxa in North Carolina and South Carolina was based primarily 


SIDA 19(4): 811 - 820. 2001 


812 BRIT.ORG/SIDA 19(4) 


B. Var. brevistolon 


C. Var. anal { 


Fic. 1. Growth form of the three varieties of Chrysogonum virginianum. A. Var. virginianum. B. Var. brevistolon. C. Var. 
australe 


on stolon production and his concepts and mapping are closer to what is pre- 
sented here. 

In the present study, variation patterns within the Chysogonum virginianum 
complex are reviewed and three morpho-geographic entities are recognized (Fig. 
1; Fig. 3). Their morphological distinction is based primarily on production (or 
not) of stolons, length of stolon internodes, whether or not the flowering stems 
are leafy, and height of flowering stems. Variation in other features (vestiture, 


NESOM, TAXONOMIC REVIEW OF CHRYSOGONUM 813 


leaf morphology, capitular and floral morphology, and cypsela and elaiosome 
morphology) is overlapping and/or parallel among the three taxa. 

Data for the present study are from the field and from herbarium speci- 
mens. Map recordsare from BRIT, MO, NCU, OS, SMU, TEX-LL, USCH, and VDB, 
with some distribution points added from various references (Strausbaugh & 
Core 1977: Harvill et al. 1981; Fisher 1988; Brown & Brown 1984; Rhoads & Klein 
1993). 


TAXONOMY 
Chrysogonum L., Sp. Pl. 2:920. 1753. Type: Chrysogonum virginianum L. 
Diotostephus Cass., Dict. Sci. Nat. 48:543. 1827. TyPE: Diotostephus repens Cass. (see comments 

below, under var. brevistolon). 
Chrysogonum virginianum L., Sp. PI. 2:920. 1753. Type: US.A. VIRGINIA: [No other data]. 

J. Clayton 298 (LECTOTYPE, Stuessy 1977 [or ISOLECTOTYPE” fide Reveal 1989 by annotation}: BM, 

John Clayton Herbarium image!). 
Plants herbaceous, perennial, arising from a fibrous-rooted rhizome up to 5cm 
long, sometimes stoloniferous, sometimes producing basal offsets. Stems erect 
to erect-ascending (flowering), mostly 10-30 (rarely to 50) cm, or prostrate (non- 
flowering stolons), up to 50 cm, villous. Leaves opposite, basal and cauline, 3(- 
5)-nerved from the base, elliptic-ovate to deltate-ovate, 2.5-10 cm, petiolate, 
crenate-dentate to subentire, minutely strigose-hirsutulous to hirsute-villous, 
also minutely stipitate-glandular above and beneath with hairs ca. 0.1 mm long. 
Capitula axillary, 1-2 per node, solitary on naked peduncles 2-22 cm or ap- 
pearing to arise from the basal rosette; involucres cupulate-hemispheric, 7-10 
cm wide; phyllaries 10 in 2 series, outer oblong-oblanceolate, distal half folia- 
ceous and spreading, inner greenish, rigidly erect and somewhat scarious, re- 
ceptacles flat to shallowly convex; paleae oblanceolate, ca. 4 mm long, scarious. 
Ray florets 5(-6), pistillate, fertile, corolla laminae yellow, broadly elliptic, 6-17 
mm, apically 3-toothed; cypselae each basally adnate to a single inner phyl- 
lary, 3 paleae, and 3 sterile disc florets, this “cypsela-complex” deciduous as a 
unit. Disc florets ca. 25-50, functionally staminate, corollas yellow, 2.5-2.7 mm, 
tubes much shorter than the narrowly funnelform throats, lobes 5, deltate; ova- 
ries sterile; anthers brown to black; styles filiform, the branches mostly undi- 
vided: ovaries filiform, epappose. Cypselae (ray) obovoid, strongly flattened ra- 
dially, 3-4.5 mm x 2-2.5 mm, blackish-brown; pappus a low, irregular, 
‘half-cup-shaped’ crown, incomplete on the adaxial face. Base chromosome 
number, x = 16. 

An elaiosome is developed at the base of the cypsela-complex (formerly 
termed “achene-complex”). This is a fleshy, oil-bearing structure elaborated at 
the base of the phyllary and the two associated outer paleae (Fig, 2). All three 
taxa of Chrysogonum produce elaiosomes; variation in the degree of elabora- 
tion of the elaiosome tissue occurs but without apparent geographical pattern. 


814 BRIT.ORG/SIDA 19(4) 


pappus 
pappus oe 


Wee pale 
pale ey 
LHe ovary 
Me | ie 
LAT cypsela 
phyllary phyllary 
elaiosome —— “ g—— elaiosome 


adaxial 


Fic. 2. Cypsela 


Native species of ants carry the cypsela-complexes into their nests, chew off 
the elaiosomes, and return the undamaged cypselae or the whole complexes to 
the surface (Nesom 1978). 

The stolons of var. brevistolon and var. australe presumably are derived from 
erect leafy stems (as in var. virginianum) where developmental processes have 
re-oriented them into lateral, prostrate or decumbent branches. In var. australe, 
the stolons commonly produce leaves and rooting plantlets only at the apex; 
another pair of leaves less commonly is produced proximal to the stolon apex. 
In var. brevistolon, the stolons are shorter and produce leaves more frequently 
at rooting nodes along the length. The basal offsets that are uncommonly pro- 
duced in var. virginianum apparently are essentially subterranean rhizomes 
produced from the caudex or main rhizome without the herbaceous nature that 
would indicate they are directly derived from erect branches. 

In all three varieties of Chrysogonum virginianum, the first (earliest-flow- 
ering) capitula are borne on leafless, ebracteate scapes. These scapes apparently 
are homologous with the leafless and ebracteate peduncles of later capitula that 
are raised on leafy stems in var. virginianum and var. brevistolon. In this view, 
the permanently ‘acaulescent’ flowering stems of var. australe remain in an 
early development stage and can be interpreted as neotonous. 

Chromosome numbers are invariantly 2n = 32 for var. virginianum and 
var. brevistolon. A count has not been made for var. australe sensu stricto. Chro- 
mosome counts reported below as “Nesom unpublished” were made by the au- 
thor in 1975 and 1976 from field-collected meiotic materials from North Caro- 


NESOM, TAXONOMIC REVIEW OF CHRYSOGONUM 815 


lina localities. In all observations, meiosis and tetrad formation were regular: 
16 bivalents were observed in populations from Lee, Montgomery, Moore, and 
Orange cos. (var. virginianum) and Wilkes Co. (var. brevistolon). Eight 
tetravalents were regularly produced in one population from Orange Co., N.C. 
(UNC campus); in one population from Wake Co., N.C. (Umstead State Park), 
four to eight tetravalents were commonly formed. 

All three taxa of Chrysogonum are now used as garden plants and are sold 
in nurseries under various horticultural names. 


KEY TO THE VARIETIES OF CHRYSOGONUM VIRGINIANUM 


1, Plants occurrin II liest flowering stems leafless, later 
ones leafy; leafy flowering stems mostly 15-35(—50) cm high var. Virginianum 
1. Plants colonial, mat-forming by stolons; Hei stems leafless and/or leafy; leafy 
flowering sens (if present) mostly 15-25 cm high 
Earliest f g stems leafless, nest 10 cm high, later ones leafy, 15-25 c 
high; longest stolon internodes 2-6 c var. aeMresioion 


2. All flowering stems leafless, 2-10 cm fick longest stolon internodes 12-60 cm 
var. australe 


la. Chrysogonum virginianum L. var. virginianum (Fig. 1) 


Ch rysogonum virginianum L. var. dentatum A. Gray, Bot. Gaz. 7:31. 1882. TYPE: U.S.A. MARYLAND. 
MONTGOMERY Co.: High Island in the Potomac River, 6 Jun 1881, J.D. Smith s.n. HOLOTYPE: GH)). 


Plants without stolons, not colonial or mat-forming. Stems (flowering) 15-35(- 
50) cm tall, the earliest leafless, becoming leafy. Leaves: blades deltate-ovate, 
basally abruptly cuneate to truncate or subcordate, minutely strigose-hirsutu- 
lous with ascending-appressed hairs 0.1-0.3 mm long. Capitula: peduncles 2- 
22 cm long; outer phyllaries 6-11(-15) mm long, 3-7 mm wide. Chromosome 
number, 2n = 32 (Montgomery Co., Md.-Kovanda 1978; Lee Co., Montgomery 
Co., Moore Co., Orange Co., and Wake Co., N.C.-Nesom unpublished). 

Flowering spring-early summer or longer with moisture (late Mar-Jun, - 
Jul and later). Moist to dry woodlands and forests, usually in partially sunny 
sites, especially in clearings and along edges; D.C, Ky., Md., N.Y., N.C,, Ohio, Pa., 

C., Va., W.Va. The report from New York (Mitchell & Tucker 1999) is noted as 
an escape from cultivation. 

Plants of var. virginianum do not produce leafy stolons, but basal offsets 
sometimes increase the size of the plant. Elongation of basal offsets or the primary 
rhizome may produce the effect of separate but interconnected plants, but such 
offsets/rhizomes are not commonly produced, except (apparently) in Maryland, 
D.C. and adjacent regions of Virginia. Plants of var. virginianum occur as dis- 
crete individuals but they are usually clustered, probably asa result of the pat- 
tern of cypsela-complex dispersal by ants. The flowering stems are erect; the 
first are leafless; they are later leafy as the lower internodes elongate. 


4) 


( 


BRIT.ORG/SIDA 19 


a 


fe Me 
aa 
\ 

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en 
+. 


f S 


O © var. virginianum 


A a Var. brevistolon 


@ var. australe 


MILES 


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Fic. 3. Geographic distribution o 


County, Tenn., 


comm.) and is not included in the present account. 


NESOM, TAXONOMIC REVIEW OF CHRYSOGONUM 817 


1b. Chrysogonum virginianum var. Dates as var now (Fig. 1). Type: 

U.S.A.SOUTH CAROLINA. LEXINGTON Co. Riverban old Saluda Mill 

site, shaded slope near ravine just S of tram line ee occasional herbs, somewhat trailing 

ut with flowering stems generally erect, 7 May 1993, J.B. Nelson 14,062 (HOLOTYPE: BRIT! 
ISOTYPE: USCH)). 


lonib t habitu coloniali; differt a var. australi stolonibus brevioribus 


Differt a var. 
foliaceis internodiis brevibus. 


mags repens Cass., Dict. Sci. Nat. 48:544. 1827. Chrysogonum diotostephus DC,, Prodr. 5:510. 
m. nov. illegit.). TYPE: “Nous avons fait cette description, générique et spécifique, sur 

un trés petit énchantillon sec, incomplet, en mauvais état, et dont nous ignorons T’origine” 
(HOLOTYPE: P?). As noted by a. RO Re of this taxon is explicit enough 
that its identity as C. virginianum seems reasonable, and de Candolle, who saw the specimen, 
treated it within Chrysogonum (as C. diotostephus, see eee lisa association here with var. 
brevistolon rather than var. australe is based on de Candoll t f its habitat as “in 


ya 


siccis pinguibus Carolinae.” 


Plants stoloniferous, colonial and mat-forming; stolons (leafy above-ground 
runners) with longest internodes 2-6 cm, these often becoming lignescent, par- 
tially to fully buried and rhizome-like. Stems (flowering) mostly 2-25 cm tall, 
the earliest leafless, becoming leafy. Leaves: blades deltate-ovate, basally 
abruptly cuneate to truncate or subcordate, strigose-hirsutulous with hairs 0.5- 
0.3mm long. Capitula: peduncles 2-22 cm long; outer phyllaries 6-10 mm long, 
3-4 mm wide. Chromosome number, 2n = 32 (Clarke Co., Ga.-Jones 1968; Lan- 
caster Co., S.C. -Solbrig et al. 1972; Rutherford Co., N.C.-Morton 1981; Wilkes Co., 
N.C.-Nesom unpublished). 

Flowering spring-early summer (Mar-May, -early Jun). Moist to dry wood- 
lands, usually in partially sunny sites; Ala. Ga., Ky. N.C, S.C, Tenn. 

Variety brevistolon differs from var. virginianum primarily by elongation 
and lateral re-orientation of the basally produced stems of the latter into sto- 
lons or stoloniform branches. Stolon production in var. brevistolon results in 
colonies of closely interconnected plants (often densely matted), in contrast to 
the individuals of var. virginianum. The distinction between var. virginianum 
and var. brevistolon usually can be determined from herbarium collections, but 
early-flowering plants commonly produce only a single head from a basal ro- 
sette of leaves and observation of later growth is necessary to observe stolon 
production. Ina few instances, it is possible that inter and infra-popu- 
lation variation occur (e.g., in NCU and USCH collections from Darlington and 
Florence cos., S.C.), and lignescent stolons (becoming partially buried and 
rhizome-like) of var. brevistolon may be very similar to structures in var. 
virginianum that originate as basal offsets or rhizomes. Using the criteria dis- 
cussed here for their distinction, however, var. virginianum and var. brevistolon 
have closely contiguous, non-overlapping geographic ranges. 

Stolon length appears to vary in var. brevistolon, although the stolons are 
never like those of var. australe and intermediates between var. brevistolon and 


—_— 


818 BRIT.ORG/SIDA 19(4 


var. australe have not been observed. In earlier phases of the present study, how- 
ever, no distinction was made between var. brevistolon and var. australe (as 
recognized here), and all were annotated as var. australe, based on their pro- 
duction of distinct stolons (in contrast to var. virginianum). 

The erect stems of var. brevistolon tend to be shorter than in var. virginianum 
but are often otherwise indisti ishable. In both taxa, the erect stems become 
leafy, with 2-3 internodes. 

In var. virginianum, a flower bud is produced in each leaf axil, and two 
capitula on naked peduncles are produced at each node (from a pair of oppo- 
site leaves). On the prostrate to decumbent stoloniform branches of var. 
brevistolon, one of the axillary flower buds commonly is suppressed and a node 
produces only a single capitulum, which is held erect. This mechanism for bud 
suppression tends to be active even on the erect branches of var. brevistolon, 
and on a plant collected without complete lower parts, observation of a single 
axillary shoot per node on erect stems isa reliable indication of its identity. The 
converse is not true, because plants of var. brevistolon may also produce both 
axillary buds at a node. 


le. ae tes virginianum Var. australe enue ex coma) Ahles, J. Elisha 

itchell Sci. a OU 173. 1964. (Fig. 1 1).c gon) trale Alexander ex Small, 

Southeast. FL, 1415, 1509, 1933. Type: U.S.A. ‘ is ean JACKSON Co. dry woods, some- 

what calcareous, aa a mile E of Marianna, 16 Mar 1925, R.M. Harper s.n. (HOLOTYPE: NY, 
ISOTYPES: GH}, USI). 


Plants stoloniferous, mat-forming; stolons (leafy above-ground runners) with 
longest internodes 12-60 cm, apparently remaining herbaceous. Stems (flow- 
ering) 2-10 cm tall, leafless. Leaves: blades elliptic-ovate, gradually tapered to 
the petiole, villous-hirsute with spreading hairs 1-3 mm long. Capitula: pe- 
duncles 2-10 cm long; outer phyllaries 7-LO mm long, 2.5-4 mm wide. Chro- 
mosome number unknown. 

Flowering spring (mid Mar-Apr, -May). Pine-oak, longleaf pine, beech-oak- 
magnolia woods, ravine slopes, limestone outcrops, floodplains and terraces, 
sand or sandy loam; Ala., Fla., Ga., La., Miss. 

Variety australe, as recognized here, corresponds in geographic range al- 
most exactly to the “East Gulf Coastal Plain” phytogeographic pattern outlined 
by Sorrie and Weakley (2001). Plants of this taxon are recognized by their leaf- 
less [lowering stems and whip-like stolons with long internodes. They are so 
strikingly different in appearance from those of var. virginianum that treat- 
ment of var. australe as a separate species might be justified. This is further 
emphasized by the apparent allopatry (and corresponding genetic isolation) of 
var. australe. The existence of a broad region of morphologically and geographi- 
cally intermediate populations (var. brevistolon), however, makes it reasonable 
to view var. australe and var. virginianum as extremes of a three-segmented 
array of variation within a single species. 


NESOM, TAXONOMIC REVIEW OF CHRYSOGONUM 819 


Besides the features of stolons and flowering stems, var. australe is distin- 
guished from the other two varieties by a tendency for the leaf blades to be ba- 
sally attenuate, a tendency for stems and petioles to be more villous (with longer 
trichomes), anda tendency for the phyllaries to be narrower. Variability in these 
features of var. australe overlaps with that in the other two varieties. 


ACKNOWLEDGMENTS 


lam grateful to the NCU and TEX-LL staffs for help during recent work there, 
USCH for a loan of specimens, the staffs at GH, MO, OS, and US for help during 
Chrysogonum study there in past years, the staffs of GH, KY, LTU, NY,and TENN 
for sending information and photocopies of specimens. John Strother and Bob 
Kral provided helpful reviews of the manuscript. 


REFERENCES 


AHLES, H.E. 1968. Asteraceae. In: Radford, A.E., H.E. Ahles, and C.R. Bell. Manual of the vascu- 
lar flora of the Carolinas. University of North Carolina Press, Chapel Hill. Pp. 1009-1139. 

Brown, R.G. and M.L. Brown. 1984. Herbaceous plants of Maryland. The Book Center, Univ. 
of Maryland, College Park. 

Cuester, E.W., B.E. Worrorb, and R. Kral. 1997. Atlas of Tennessee vascular plants. Volume 2. 
Angiosperms: Dicots. Misc. Publ. 13, Center for Field Biology, Austin Peay State Univ., 
Clarksville, TN. 

Cronauist, A. 1980.Vascular flora of the southeastern United States, Vol.|. Asteraceae. Univ. 
of North Carolina Press, Chapel Hill. 

FisHer, R.T. 1988. The dicotyledoneae of Ohio. Part 3. Asteraceae. Ohio State Univ. Press, 
Columbus. 

Haavitt, A.M., T.R. Brabtey, and C.E. Stevens. 1981. Atlas of the Virginia flora. Part Il. Virginia 
Botanical Associates, Route 6, Box 130, Farmville. 

Jones, $.B.1968.Chromosome numbers in southeastern United States Compositae, ll. Bull 
Torrey Bot. Club 95:488-494, 

Kovanpa, M. 1978. Chromosome numbers of miscellaneous United States dicotyledons. 
Rhodora 80:43 1-440. 

MitcHett, R.S. and G.C. Tucker. 1997. Revised checklist of New York state plants. New York 
State Museum Bull.490. New York State Museum, Albany. 

Morton, J.K. 1981.Chromosome numbers in Compositae from Canada and the U.S.A. Bot. 
J. Linn. Soc. 82:35 7-368. 

Naturat History Museum. The John Clayton Herbarium. The Natural History Museum, Lon- 
don. http://www.nhm.ac.uk/botany/clayton/index.htm! Accessed 1 Sep 2001. 

Nesom, G.L. 1978. Myrmecochory in Chrysogonum virginianum L. (Compositae). Bot. Soc 
Amer. Abstr., Misc. Ser. Pub. 156:21. 

RHoaps, A.F. and W.M.Kiein Jk. 1993.The vascular flora of Pennsylvania: Annotated checklist 
and atlas. American Philosophical Society, Philadelphia, Pa. 

Smatt, J.K. 1933. Manual of the southeastern flora. Univ. of North Carolina Press, Chapel Hill. 


820 BRIT.ORG/SIDA 19(4) 


Soisric, O.T., D.W. KyHos, M. Powel, and PH. Raven. 1972. Chromosome numbers in Compositae 
Vill: Heliantheae. Amer. J. Bot. 59:869-878. 

Sorrie, B.A. and A.S.Weaktey. 2001.Coastal plain vascular plant endemics:Phytogeographic 
patterns. Castanea 66:50-82. 

STRAUSBAUGH, P.D. and E.L. Core. 1977. Flora of West Virginia. Part 4 (Rubiaceae through 
Compositae) (ed. 2). 

Stuessy, LF. 1977.Revision of Chrysogonum (Compositae, Heliantheae). Rhodora 79:190-202. 


TAXONOMY OF THE DICHOTOMA GROUP OF 
DICHANTHELIUM (POACEAE) 
Richard J.LeBlond 
North Carolina Natural Heritage Program 
Richlands, fae ee U.S.A. 


ABSTRACT 


bn; -] Py ae Cee: [aac Pete Er 


Eleven taxa associated with the Dichotoma group of 
species and three varieties. A summary of recent taxonomic treatment of the BroLip is srowided, one 
with a key to the species and varieties, plus synonymy, typification, discussion, distribution, and 
ecology. Six new combinations are made: Dichanthelium annulum (Ashe) RJ. LeBlond, D. 
dichotomum (L.) Gould var. nitidum (Lam.) RJ. LeBlond, D. dichotomum var. ramulosum (Torr) RJ. 
LeBlond, D. dichotomum var. roanokense (Ashe) RJ. LeBlond, D. lucidum (Ashe) RJ. LeBlond, and D. 
sphagnicola (Nash) RJ. LeBlond. 


RESUMEN 


Se reconocen once taxa avorlades con el Brune prenctoms de Dichanthelrum, que ineluyen ocho 
especies y tres junto 


con una clave para las sean eee eee i imias, tipificacion, discusion, een ion 

, ecologia. Se hacen seis combinaciones nuevas: eer paas eda annulum (Ashe) RJ. LeBlond, D. 
es aan (L.) Gould var, nitidum (Lam.) RJ. LeBlond, D. dichotomum var. ramulosum (Torr) RJ. 
LeBlond, D. dichotomum var. roanokense (Ashe) RJ. LeBlond, D. 1 ne (Ashe) RJ. LeBlond, y D. 
sphagnicola (Nash) RJ. LeBlond 


INTRODUCTION 


In their seminal treatment of North American Panicum, Hitchcock and Chase 
(1910) established Dichantheliumasa subgenus, which they further subdivided 
into 17 “minor groups. ... These names are not intended to be formal and should 
have no nomenclatural standing.” Dichotoma is one of these “minor groups,” 
though it should be noted that Dichotoma has been used to name a section that 
includes all North American species of Dichanthelium found north of Mexico 
(Hsu 1965: Crins 1991). Dichanthelium was elevated to generic rank by Gould 
(1974) based on morphological characters, an interpretation more recently sup- 
ported by molecular data (Giussani et al. 2000). 

though most current treatments do not recognize the Hitchcock and 
Chase groups, the implied relationships are reflected in key groupings, infraspe- 
cific alignments, and synonymy. Hitchcock and Chase recognized 14 taxa in their 
Dichotoma group, allat the rank of species: Panicum annulum Ashe, P. barbulatum 
Michx., P. boreale Nash, P. caerulescens Hack. ex Hitchc., P. clutei Nash, P. 
dichotomum L.,P lucidum Ashe, P mattamuskeetense Ashe, P microcarpon Muh. 


SIDA 19(4): 821 - 837. 2001 


822 BRIT.ORG/SIDA 19(4) 


ex Elliott, P multirameum Scribn., P nitidum Lam., P. roanokense Ashe, P 
sphagnicola Nash, and P. yadkinense Ashe. Panicum nudicaule Vasey was added 
by synonymy in later treatments of Dichotoma taxa (Gould & Clark 1978; 
Clewell 1985; Hansen & Wunderlin 1988: Wunderlin 1998) but evidently is more 
closely related to tropical species or to members of Hitchcock and Chase’s 
Laxiflora group. Gould and Clark (1978) treated Dichanthelium boreale (Nash) 
Freckmann (=Panicum boreale) as a species separate from their concept of the 
Dichotoma group, regarding it as appearing “to be close to and possibly 
intergrading with D. dichotomum var. dichotomum and D. commutatum.” Gould 
and Clark also placed Panicum bicknellii Nash and P.calliphyllum Ashe, mem- 
bers of Hitchcock and Chase's Bicknelliana group, in synonymy with D, boreale. 

This treatment recognizes 10 taxa—seven species and three varieties—from 
the original Dichotoma group: Dichanthelium annulum (Ashe) RJ. LeBlond 
(comb. nov.), D. caerulescens (Hack. ex Hitchc.) Correll, D. dichotomum (L.) 
Gould, D. dichotomum var. nitidum (Lam.) RJ. LeBlond (comb. nov.), D. 
dichotomum var. ramulosum (Torr.) RJ. LeBlond (comb. nov.) (=Panicum 
microcarpon), D. dichotomum var. roanokense (Ashe) RJ. LeBlond (comb. nov), 
D. lucidum (Ashe) RJ. LeBlond (comb. nov.), D. mattamuskeetense (Ashe) 
Mohlenbr, D. sphagnicola (Nash) RJ. LeBlond (comb. nov,), and D. yadkinense 
(Ashe) Mohlenbr. Even though its closest relatives undoubtedly lie outside the 
Dichotoma group, D. nudicaule (Vasey) B.F Hansen & Wunderlin is also treated 
here because of its morphological similarity to Dichotoma taxa (e.g,, minute 
ciliate ligules, glabrous internodes), and because of recent treatments in syn- 
onymy. Dichanthelium boreale is in need of additional study and is not treated 
here. No attempt is made to elevate the status of Dichotoma above an informal 
grouping of taxa that appear to be very closely related (corroborated by syn- 
onymy assignments in recent treatments). 


RECENT TAXONOMIC TREATMENT 


Hitchcock and Chase’s Dichotoma group species concepts were largely main- 
tained in regional manuals until 1964. nat (1950) added P lucidum var. 
opacum Fernald, treated P.cluteias P matt tense var. clutei (Nash) Fernald, 
and treated P. barbulatumas P dichotomum var. barbulatum (Michx.) A.W. Wood. 
Gleason (1952) added P annulum var. glabrescens Gleason, treated P microcarpon 
as P. nitidum var. ramulosum Torr, and synonymized P lucidum var. opacum 
with P. lucidum, P. clutei with P. mattamuskeetense, P. barbulatum with P. 
dichotomum, and P. caerulescens with P roanokense. In summary, Fernald rec- 
ognized all 12 of the Dichotoma taxa attributed to the region covered by Gray’s 
Manual of Botany, plus P. lucidum var. opacum, and Gleason recognized nine of 
the same taxa (not including P. lucidum var. opacum), plus P annulum var. 
glabrescens. Among the Hitchcock and Chase Dichotoma taxa, only Panicum 
multirameum and P. sphagnicola were not known from the northeastern US. 


LEBLOND, TAAUNUBIT UF UP VILTANTAELIOM 823 


Radford et al. (1964) reduced the 11 Dichotoma taxa known to occur in the 
Carolinas toa single taxon, Panicum dichotomum (not including P sphagnicola, 
which was included in synonymy by Radford et al. 1968, but a specimen has 
not been found). This single-species concept was also used in several subse- 
quent treatments, notably Correll and Johnson (1970), Gould and Clark (1978), 
Hansen and Wunderlin (1988), and Zuloaga et al. (1993). Gould and Clark addi- 
tionally included all 11 species of Hitchcock and Chase's Ensifolia group within 
their concept of D. dichotomum, thereby reducing Hitchcock and Chase's con- 
cept of 25 species (not including boreale but including nudicaule) to a single 
species comprising five varietal taxa. The 13 Dichotoma taxa plus nudicaule 
were treated as a single taxon, D. dichotomum var. dichotomum, with the 11 
Ensifolia species reduced to four varieties: D. dichotomum var. ensifolium (Bald- 
win) Gould & C.A. Clark, var. tenue (Muhl.) Gould & C.A. Clark, var. glabrifolium 
(Nash) Gould & C.A. Clark, and var. breve (Hitchc. & Chase) Gould & C.A. Clark. 
Ensifolia taxa were separated from Dichotoma taxa at the species level in later 
treatments (Godfrey & Wooten 1979; Lelong 1984, 1986; Hansen & Wunderlin 
1988; Gleason & Cronquist 1991; Zuloaga et al. 1993; Wunderlin 1998). Clewell 
(1985), however, exceeded even Gould and Clark’s reduction by lumping all 
Dichotoma and Ensifolia taxa into a single taxon, Dichanthelium dichotomum 

Lelong (1984) argued that some taxa in Hitchcock and Chase’s Dichotoma 
group deserved a better fate than “being buried in synonymy,” and resurrected 
six former species as varieties of Panicum dichotomum: var. lucidum (Ashe) 
Lelong (including P. sphagnicola and P. lucidum var. opacum), var. 
mattamuskeetense (Ashe) Lelong (including P annulum and P. clutei), var. 
nitidum (Lam.) A.W. Wood, var. ramulosum (Torr.) Lelong, var. roanokense 
(Ashe) Lelong (including P caerulescens), and var. yadkinense (Ashe) Lelong. 
Neither Panicum boreale nor its synonyms, P. bicknellii and P.calliphyllum, are 
included in synonymy under Lelong’s concept of Dichotoma taxonomy. Lelong 
(1986) included P. barbulatum in synonymy with P. dichotomum var. 
dichotomum, and recognized Panicum nudicaule at the rank of species. The only 
other Hitchcock and Chase Dichotoma entity is Panicum multirameum, known 
from Mexico to Venezuela and Jamaica. It is treated in the present study as a 
synonym of D. dichotomum var. nitidum. 


DISTRIBUTION AND ECOLOGY 


All 10 Dichotoma group taxa treated here, plus Dichanthelium nudicaule, are 
centered in the eastern US., and five are endemic: Dichanthelium annulum, D. 
lucidum, D. mattamuskeetense, D. nudicaule, and D. sphagnicola. Distribution 
of species and populations in the US. is concentrated in the Atlantic and Gulf 
Coast states from New Jersey south to Florida and west to Texas, and extending 
as far inland as Illinois, Missouri, and Oklahoma. Dichanthelium dichotomum 
var. dichotomum is the only taxon reaching Canada (New Brunswick and 


824 BRIT.ORG/SIDA 19(4) 


Ontario). Three taxa range southward to the Bahamas and West Indies: D. 
caerulescens, D. dichotomum var. nitidum, and D. dichotomum var. roanokense 
(the last not in the Bahamas). Two reach Mexico: D. dichotomum var. nitidum 
and D. yadkinense, with D. dichotomum var. nitidum reaching Central America 
and South America (Venezuela). 

Within the US, three are endemic to the Atlantic and/or Gulf coastal plains: 
D. mattamuskeetense, D. nudicaule, and D. sphagnicola. Dichanthelium 
caerulescens and D. dichotomum var. roanokense are restricted to the coastal 
plain in the US. portion of their ranges. Dichanthelium dichotomum var. 
nitidumand D. lucidumare primarily found on the coastal plain in the US., but 
range inland: nitidum to the Appalachians, and lucidum to the Great Lakes. 
Dichanthelium dichotomum var. dichotomum, D. dichotomum var. ramulosum, 
and D. yadkinense are widespread from the Atlantic and Gulf coasts to the Mid- 
west, though D. yadkinense appears to be less frequent. Only Dichanthelium 
annulum is primarily an inland species, nearly restricted to the Appalachian 
Province with very few coastal plain occurrences. 

Literature sources for range data in the descriptions are Hitchcock and 
Chase (1910, 1915), Fernald (1950), Hitchcock (1951), Correll and Correll (1982), 
Lelong (1986), Zuloaga et al. (1993), Homoya et al. (1995), Angelo and Boufford 
(1998), and Schuyler (2000). All reported U.S. states of occurrence are listed, 
including District of Columbia (DC). 

The majority of Dichotoma group taxa are adapted to wet habitats, includ- 
ing swamps, marshes, and wet pine savannas and flatwoods. Dichanthelium 
dichotomum var. ramulosum and D. dichotomum var. nitidum, primarily plants 
of wet habitats, can occasionally be found in dry situations (with var. nitidum 
known only from dry habitats south of the U.S.). Dichanthelium dichotomum 
var. dichotomum is primarily a plant of wet-mesic to dry woodlands. 
Dichanthelium annulum isa plant of dry rocky or sandy open woods and cal- 
careous grasslands (barrens) and is thus distinguished by habitat and range as 
well as by morphology from other Dichotoma taxa. All of the Dichotoma taxa 
(plus D. nudicaule) primarily or exclusively found on the Atlantic and/or Gulf 
coastal plains are restricted to wetland habitats. 


SYSTEMATICS 


Dichanthelium isa frustratingly complex genus. Familiarity with living popula- 
tions is helpful in understanding taxonomic concepts, but lack of in situ familiar- 
ity with taxa outside of one’s area of work can lead to parochial treatments. Many 
Dichanthelium taxa are clinal, distinct in one part of their range and indistinct 
in another, adding to the difficulty of a range-wide approach. Intermediates 
can be found even among apparently more stable taxa. Despite these difficulties, 
most Dichotoma specimens, and especially living plants, are readily assignable 
toa taxon and most taxa display fidelity toa limited range of habitats. 


LEBLOND, TRAUNUIMIT UF UP VICAANTAELIOUN 825 


The following key and discussions are proposed as a foundation for a range- 
wide treatment of Dichotoma group taxa. They result from field and herbarium 
study beginning in 1985, and the accumulated efforts of other students of the 
genus. Field familiarity includes much of the Atlantic and Gulf coastal plains 
from Massachusetts to Mississippi, and Andros Island in the Bahamas. Since 
1990 Lhave been a fulltime field botanist in the coastal plain of North Carolina, 
where nine of the 10 Dichotoma taxa are found. The treatment is based prima- 
rily ona study of the extensive collections at US, NCU and DUKE, specimens in 
my own collection, and living populations. As much as possible, it relies on 
features of the spikelet to distinguish taxa, particularly spikelet and first glume 
length, pubescence, and fertile lemma length and width. Taxa with consistent 
diagnostic characters throughout their range are treated as full species, even if 
sometimes appearing intermediate toward other taxa in some features (e.g, D. 
mattamuskeetense, D. sphagnicola). 


KEY TO DICHANTHELIUM DICHOTOMA GROUP TAXA AND D. NUDICAULE 
1. Nodes, at least lower, bearded. 
Culm internodes (at least middle and ia ane pecunde paisa to moder- 
ately spreading short-hairy, sometim blades vel 
vety-pubescent on both surfaces, often dence SO; spikelee (1.5-)1.8-2.1 mm 


: 


long; dale ol ay Ieee or Bae soil and barrens 1.D.annulum 
. Culm t least middle and upper cauline blades glabrous (if 
pubescent, then ankles 5 2.0-2 rie m long in D. mattamuskeetense, or fertile 


lemma densely papillose in D. lucidum); spikelets 1.4-2.8 mm long; plants mostly 
of wet soils and mesic to dry woodlands. 
3. area glabrous. 
4, Spikelets 1.8-2.3 mm long;first glume 0.6-1.1 mm long; fertile ae 0.8- 
1.0 mm wide; widest vernal blades 3—8(—10) mm wide D. dichotomum 
pie eal 
4. Spikelets 1.4-1.9 long;first glume 0.3-0.6(-0.7) mm long; fertile lemma 
~0.8 mm wide; widest vernal blades 7-15 mm wide 5.D.dichotomum 


var.ramulosum 
3. Spikelets pubescent (rarely in D. dichotomum var. ramulosum) 
5. Spikelets (2.0-)2.2-2.8 mm long;first glume 0.5-1.3 mm long; fertile lemma 
8-2.3 mm long;lowest vernal cauline blades pul at least abaxially 
8.D. mattamuskeetense 
5. Spikelets 1.4-2.2 mm long; first glume 0.3-0.9 mm long;fertile lemma 
1.7 mm long; lowest vernal cauline blades glabrous. 
6. Spikelets 1.7-2.2 mm long; first glume 0.6-0.9 mm long; fertile lemma 
7-1.0 mm wide 4.D.dichotomum var. nitidum 
6. Spikelets 1.4-1.9 mm long; first glume 0.3-0.6(-0.8) mm long; fertile 
lemma 0.6-0.8 mm wide 5.D.dichotomum var. ramulosum 
. Nodes beardless, though sometimes short-pubescent. 
Cauline leaves mostly basally disposed, strongly ascending, much larger than 
the 2-3 remote middle and upper cauline leaves of fertile culms; spikelets 2.4— 
2.9 mm long; culms branch from basal and lower nodes, but are not known to 
produce autumnal inflorescences 9.D.nudicaule 


oat 


A 


™ 


BRIT.ORG/SIDA 19(4 


7. Cauline leaves well-distributed along culm, more than three, gradually reduced 
upwards and often spreading; spikelets 1.4-2.6 mm long; culms branch either 


from all nodes, or from middle and/or upper nodes, producing autumnal inflo- 
rescences 


8. Widest vernal cauline blades 7-15 mm wide; upper sheaths often oe 
warty;spikelets 2.1-2.6 mm long,some or most acute to beaked 


glum 
and sterile lemma extending 0.3-0.5 mm beyond fertile lemma in at ‘en 
some spikelets 


co 


. Widest vernal cauline blades 3-10 mm wide; upper sheaths not See 
warty; spikelets 1.4-2.7 mm long, blunt to subacute, second glume and 
sterile lemma equal to or shorter than fertile lemma, or extending less than 
0.3 mm beyond it. 

9. Spikelets 1.4-1.8 mm long; first glume 0.3-0.8 mm long; fertile lemma 1.3- 
1.5 mm long, smooth; mature vernal sas ea short-exerted with 
ascending branches; fresh foliage bluish-glaucous 2.D. caerulescens 

9. Spikelets (1.5-)1.7-2.7 mm long, if as short as - 5 mm, then with densely 
papillose Bee set (D. lucidum); first oon 0.6-1.4 mm Ong: fertile 
lemma 1.5- m long,smooth ord 


Or MALTS | 


ell lal dl icles 


exerted a oe branches; fresh foliage not bluish-glaucous 
10. Culms weak, soon sprawling over other vegetation; spikelets athel pu- 
bescent or fertile lemma and palea minutely but densely papillose at 
20x 


1 Spikelets 2.2-2.7 mm long, pubescent;first glume 1.0-1.4 mm long; 
fertile lemma and palea smooth or with a few weak papillae at 20 


D. sphagnicola 
ae J- . 3mm tong, glabrous (rarely pubescent); first 
Im rtil 


11. Spikelets (1.5 
g : lemma and palea densely papillose 


at 20x 


10. Culms stiffer, erect to ascending; spikelets glabrous, fertile lemma with 
no or few papillae 


7.D. lucidum 


12. Vernal a pases stiffly erect; plants of wet pine savannas and 


open D. dichotomum var. roanokense 
12. a ae blades spreading to deflexed, flexuous; plants of wet- 
mesic to dry woods and thickets 3. D. dichotomum var. dichotomum 


1. Dichanthelium annulum (Ashe) R.J. LeBlond, comb. nov. Panicum annulum Ashe 
J. Elisha Mitchell Sci. Soc. 15:58. 1898. Panicum bogueanum Ashe, J. Elisha Mitchell Sci. Soc. 
16:85. 1900, nom. illegit. Type: U.S.A. DISTRICT OF COLUMBIA: Dry rocky woods, Woodley 
Park, 1882, Ward s.n. (LECTOTYPE: USk SYNTYPE: US). 
P ae ] Pan ] re hI ry | ay, ea \RAALI ] 


This poorly known species has been infrequently collected and is presumably 
rare over its range. It has been treated in synonymy with D. dichotomum var. 
dichotomum, specifically with what is here treated as D. mattamuskeetense 
because of plants from southeastern Massachusetts. Dichanthelium annulum 
is a plant of dry rocky or sandy soil of open woods and calcareous grasslands 
(barrens), while D. mattamuskeetense inhabits wet sandy or peaty acidic soils 
of wet pine savannas and meadows, and wet borders of pocosins and thickets. 
As indicated in the key, the middle and upper internodes and peduncle of D. 


1. D. yadkinense 


) 


LEBLOND, TRAUNUMT UF UP VILAANTACLIVUM 827 


annulumare sparsely to moderately spreading pubescent with hairs 0.1-0.3 mm 
long. All internodes of D. mattamuskeetense are glabrous. Additionally, the 
internodes of D.annulumare typically dull, while those of D. mattamuskeetense 
are shiny. Spikelets of D. annulum are (1.5-)1.8-2.1 mm long with first glumes 
0.6-0.9 mm long. In D. mattamuskeetense, spikelets are 2.0-2.8 mm long with 
first glumes 0.5-1.3 mm long. Fertile lemmas are (1.2-)1.4-18 mm long and 0.8- 
1.0 mm wide in D. annulum and 1.8-2.3 mm long and (0.8-)1.0-1.3 mm wide in 
D. mattamuskeetense. Plants from southeastern Massachusetts with all blades 
pubescent on both surfaces, glabrous internodes, spikelets 2.2-2.5mm long, and 
occurring on moist ground, were treated by Gleason (1952) as P. annulum var. 
glabrescens Gleason. These plants may be the source of Fernald’s (1950) decla- 
ration that P annulum is “perhaps better merged with [mattamuskeetensel.” 
Glabrous internodes, spikelet length, and habitat are a much better fit for D. 
mattamuskeetense, as is range. Based on these conditions and character states, 
Pannulum var. glabrescens is here synonymized with D. mattamuskeetense. 

Dichanthelium annulum is one of the more distinctive entities within the 
Dichotoma group. Its combination of short-hairy upper internodes and velvety- 
pubescent leaves (upper as well as lower) distinguish it from all other taxa in 
the Hitchcock and Chase Dichotoma group. I have not found a treatment that 
specifically describes the internode pubescence character, but Ashe (1898) 
described the culmsas “below pilose, above generally glabrous,” which is oppo- 
site of the condition expressed in the two types at US, which are cited in Ashe’s 
paper. Culm internodes and the panicle axis are often glandular in D.annulum, 
with resinous nodules frequently in the grooves between upper internode 
longitudinal ridges, and with pellucid spots often present on the peduncle and 
panicle axis (both types at US express these characters). While glutinous-warty 
spots are encountered on sheaths in the Dichotoma group (notably D. 
yadkinense and D. dichotomum var. nitidum), and occasionally pellucid spots 
in the panicle axis and branches of D. dichotomum var. nitidum, further study 
may show that these glandular expressions provide additional distinguishing 
characters. 

Distribution and habitat—Distribution of Dichanthelium annulum is pri- 
marily in the Appalachian Province with very few occurrences on the coastal 
plain, from New Jersey to Georgia, west to Mississippi, and in Tennessee, south- 
east Missouri, and southern Indiana (AL, DC, DE, GA, IN, MD, MO, MS, NC, NJ, 
PA, TN, VA). Recent collections have been made in Lee Co., VA, on July 22, 1996 
(J.C. Ludwig 2872 & G. Fleming, pers. herb. RJ. LeBlond), and in Harrison Co., 
IN, on May 19, 1991 (Homoya et al. 1995). A specimen at US (Hanes & Hanes s.n., 
3 Jul 1934) that may be the source of Michigan’s inclusion in the Hitchcock (195) 
range has sparsely bearded nodes, glal internodes, and leaves that are mem- 
branous, subcordate, and pubescent. This entity may be more closely related to 
D.commutatum (J.A. Schultes) Gould. A specimen at US (Neill 5021) that may 


828 BRIT.ORG/SIDA 19(4) 


be the source of Florida’s inclusion in the Hitchcock (1951) range has pubes- 
cent leaves and pubescent spikelets 1.8-2.0 mm long, but a glabrous and shiny 
culm. It is not typical and may be more closely related to D. dichotomum var 
nitidum. No other specimens of D. annulum have been seen from Michigan or 
Florida and these states are not here included in the distribution. Dichanthelium 
annulum is a plant of dry rocky or sandy soil of open woods and calcareous 
grasslands (barrens). 


2. Dichanthelium caerulescens (Hack. ex Hitchc.) Correll, J. Arnold Arbor, 60:154. 
ga k.ex Hitche., U.S. Natl. Herb. Contr, 12:219, 1909. Type: U.S.A 
“LORIDA. ‘Date Co:" ‘In glade among Spartina, etc.,” 3 Apr 1906, Hitchcock 706 (HOLOTYPE: 


This species has the smallest spikelets and fertile lemmas among the Dichotoma 
taxa with beardless nodes and non-papillose fertile lemmas. Combined with 
the usually narrow and short-exerted vernal panicles, bluish-glaucous fresh 
foliage, and usually erect vernal blades, these characters readily distinguish 
Dichanthelium caerulescens from other Dichotoma taxa. It is most similar in 
habit to D. dichotomum var. roanokense, which has spikelets 1.8-2.2 mm long 
(14-18 mm in D. caerulescens) and fertile lemmas 16-18 mm long (1.3-1.5 mm 
in D. caerulescens). The culms of D. dichotomum var. roanokense on average are 
5-10 dm tall while those for D. caerulescens are 3-7.5 dm tall and not as stout. 
Dichanthelium caerulescens also prefers less acidic soils than D. dichotomum 
var. rodnokense. 

Distribution and habitat.-The documented range of Dichanthelium 
caerulescens is discontinuous and restricted to near the coastal edge in New 
Jersey, Virginia, and North Carolina, and from both coasts of Florida through 
southern Alabama and Mississippi to Louisiana. It also occurs in the Bahamas, 
western Cuba, and Puerto Rico. Dichanthelium caerulescens inhabits marshes, 
swamps, wet pinelands, maritime grasslands, and damp sandy soil near the 
coast. It is most frequently found in habitats where an alkaline or calcareous 
influence is evident and appears to be one of the least collected members of the 
Dichotoma group. 


3. Dichanthelium dichotomum (L.) Gould var. dichotomum, Brittonia 26:59. 1974. 
Panicum dichotomum L., Sp. PL. 58. 1753. Chasea dichotoma (L.) Nieuwl, Amer. Midl. Natural- 
ist 2:64. LOL. Type: U.S.A. VIRGINIA: “Habitat in Virginia,” Clayton 458 (LECTOTYPE: BM; 
ISOLECTOTYPE: US), fragment). 

Panicum barbulatum Michx. 1803, Fl. Bor. Amer. 1:49. 1803. Panicum dichotomum var. barbulatum 
(Michx.) A.W. Wood, Class-Book Bot., ed. 3:786. Panicum pubescens Lam. var. barbulatum 
(Michx.) Britton, Cat. Pl. New Jersey 280. 1889. Panicum nitidum var. barbulatum (Michx.) 
Chapm., FL. South. U.S., ed. 3:586. 1897. Type: CANADA: “Hab. in Canada P. capillari affine. Ad 
ripas amnis: Rivierre a a Cartier dicti legi,” Michaux s.n. (LECTOTYPE: P-MICH 
ISOLECTOTYPE: US!, fragme 

Panicum angustifolium cae ex Torr, Cat. Pl. New York 91. 1818, non Elliott 1816. Type: 


LEBLOND, TAAVINVUINIT UE VP VIRTANTACLIUIVE 829 


unknown. “A vernal specimen in the Torrey Herbarium penciled ‘angustifolius (nitid. var.) 

but without data may be the type” (Hitchcock & Chase 1910). 

Panicumtremulum S$ Spreng., Neue Entd. 2:103. 1821. Type: U.S.A. without data, Muhlenberg Herb. 
s.n. (LECTOTYPE: B, destroyed; ISOLECTOTYPE: US, fragment). 

Panicum dichotomum L. var. divaricatum Vasey, U.S.D.A. Div. Agrost. Bull. 8:30. 1889. TYPE: U.S.A. 
MISSISSIPPL SCOTT Co.: Lake, Tracy 127 (LECTOTYPE: USI). 

Panicum dichotomum L. var. viride Vasey, U.S.D.A. Div. Agrost. Bull. 8:30. 1889. Panicum nitidum 
Lam. var. viride (Vasey) Britton, Trans. New York Acad. Sci. 9:14. 1889. Panicum ramulosum 
Michx. var. viride (Vasey) Porter, Bull. Torrey Bot. Club 20:194. 1893. TyPE: U.S.A. DISTRICT 
OF COLUMBIA: Woodley Park, 1881, Ward s.n. (LECTOTYPE: US). 

Panicum nitidum Lam. var. saan orum Button, — New York Acad. Sci. 9:14. 1889. TYPE: U.S.A: 
NEW JERSEY: Morris Co. J | |, Britton s.n. (HOLOTYPE: NY). 

Panicum oo L. var. commune S. Watson & J. M. Coulter in A. Gray, Manual, ed. 6. 633. 

E: unknow 

Panicum a oe & Chase, Rhodora 8:205. 1906. Type: U.S.A. DELAWARE: NEWCASTLE 
Co. “in sandy, rather dry woods, on the old Commons farm, between Centerville and Mt. Cub 
30 im 1906, Chase 3620 (HOLOTYPE: US). Placed in synonymy with Panicum barbulatum 
Michx. by Hitchcock and Chase (1910 


o 


ba,” 


Recognition of the several species and varieties in this treatment reduces but 
by no means eliminates the apparent genetic diversity in Dichanthelium 
dichotomum var. dichotomum. Robust plants with bearded nodes, leaves to 10 
mm wide, and top-heavy autumnal inflorescences are referable to Panicum 
barbulatum (P dichotomum var. barbulatum), but intermediates abound. Plants 
frequently are found with bearded nodes, but with the smaller leaves and culms 
typical of the glabrous node form. 

Distribution and habitat.—Dichanthelium dichotomum var. dichotomum 
ranges from southern Canada (N.B., Ont.) south to the Gulf Coast, occurring 
throughout the eastern U.S. (AL, AR, CT, DC, DE, FL, GA, IL, IN, KY, LA, MA, 
MD, ME, MI, MO, MS, NC, NH, NJ, NY, OH, OK, PA, RI, SC, TN, TX, VA, VT, WV). 
Gould (1980) included this in synonymy in his treatment of D. dichotomum 
var. dichotomum sensu lato in Mexico, but var. dichotomum sensu stricto needs 
confirmation. Dichanthelium dichotomum var. dichotomum is the most wide- 
spread and frequent taxon within the Dichotoma group in the U.S, occurring in 
all states from which other members of the group have been documented. It is 
found primarily in wet-mesic to dry woods, thickets, and woodland openings. 


4. Dichanthelium dichotomum (L.) Gould var. nitidum (Lam.) RJ. LeBlond, comb. 
nov. Panicum nitidum Lam.,, Tabl. Encycl. 1172. 1791. Panicum dichotomum L. var. nitidum 
(Lam.) A.W. Wood, Class-Book Bot., ed. 3:786. Dichanthelium nitidum (Lam.) Mohlenbr., 
Erigenia 6:26. 1985. TYPE: U.S.A. “E. CAROLINA” Fraser s.n. (HOLOTYPE: P-LAM; ISOTYPE: USI, frag- 
ment and photo). 


Panicum nodiflorum Lam., Encycl. 4:744. 1798. Panicum dichotomum var. nodiflorum (Lam.) 
iseb., Cat. Pl. Cuba 234. 1866. TYPE: U.S.A.“ CAROLINE.” Fraser s.n. (HOLOTYPE: P-LAM; ISOTYPE: 
ie fragment and to). 
Panicum multirameum A U.S.D.A. Div. Agrost. Circ. 19:2. 1900. TyPE: MEXICO: VERACRUZ: 
near Jalapa, 1889, Pringle 7882 ( LECTOTYPE: USI, ISOLECTOTYPE: MO). 


830 BRIT.ORG/SIDA 19(4) 


Panicum subbarbulatum Scribn. & Merr, U.S.D.A. Div. Agrost. Circ. 29:9. 1901. TyPE: U.S.A. Elliott 
Herb. s.n. (HOLOTYPE: CHARLE: ISOTYPE: US|. fragment). 

Panicum multirameum, recognized by Hitchcock and Chase (1910, 1915), is an 
entity from Mexico, Guatemala, and Jamaica. Hitchcock and Chase (1910) de- 
scribe it as “allied to P nitidum,’ and a handwritten note (by J.R. Swallen, per P. 
Peterson, pers.comm.) on the outside of the Mexico-CA Panicum nitidum folder 
at US reads: “There are no consistent characters to distinguish P multirameum 
from P. nitidum. Neither the habit of the autumnal phase nor the viscid-spot- 
ted sheaths are constant characters. The type of P multirameum is from dry 
hills, which would account for its aspect. 5/15/50.” I concur with this conclu- 
sion after review of the P. multirameum specimens at US, although an appar- 
ent difference in habitat may warrant further study. 

Distribution and habitat.—In the U.S., Dichanthelium dichotomum vat. 
nitidum is found in the coastal plain from Pennsylvania and New Jersey south 
to Florida and west to east Texas; also in southeastern Missouri (DE, FL,GA,LA, 
MO, MS, NC, NJ, PA, SC, TX, VA). There are a few scattered mountain and pied- 
mont records from Virginia to Georgia. It is also known from the Bahamas and 
West Indies, and from Mexico to Venezuela. Dichanthelium dichotomum var. 
nitidum occurs on moist sandy or peaty soil of wet pine savannas and pocosin 
ecotones, in wet meadows near the coast, and reportedly from swamps and 
marshes. The Panicum multirameum specimensare from dry hills and gravely 


banks. 


5. Dichanthelium dichotomum (L.) Gould var. ramulosum (Torr.) RJ. LeBlond, 
MD. NOV. Panicum nitidum Lam. var. ramulosum Torr. Fl. N. Middle United States 146. 
1824. Panicum dichotomum L. var. ramulosum (Torr) Lelong, Brittonia 36:265. 1984. TyPE: 

S.A. NEW JERSEY: near Quaker Bridge, Jun 1818, Torrey s.n. (HOLOTYPE: NY). 

Panicum microcarpon Muhl. ex Elliott, Sketch Bot. S. Carolina 1:127. 1816. Dichanthelium 
microcarpon (Muhl. ex Elliott) Mohlenbr, Erigenia 6:26. 1985. TYPE: U.S.A. GEORGIA: Bald- 
vin s.n. (LECTOTYPE: CHARLD. Two fragments at US! labeled as isolectotypes of P. microcarpon 
are assignable to P. polyanthes Schult. and are examples of the nomenclatural problem dis- 
cussed by Hitchcock and Chase (1910) for this name. 

The infrequent occurrences of pubescent spikelets or first glumes to 0.8 mm in 
length in Dichanthelium dichotomum var. ramulosum suggest intergradation 
with another taxon, possibly D. dichotomum var. nitidum. The width of the fertile 
lemma appears to be a more reliable character than the width of the spikelet as 
a whole (the latter used by Fernald 1950), especially in herbarium specimens. 

Distribution and habitat—Dichanthelium dichotomum var. ramulosum is 
found from Massachusetts west to Michigan and south to Florida and Texas 
(AL, AR, CT, DC, DE, FL, GA, IL, IN, KY, LA, MA, MD, MI, MO, MS, NC, NJ, NY, 
OH, OK, PA, RI, SC, TN, TX, VA, WV). Gould (1980) included this in synonymy 
(as Panicum microcarpon) in his treatment of D. dichotomum var. dichotomum 


LEBLOND t VP DINAANTACLIUN 831 


in Mexico, but its occurrence there needs confirmation. Dichanthelium 
dichotomum var. ramulosum inhabits floodplain forests, swamps, openings, and 
borders of streams and ponds, and is occasionally found in dry upland woods. 


6. Dichanthelium dichotomum (L.) Gould var. roanokense (Ashe) RJ. LeBlond, 
b. nov. Panicum roanokense Ashe, J. Elisha Mitchell 2 Soc. 15:44. 1898. Panicum 
dichotomum L. var. roanokense (Ashe) Lelong, Brittonia 36:265. TyPr: U.S.A. NORTH CARO- 
LINA: DARE Co. Roanoke Island, Jun 1898, Ashe s.n. (PARATYPE: a NCU)). The specimen at 
NCU is labeled “Panicum roanokense Type” in Ashe’s handwriting and was collected by him, 

but lacks a date and locale. 


Pani j i Ashe, J. Elisha Mitchell Sci. Soc. 16:85. 1900. TyPE: U.S.A. MISSISSIPPI: JACK- 
SON CO.: Petit Boetdand 8 May 1898, Tracy 4584 USOLECTOTYPE: US!). The lectotype, suspected 
to be at NCU (Hansen & Wunderlin 1988), could not be located. 


Dichanthelium dichotomum var. roanokense is quite distinctive in the field with 
its stiff and erect vernal blades, but I can find no other character to consistently 
separate it from D. dichotomum var. dichotomum. Other treatments (Hitchcock 
& Chase 1910; Small 1933; Hitchcock 1951; Fernald 1950; Gleason 1952) vari- 
ously have used node annulus length, grayish olive-green blade color, spikelet 
turgidity, glume nerves, and glume basal coloring to separate var. roanokense 
from other D. dichotomum taxa, especially var. dichotomum. Fernald described 
the node annulus (the disk-like cartilaginous structure separating the intern- 
odes) as two-thirds or more as long as wide in var. roanokense, compared with 
“rarely” one-third as long as wide in var. dichotomum. Plants otherwise match- 
ing var. roanokense but with an annulus less than one-third as long as wide are 
common. Plants matching var. roanokense with olive-green blades are frequent, 
but plants with green and purplish-tinged blades are also encountered. Spike- 
lets tend to be more turgid than in var. dichotomum, and glume and sterile 
lemma nerves tend to be more pronounced, but there is too much variability in 
both taxa for this to be of use. Purplish-based glumes are frequent, but also oc- 
cur in var. dichotomum. Dichanthelium dichotomum var. roanokense is perhaps 
transitional between var. D. dichotomum var. dichotomum and D. caerulescens. 

Distribution and habitat.-The primary range for Dichanthelium 
dichotomum var. roanokense in the US. is near the coast from Delaware and 
Virginia south to Florida and west to Texas (AL, DE, FL,GA, LA, MS, NC, SC, TX, 
VA). It also occurs in Jamaica. Gould (1980) included this in synonymy in his 
treatment of D. dichotomum var. dichotomum in Mexico, but its occurrence there 
needs confirmation. This taxon is found in wet pine savannas, swamp open- 
ings, and wet peaty meadows. 


7. Dichanthelium lucidum (Ashe) RJ. LeBlond, comb. nov. Panicum lucidum Ashe, J. 

Elisha Mitchell Sci. Soc. 15:47. 1898. Panicum Seale L. var. lucidum (Ashe) Lelong, 

Brittonia 36:265. 1984. TyPE: U.S.A. NORTH CAROLINA: DARE Co. swamps bordering Lake 
Mattamuskeet, Jun 1898, Ashe s.n. (LECTOTYPE: a 


832 BRIT.ORG/SIDA 19(4) 


Panicum lucidum Ashe var. opacum Fernald, Rhodora 39:386. 1937. Type: U.S.A. VIRGINIA: PRINCE 
GEORGE Co. boggy depression one of Gary Church, 25 Aug 1936, Fernald and Long 6484 
CHO OTYPE: oe ISOTYPES: Me S!). 


p : NDi-l rig | 72 a § ors lat NI hac te 
yf 


Freckmann (LEC- 
TOTYPE. NGUY, 
The densely papillose fertile lemma and palea readily separate Dichanthelium 
lucidum from all other Dichotoma taxa. Fertile lemmas in other taxa in the 
group can havea few scattered papillae or longitudinally elongate reticulations, 
but are otherwise smooth. The papillae in D. lucidum cover the entire surface 
and are most easily seen at 20x or greater, with the pebbled texture discernable 
at 1Ox. Fernald (1950) described the fertile lemma as “obviously cellular-reticu- 
late,” but the papillate structure is evident when the edge of the fertile lemma 
is viewed at 20x. Dichanthelium lucidum is recognized in the field by its habit 
of sprawling over other vegetation (also see discussion under D. sphagnicola). 
Fernald (1937) described Panicum lucidum var. opacum from “an extensive 
boggy depression” in southeastern Virginia, distinguishing it from the typical 
variety by opaque strigose-pilose leaves (vs. lustrous and glabrous), and spike- 
lets L.5-1.8 mm long (vs. 1.9-2.1 mm). The isotype at US has leaves pilose to gla- 
brous, but the vestiture cannot be described as strigose. However, this character 
state may be present in situ, and may be lost over time in herbarium specimens 
The lustrous (satiny) leaf surface character is often absent (at least when dried) 
among specimens with glabrous leaves and larger spikelets. Gleason (1952) 
stated that “G)ntermediates exist with glabrous leaves and small spikelets, or 
with puberulent leaves and typical spikelets.” Glabrous plants with spikelets 
as short as 1.7-1.8 mm are frequent from the Carolinas to the Gulf Coast. 
Distribution and habitat—Dichanthelium lucidum is found primarily 
along the coast from Massachusetts south to Florida and west to Texas and 
Arkansas; it also occurs near the Great Lakes in Michigan and Indiana (AL, AR, 
CT, DC, DE, FL, GA, IN, LA, MA, MD, MI, MS, NC, NJ, NY, PA, SC, TN, TX, VA). 
This species inhabits wet meadows, sphagnous swamps, bogs, wet woods, and 
sphagnous streamheads (known as streamhead pocosins in the Carolina 
Sandhills and baygalls in the Gulf Coast). 


8. Dichantheli tt keet (Ashe) Mohlenbr., Erigenia 6:26. 1985. Pani- 
cum mattamuskeetense Ashe, Journ. Elisha Mitchell Sci. Soc. 15:45. 1898. Panicum dichotomum 

L. var. cle sealed she) Lelong, Brittonia 36:265. Type: U.S.A. NORTH CAROLINA: 

): Lake Mattamuskeet, 10 Jun-6 Jul 1898, Ashe s.n. (LECTOTYPE: US! ISOLECTOTYPE: NY). 


Panicum clutei Nash, Bull. Torrey Bot. Club 26:569, 1899, Panicum matiamuskeetense Ashe var. 
clutei (Nash) Fernald, Rhodora 39:386. 1937. Panicum dichotomum var. clutei (Nash) C.F Reed, 
Phytologia 67:452. Type: U.S.A. NEW JERSEY: BURLINGTON Co.: Tuckerton to Atsion, 3-6 Jul 
1899, Clute s.n. (HOLOTYPE: NY). 
Panicum flexuosum Muhl. ex Scribn. & Merr, U.S. Dept. Agric. Div. Agrost. Circ. 27:3. 1900, nom. 
illegit., non Retz. 1791. GsotTyPr: US!, fragment). 


LEBLOND OF DICHANTHELIUM 833 


inicumannulum Ashe var. glabrescens Gleason, Phytologia 4:21-22. 1952. TyPE: U.S.A. MASSA- 
CHUSETTS: BARNSTABLE Co.: Along roads leading from eee bogs east of Slough Road, 
Harwich, 18 Aug 1918, Fernald & Long 16044 (HOLOTYPE: NY!). At US! is a specimen labeled as an 
isotype of Pannulum var.glabrescens from Barnstable Co oe ‘Along cart road from 
Stanbeny Pegs east ra mioues Re ad, Bapsvich ” 9Jul 1918, Fernald 16043, but there is no in- 
t itself that Gleason designated this as a type. 


One of the most ‘einen of the Dichter taxa, D.mattamuskeetense is larger 
not only in spikelet and fertile lemma length, but tends to have the tallest and 
thickest culms, and the herbage tends to be a darker and more frequent purple- 
maroon. Plants with only the lowest nodes, sheaths, and blades pubescent, and 
with smaller spikelets, have been treated as Panicum clutei (Hitchcock & Chase 
1910; Hitchcock 1951) or P. mattamuskeetense var. clutei (Fernald 1950). Hitch- 
cock and Chase (1910) describe P clutei as “but doubtfully distinguished from 
P mattamuskeetense,” and there is considerable variability relative to plant pu- 
bescence and spikelet size throughout the range. Plants with glabrous middle 
and upper portions and with larger spikelets (2.4-2.8 mm long) are frequent in 
the southern portion of the range, and more pubescent plants with smaller 
spikelets (2.2-2.5 mm long) are known from southern New England. Also see 
discussion under Dichanthelium annulum. 

Distribution and habitat—Dichanthelium mattamuskeetense is found in 
the Atlantic coastal plain from southeastern Massachusetts south to northeast- 
ern South Carolina (DC, DE, MA, MD, NC, NJ, NY, RI, SC, VA). In the northern 
part of its range (Massachusetts to New Jersey), it is frequently found in the 
margins of cranberry bogsand in margins and openings of red maple-Atlantic 
white cedar swamps. Farther south it is found in damp to wet sandy or peaty 
soil of wet pine savannas and meadows, and wet borders of pocosin shrub 
swamps and thickets. 


9. Dichanthelium nudicaule (Vasey) B.- Hansen @ Wunderlin, Novon 11:367. 
Ol. Panicum nudicaule Vasey, U.S.D.A. Div. Bot. Bull. 8:31. 1889. TYPE: U.S.A. FLORIDA: 
swamps, Santa Rosa Co.: May 1886, Curtiss 3583 (LECTOTYPE: US! ISOLECTOTYPES: NY, TAES, 

US) 


Panicum nudicaule was “tentatively” placed in a single-taxon group, 
Nudicaulia, by Hitchcock and Chase (1910) based on “the narrow, enveloping 
base of the blades, and the nearly naked culms.” They allowed that the “techni- 
cal characters” suggest placement in the Dichotoma group, and D. nudicaule 
has been synonymized with D. dichotomum by recent authors (Gould & Clark 
1978; Clewell 1985; Hansen & Wunderlin 1988; Wunderlin 1998). The blades 
gradually narrow proximally and frequently the base is no wider than the sum- 
mit of the sheath, although envelopment of the culm by the basal portion of 
the blade is not a consistent character. Blade bases are frequently conduplicate 
(a continuation of the sheath folding), but are as likely to be free of the culm as 


834 BRIT.ORG/SIDA 19(4 


to enclose it. (In their 1910 description of P nudicaule, Hitchcock and Chase 
modified this Nudicaulia group character by describing the blade as “some- 
what enveloping the culm at base.”) The elongate lower culm blades are nu- 
merous and crowded and the middle and upper blades of fertile culms are few, 
distant, and markedly smaller. This leaf arrangement is due to short lower culm 
internodes and elongate upper culm internodes. Lower sheaths tend to be much 
shorter than the blades while middle and upper sheaths are often much longer 
than the blades. 

Other important morphological characterstics of Dichanthelium nudicaule 
suggest its closest relatives lie outside the Dichotoma group. It is peculiar among 
US. Dichanthelium taxa for not producing autumnal inflorescences; at least 
none has ever been seen or collected, not even during visits to 48 D. nudicaule 
sites from 1995-1999 by B.A. Sorrie and myself. However, D. nudicaule branches 
from the base and lower nodes, with the axillary branches persistent and the 
lower portions of previous-year culms at least sub-persistent. These character 
states suggest an affinity with Dichanthelium taxa found in the tropics and 
South America. Floral dimorphism in Dichanthelium taxa is often absent in 
tropical America, and “[mlain and axillary culms are persistent in Mesoameri- 
can and South American species” while “only the rosette usually persists” in 
North American species (Zuloaga et al. 1993). Basal branching and numerous 
elongate basal leaves also suggest an affinity with such taxa as D. laxiflorum 
and D. strigosum, both of which occur in North America and Mesoamerica. 

Distribution and habitat.—Dichanthelium nudicaule is known only from 
northwestern Florida, southern Alabama, and southern Mississippi, where it 
occurs in saturated areas of pitcher plant bogs (especially at the bases of seep- 
age slopes) and boggy ecotones between wet savannas and cypress streamheads. 


Representative specimens: ALABAMA. Baldwin Co.: Gateswood, | May 1903, S.M. Tracy 8432 (GH); 
Lillan Swamp, on tree stump along west bank of south fork of Caney Bayou, directly south of 
confluence with north fork, 18 Sep 1980, M. Lelong N532 (USA). Mobile Co.: pitcher plant bog along 
Rte 59,0.5 mi N of Fowl River, 24 Apr 1972, M. Lelong 6492 ((NCU, USA, VDB). FLORIDA. Bay Co. 
mucky sand of grass savanna between Sand Creek Rd and Mule Creek, ca.8.5 air mi ESE sie allawiay 
9 May 1995, L.C. Anderson 15473 (FSU). Santa Rosa Co.: Blackwater River ee May 1886, A.H. 
Curtiss 3583B (US, holotype). Walton Co.: Eglin Air Force ns New i Bogs, 17 May 1996, B.A. 
Sorrie 8812 (GA, NCU). MISSISSIPPI. Greene Co.: sohagnum bog ca mis “ State Line, 5 Jun 1980, 

K.L.Fordon 1903 and J. Burris (IBE). Harrison Co.: DeSoto cra ne pitcher plant bogs, Tuxachanie 
Trail,4 May 1974, K.E. Rogers 9537-A (NCU).Stone Co.: University of Mississippi Forest Lands, seepage 
bog at headwaters of Little Railroad Creek, 16 May 1997, B.A. Sorrie 9226 (GH, NCU) 


10. Dichanthelium sphagnicola (Nash) R.J. LeBlond, comb. nov. Panicum sphagnicola 
Nash, Bull. Torrey Bot. Club 22:422. 1895. Type: U.S.A. FLORIDA: COLUMBIA Co. sphagnum 
bog, Lake City, 29-31 Aug 1895, Nash 2500 (LECTOTYPE: NY; ISOLECTOTYPES: NY, US!) 

Soon reclining, Dichanthelium sphagnicola is similar in aspect to D. lucidum, 

but it has longer spikelets and first glumes that are typically pubescent and 

has smooth fertile lemmas. A few specimens from Florida have glabrous to 


LEBLOND, TAAUIIVIVIT VE VP VINTIANTTIOCLIUNG 835 


glabrate spikelets and weakly papillose fertile lemmas, suggesting intergrada- 
tion with D. lucidum. However, even these are readily separated from D. lucidum 
based on spikelet and glume lengths and the nearly smooth fertile lemmas. 
Hitchcock and Chase (1910) describe the autumnal habit of D. lucidumas “much 
more slender, more leafy, and bright green and shining” compared with D. 
sphagnicola. 

Distribution and habitat.—Dichanthelium sphagnicola is restricted to 
southeast Georgia and northern to south-central Florida. Although included 
in synonymy with Panicum dichotomum by Radford et al. (1968), no specimen 
has yet been seen from North or South Carolina. It is found along edges of cy- 
press swamps, in sphagnous bogs, and similar moist, shady places (Hitchcock 
& Chase 1910). 

Representative specimens: FLORIDA. Columbia Co.: Lake City, 29/31 Aug 1895, G.V. Nash 2500 (lec- 

totype, US). Levy Co.: fertile black flat woods near ponds, Bronson, 6 Sep 1898, R. Combs 838 (US). 

Volusia Co.: sandy bank along road about 10 mi SW of Daytona Beach, 24 Oct 1940, WA. Silveus 

6726 (US). GEORGIA. Chatham Co:: river swamp near Chatham Tower, 14 May 1940, DF. Fyles 5628 

(US). 

11. Dichanthelium a athe (Ashe) Mohlenbr, Erigenia 6:27. 1985. Panicum 
yadkinense Ashe, J. Elisha Mitchell Sci. Soc. 16:85. 1900 (nom. nov.). Panicum maculatum Ashe, 
J. Elisha Mitehev oat soe 15:44. 1898, nom. — not Aublet 1775. Panicum dichotomum L. 
var. yad} Lelong, Brittonia 36:26 E: U.S.A. NORTH CAROLINA: WAKE Co:: 
Raleiel Mey 1895, Ashe s.n. (LECTOTYPE: US). 


The combination of beardless nodes, large vernal blades (to 15 mm wide), and 
acute spikelets 2.1-2.6 mm long are definitive. When present, the large yellow- 
ish glandular “warts” on the (usually upper) sheaths are also helpful. However, 
they are often absent and occasionally occur on other Dichotoma group taxa 
(notably D. dichotomum var. nitidum). Within an inflorescence, some spikelets 
may be blunt apically, but many or most will be acute, with the second glume 
and sterile lemma surpassing the fertile lemma by as much as 0.5 mm. These 
characters, combined with no observed intergradation with other Dichotoma 
taxa, have led to recognition here at species rank. However, care must be taken 
not to confuse D. yadkinense with the superficially similar D. cryptanthum 
(Ashe) RJ. LeBlond ined., which has a similar appearance and acute spikelets 
of similar length, but with a membranous ligule (vs. hairy) and a scabrous pe- 
duncle and panicle axis (vs. smooth in D. yadkinense). 

Distribution and habitat—Dichanthelium yadkinense is reported from 
New Jersey to Michigan and south to Georgia and Texas (AL, DC, DE, GA, IL, IN, 
KY, LA, MD, MI, NC, NJ, OH, PA, SC, TN, TX, VA, WV). It also occurs in Mexico 
(Hitchcock & Chase 1915). This species appears to be scattered throughout most 
of its range, though there are 22 counties of occurrence in North Carolina. 
Dichanthelium yadkinense is found in floodplain forests, thickets, bottomlands, 
and swamps, often on alluvial deposits. 


836 BRIT.ORG/SIDA 19(4) 


ACKNOWLEDGMENTS 


Field work that substantially contributed to this study was sponsored prima- 
rily by the N.C. Natural Heritage Program, Division of Parks and Recreation, 
Department of Environment and Natural Resources. Bruce A. Sorrie introduced 
me to Dichanthelium at ponds on Cape Cod and has continued to be a learned 
counselor and great field companion. Alan S. Weakley is grudgingly thanked 
for entrusting me with the treatment of Dichanthelium in his draft of the Flora 
of the Carolinas and Virginia, an exercise that proves to be an unending learn- 
ing experience. | am indebted to Guy L. Nesom, Robert W. Freckmann, and 
Barney Lipscomb for reviewing this study in draft, and I thank the curators of 
US, NY, CHARL, and NCU for their assistance in the examination of types. 


REFERENCES 


Anaeto, R. and D.E. Bourrorp. 1998. Atlas of the flora of New England: Poaceae. Rhodora 
100:101-233. 

AsHe,W.W. 1898. The dichotomous group of Panicum in the eastern United States. J. Elisha 
Mitchell Sci. Soc. 15:22-62. 

Crewetl, A.F. 1985. Guide to the vascular plants of the Florida Panhandle. Florida State 
University Press, Tallahassee. 

Corrett, D.S. and H.B. Corrett. 1982. Flora of the Bahama Archipelago (including the Turks 
and Caicos Islands). J. Cramer, Vaduz, Germany. 

Corrett, D.S.and M.S. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research 
Foundation, Renner 

Crins,W.J.1991.The genera of Paniceae (Graminae: Panicoideae) in the southeastern United 
States. J. Arnold Arbor., Suppl. Ser. 1:171-312. 

FerNaco, M.L. 1937. Plants of the inner coastal plain of Virginia. Rhodora 39:32 1-366, 379- 
415, 433-459, 465-491, 

Fernato, M.L. 1950. Gray’s manual of botany. 8th ed. D. Van Nostrand, New York. 

Giussanl, L.M.,J.H. Cota-Sancuez, F. ZuLoaca, and E.A. Kettoac. 2000.A molecular phylogeny of 
the subfamily Panicoideae (Poaceae) using ndhF sequences. Amer. J. Bot. 87(6 
supp.):129. 

Gteason, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United 
States and adjacent Canada. 3 vols. New York Botanical Garden and Hafner Press, New 
York. 

GLeAson, H.A.and A. Cronauist. 1991.Manual of the vascular plants of northeastern United 
States and adjacent Canada. New York Botanical Garden and Hafner Press, New York. 

Goorrey, R.K.and J.W. Wooten. 1979. Aquatic and wetland plants of the southeastern United 
States: Monocotyledons. University of Georgia Press, Athens. 

Gouto, FW. 1974. Nomenclatural changes in Dichanthelium (Poaceae). Phytologia 39: 
268-271. 


LEBLOND 


VP VICAANTACLIUM 837 


Goup, FW. 1980. The Mexican species of Dichanthelium (Poaceae). Brittonia 32:353-364. 

Goutp, .W.and C.A.Ciark. 1978. Dichanthelium (Poaceae) in the United States and Canada. 
Missouri Bot. Gard. 65:1088-1132. 

Hansen, B.F., and R.P. Wunber.in. 1988. Synopsis of Dichanthelium (Poaceae) in Florida. Ann. 
Missouri Bot. Gard. 75:1637-1657. 

HitcHcock, A.S.1951.Manual of the grasses of the United States, second edition, revised by 
A. Chase. U.S.D.A. Misc. Pub. No. 200.Washington, DC. 

HitcHcock, A.S.,and A.CHase. 1910.The North American species of Panicum.Contr. U.S. Natl. 
Herb. 15:1-396. 

HitcHcock, A.S.and A. CHase. 1915. Tropical North American species of Panicum. Contr. U.S. 
Natl. Herb. 17:459-539. 

Homoya, M.A., D.B. Asrett, C.L. Heoce, and R.L. Heoce. 1995. Additions to the flora of southern 
Indiana, V and VI. Proc. Indiana Acad. Sci. 104:213-221. 

Hsu, C.-C. 1965. The classification of Panicum (Gramineae) and its allies, with special refer- 
ence to the characters of lodicule, style-base and lemma. Jour. Fac. Sci. Univ. Tokyo, Il, 
Vol. 9:43-150. 

Letonc, M.G. 1984. New combinations for Panicum subgenus Panicum and subgenus 
Dichanthelium (Poaceae) of the southeastern United States. Brittonia 36:262-273. 
Letonc, M.G. 1986.A taxonomic treatment of the genus Panicum (Poaceae) in Mississippi. 

Phytologia 61:251-269. 

Raproro, A.E., H.E. Antes, and C.R. Bett. 1964. Guide to the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill. 

Raprorb, A.E., H.E. AHLes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill. 

ScHuyter, A.E. 2000. Personal communication: report of specimens annotated as Panicum 
nitidum from Delaware (coll. by A. Chase) and Pennsylvania at PH. 

Smatt, J.K.1933.Manual of the southeastern flora. University of North Carolina Press, Chapel 
Hill. 

Wunpertn, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, 
Gainesville. 

ZuLOAGA, F.O., R.P. Ets, and O. Morrone. 1993. A revision of Panicum subg. Dichanthelium 
(Poaceae: Panicoideae: Paniceae) in Mesoamerica, the West Indies, and South America. 
Ann. Missouri Bot. Gard. 80:119-190. 


838 BRIT.ORG/SIDA 19(4) 


Book REVIEW 


Epwarb A. Corr. 2001. Muenscher’s Keys to Woody Plants: An Expanded Guide 
to Native and Cultivated Species. (ISBN 0-8014-8702-1, pbk.). Cornell Uni- 
versity Press, Sage House, 512 F. State Street, Ithaca, NY 14850). $22.95, 377 
bp. 6170" x9 1": 


ace wie for ieee of us who work in Le banized f1 I d to identify non-native 

y speci ever-present. I would have used this volume many li contexts over 
the | ast ae had it been available. The keys are simple but efficacious and well-polished and, 
clearly, most of the woody species to be encountered in temperate eastern North America are in- 


cluded. The a coverage s the! area east of the Mississippi River to the Atlantic Ocean, 
north tothe Arcti d htoi Mississippi, Alabama, and Georgia, and all of South 
Carolina except for five southeastern counties...” Even with this, all but five of the tree species on my 
tentative list of those native, naturalized, and auleivaced in Fort Worth Sexo NE in Cope’s treatment. 
C ope’s revision at 


x ion of the last edition of Muenscher's Keys (1950) treats 335 genera 
(174 native; 161 cultivated or - naturalized) = 1171 species (589 native; 582 cultivated, sera 198 
paralized) Some of the additions are “southern native species,” but “about 80 percent of [the] addi- 


ions are cultivated species. ... Of these, 198 species are naturalized from horticultural or ... inadvert- 
ent oe uctions of non-native species, ne h about half are only occasionally or rarely natural- 
ized.” “Additions of native woody plant species are mostly accounted for by a more precise and 
somewhat oe southern boundary of the geographical range covered, [a] tendency to include 
more of the locally endemic species, and the inclusion of woody grasses or herbs that appear to be 
nee such as Polygonum cuspidatum.” Examples of others included but on the border between 
‘woody’ and ‘herbaceous’ are CHEM Geos, pasos lata, C il ade ED PD periome =EP ener rid 
ete Rubus spp., Epigaea repens, C Clematis spp 

inca SPP., el, 


re 


Past the introductory material, the book is divided into the following sections: 
Keys to Genera—Inclusive [all native, naturalized, and cultivated genera that might be found 
outside of a otan ical perden or arboretum] 
Keys to Genera naturalized genera] 


ae to specie ponte arranged alphabetical i for easy access] 


1 by family, following the Cronquist 
r y i a oO q 
system| 


Glossary [technical terminology kept toa mininum], and Index. 


Each of the two ‘Keys to Genera’ is divided into seven smaller keys: 
Plants with needlelike, scalelike, or awl-shaped leaves 


Plants with ops or a ae e leaves 


Plants with oppo 


s P] leaves 


Plants with oo nate simple ae 
Ple te ith 
q 


tS eaves 
Plans a eaves, with the leaf scars opposite or whorled 
Plants lacking leaves, with the leaf scars alternate 
If th 5 i ion has several species with deciduous leaves, two ‘keys to species’ are of ten pro- 
aude summer hey and a winter key. “The summer keys are based primarily on leaves and fruits: 


the winter keys are based primarily on bud and twig characters, supplemented by those of bark, 
general habits, and fruits.” Floral features are not used in the keys. 

Highly recommended.—Guy L. Nesom, Botanical Research Institute of Texas, 509 Pecan Street, 
Fort Worth, TX 76102-4060, U.S.A. 


SIDA 19(4): 838. 2001 


THE GENUS HESPEROYUCCA (AGAVACEAE) IN THE 
WESTERN UNITED STATES AND MEXICO: NEW 
NOMENCLATURAL COMBINATIONS 


Karen H. Clary’ 
Department of Integrative Biology 
Plant Resources Center, Bio Labs 311 
University of Texas, Austin, TX 78712, U.S.A. 


ABSTRACT 
The g Hes, Ag consisting of three species, is alee as distinct On oes 
gee Ge ices hal eeaee d pollinator cl 


erations and recent DNA analyses sale ta whipplei is recognized and new Pembina for 
Ww e made: H. newberryi and H. peninsularis. A key that distinguishes Yucca from 
oe a the three species of ae. is included 


RESUMEN 


Se reconoce a Hesperoyucca (Agavaceae) como género distinto de Yucca en base a caracteres 
mortfoldgicos, fenoldgicos y del polinizador, a consideraciones biogeograficas y analisis recientes de 
ADN. Hesperoyucca consta - tres especies. Se reconoce Hesperoyucca whipplei y se proponen dos 
panies ve nuevas para las otras d peci ae newer: ace peninsularis. Se presenta una 


r Yucca de aaa at Pp Hesperoyucca 


Key Worps: Senta Yucca whipplei, Agavaceae, DNA 


Within Yucca L., as traditionally circumscribed, is a group of species, herein 
recognized as Hep Coyuce (Engelm.) Baker, that has morphological, pheno- 
logical, and pollina tor characteristics mar kedly different from all others. These 
characteristics include a loculicidal capsule, a capitate stigma, and glabrous, 
swollen filaments that are attached to the lower part of the tepals and that draw 
away from the ovary when the tepals open. These filaments bear bulbous an- 
thers with tufted pubescence that bend toward the stigma and dehisce later- 
ally (Fig. 1). Their pollen is produced ina glutinous mass. The sole pollinator of 
Hesperoyucca is Tegeticula maculata Riley, a species of yucca moth (Riley 1892; 
Trelease 1893, 1902; Powell & Mackey 1966; Davis 1967; Segraves & Pellmyr 2001). 
The other approximately 47 species of Yucca are visited by the remaining spe- 
cies of yucca moth (Pellmyr 1999). In Yucca, the pollen is not agglutinated, but 
produced as single grains, nor are the filaments tufted. Furthermore, the stig- 
mas are not capitate, but divided into three lobes that spread outward. The pu- 
bescent filaments are appressed to the ovary and angle outward and when the 
tepals open. 


'Current address: 4701 Canyonwood Drive, Austin, TX 78735-6602, U.S.A. kjclary@flash.net 
SIDA 19(4): 839 — 847. 2001 


840 BRIT.ORG/SIDA 19(4) 


Recent analyses of DNA markers, including the internal transcribed spacer 
region (ITS) of nuclear ribosomal DNA and chloroplast DNA (cpDNA) (Hanson 
1993; Bogler 1994; Bogler & Simpson 1995, 1996; Clary 1997) support the recog- 
nition of these unique species as members of a separate genus, Hesperoyucca 
(Engelm.) Baker. In the phylogenetic parsimony and maximum likelihood trees 
from DNA analyses, these species form a branch that is sister to Hesperaloé 
(Agavaceae) and not Yucca (Fig. 2). Hesperaloé Engelm. includes a group of five 
species found in the Sonoran and Chihuahuan Deserts allied taxonomically as 
close relatives to both Agave L.and Yucca (Correll & Johnston 1979; Gentry 1972; 
Starr 1997). 

The group in question, Hesperoyucca, corresponds to Engelmann’s (1871) 
Yucca group Hesperoyucca, which ranges from California and Arizona to Baja 
California Norte and Sonora in México. In this paper, Hesperoyucca is recog- 
nized at the genus level, requiring two new species combinations. A taxonomic 
treatment of the genus will be published in Flora of North America. 

While taxonomists working with Yucca have recognized the members of 
the Hesperoyucca group as distinctive from the other Yucca species, they have not 
agreed on infrageneric and infraspecific circumscription. The taxa within Hespero- 
yucca have been classified primarily by growth form (single or multiple rosettes), 
leat morphology and whether the plants are monocarpic or polycarpic. Most 
of the disagreement centers on the significance of morphological variation in 
Y. whipplei Torr. populations in Arizona and California (Trelease 1893, 1902; 
Haines 1941; McKelvey 1947; Webber 1953; Hochstatter 2000). 

Within Yucca whipplei, Trelease (1893) recognized two varieties, Haines 
(1941) and Munz (1968) each recognized five subspecies and Webber (1953) rec- 
ognized four varieties. Others argue that growth form isa highly variable char- 
acter and question the recognition of varieties or subspecies at all (McKelvey 
1947; McKinney & Hickman 1993). 

McKelvey (1947) described Yucca newberryi from Arizona and Y. 
peninsularis from the Vizcaino Desert region in Baja California Norte. Webber 
(1953:33) believed leaf, inflorescence and capsule features used by McKelvey to 
separate Y.newberryifrom Y. whipplei were weak and within the normal varia- 
tion of Y. whipplei. Hochstatter (2000) included Y. newberryi asa subspecies of 
Y. whipplei. 

Supported by distinctive morphological characteristics, unique species 
ranges and ITS DNA analyses (Clary 1997), three species of Hesperoyucca are 
here recognized: H. whipplei, H. peninsularis (McKelvey) Clary, and H. 
newberryi (McKelvey) Clary. A key to the species follows. 

The ITS DNA analysis of Yucca, Hesperoyucca, Agave and Hesperaloé(Clary 
1997) included single individuals from four Hesperoyucca populations: H. 
whipplei from Sierra Viejo, Sonora, México (Bogler & Simpson 1996), and from 
San Diego County, California; H. peninsularis from Catavina, Baja California 


CLARY, 


Norte, México; and H. newberryi from Mohave County, Arizona. The results 
show each of these samples to be genetically distinct from the others (Clary 
1997). The other taxa attributed to H. whipplei [Y. whipplei subsp. caespitosa 
(M.EJones) A.L.Haines, Y. whipplei subsp. percursa A.L.Haines, H. whipplei 
subsp. intermedia A.L.Haines, Y. whipplei subsp. typica A.L.Haines, and Y. 
whipplei subsp. parishii (M.E Jones) A.L.Haines] were not sampled. 

Results of the phylogenetic (parsimony and maximum likelihood) analy- 
sis show atree (Fig. 2) with the Hesperoyucca clade split into two branches. One 
branch contains H. newberryi, basal to the lineage, while the other branch con- 
tains H. peninsularis, which is sister toa branch that contains both H. whipplet 
populations. These data support specific status for H. peninsularis and H. 
newberryi, but not the two H. whipplei populations since they share unique 
mutations (Clary 1997). 

Recognition of infraspecific taxa within Hesperoyucca whipplei is beyond 
the scope of this paper. Although the taxonomic treatments of H. whipplei 
within the above-cited earlier works give distinctive morphological characters 
that separate taxa within H. whipplei, the treatments, with the exception of 
McKelvey’s (1947), lack sufficient records of specimens seen to evaluate the 
hypotheses about the proposed nature of variation within Y. whipplei. Further 
systematic study of informative morphological characters and DNA of all taxa 
of Hesperoyucca is warranted to elucidate its entire phylogeny and determine 
the genetic relationships that underlie the taxonomy of this group. 


KEY TO YUCCA AND THE THREE RECOGNIZED SPECIES OF HESPEROYUCCA 


1. Capsules septicidal, the seeds dispersing laterally through openings at the locule 
tips, or the fruit indehiscent, fleshy or spongy; seeds ultimately dispersed by ani- 
mals; stigmas 3-lobed, white, the lobes papillose on the inner surface; filaments 
usually shorter than the pistil, pubescent ania clavate su everitaally turned ata 


pronounced or negligibl hastate; 
inflorescence bracts on erect, persistent; leaf blades narrowed at ai with 
spoonlike base, widening above that point before tapering to the Yucca s.l. 


= 


. Capsules loculicidal, the seeds dispersing laterally through are or Ses 
false placental septa Boece initially wind dispersed; stigmas capitate, bright green, 
lly longer than pistil, finely papillate,and more 
th,stoutest at or near tl iddle, erect or spreading out- 
al from point of attachment at anthesis, bearing tufts of papillae at the apex; 
anthers cordate; inflorescence bracts becoming reflexed above the base, easily 
detached; leaf blades widest just above basal spoon and tapering to tip Hesperoyucca 
2. Mature leaf blades usually long and slender, to 0.7-2.0(-2.5) cm wide above the 
base, to 45-60(-100+) cm long, flexible, or if rigid, then distinctly thickened. 
3. Mature capsules with conspicuous placental wings; plants with single or 
multiple rosettes; western southern California and adjacent northern Baja 
California and Sonora H. whipplei 
3. Mature capsules with rudimentary placental wings; rosettes solitary; Mojave 
County, Arizona and vicinity H. newberryi 


| {| th 


842 BRIT.ORG/SIDA 19 


— 


4) 


2. Mature leaf-blades rather short and broad, to (2.5-)3.2-4.0 cm wide, straight and 
rigid, or falcate, tapering from above basal spoon to the tip; plants forming 
clustered rosettes; Vizcaino region, Baja California Norte, México H. peninsularis 


Hesperoyucca (Engelm.) Baker, Bull. Misc. Inform. Kew 1892(61):8. 1892. Yucca 
subgen. Hesperoyucca (Engelm.) Baker. Gard. Chron. ns. 6:196. 1876. Based on Yucca (without 
rank) Hesperoyucca Engelm. in S. Wats. et al. Botany [fortieth parallel! 497. 1871 as group 2: 
HESPERO-YUCCA. Yucca Sect. Hesperoyucca (Engelm.) in McKelvey, Yuccas Southw. U.S. 2: 
1947. TyPE: Yucca whipplei Torr. in lves. 


There is disagreement regarding authorship of the genus Hesperoyucca. Engel- 
mann (1871) divided Yucca into two major groups: Eu-yucca and HEsPERO-YUCCA, 
the former with three subgroups: Sarcocarpa, Clistocarpa and Chaenocarpa. His 
group Hesperoyucca contained only Yucca whipplei. Engelmann (1873) provided 
a similar summary classification, but within his Euyucca, changed the names 
to Sarcoyucca, Clistoyucca, Chaenoyucca (note change from “-carpa” to “- 
yucca”). In 1875 Engelmann retained four equal groups under Yucca: Sarcoyucca, 
Clistoyucca, Chaenoyucca and Hesperoyucca. But at no time did Engelmann 
indicate ranks for his groups within Yucca. Baker (1876) gave the rank subge- 
nus to Engelmann’s Hesperoyucca recognizing within it a single species, Yucca 
whipplei. Greuter et al. (1993) and Greenhouse and Strother (in press) accept 
this as legitimizing Engelmann’s Hesperoyucca as a subgenus. In 1892, Baker 
noted that Y. whipplei, “had better be kept as a genus distinct from Yucca, un- 
der Engelmann’s name Hesperoyucca” but he still listed the species as Yucca 
whippleii (sic.) Torrey. Greuter et al. (1993) accept this as having erected the 
genus Hesperoyucca. (Engelm.) Baker in 1892. Greenhouse and Strother (in press) 
following ICBN Art. 34.1 (Greuter et al. 2000), do not accept this as creating a 
new genusas Baker listed the species as Yucca whipplei and thus did not accept 
the new combination. Trelease (1893:208 pted Baker's (1892) suggestion of 
the genus rank for Hesperoyucca, formally recognizing Hesperoyucca at the same 
rank as, and separate from Yucca, and distinguishing Hesperoyucca from the 
“true Yuccas.” The combination Hesperoyucca whipplei appears in the list of 
illustrations, in the Explanation of Plates (Trelease 1893:215) and as the generic 
name of variety graminifolia (Trelease 1893:215, tt. 17 & 23). This is considered 
by Greenhouse and Strother (in press) to be the first legitimate use of the name 
of the genus Hesperoyucca. However, Baker's (1892) mere suggestion in print of 
generic rank for Hesperoyucca is accepted by Names in Current Use (Grueter et 
al. 1993) and by Flora North America (FNA) to be the first valid use of 
Hesperoyucca as a genus, not Trelease’s (1893) taxonomic description. 

Distribution.—U.S.A. California, Arizona. México. Baja California Norte, 
Sonora. 


Hesperoyucca whipplei (Torr) Baker ex Trel., Ann. Rep. Missouri Bot. Gard. 4:208. 
1893. (Fig. 1). Basionym: Yucca whipplei Torr. in J.C. Ives. Rep. Colorado R. 4 (Bot.):29. 1861. 
Type: U.S.A. CALIFORNIA. SAN DiFGO Co.: San Pasqual, A. Schott s.n. (LECTOTYPE: NY). The 


CLARY, 


OKSSSR 
se 
se 
a, 
is 
> 
Ley 
3 


ae 
aoe = 
FEE 

C) 
WSs 
F5g 
= 
Zoe 


= 
eS 
ee 


OF, Ue a, va ads 
< Ne 
9 es 


Fic. 1 We + o.. ru } mn TAL. A Growth habit, L ht gt tt £ J: £1 TE, i a 
voucher 148731 (MO))]; R cl | $l I rT re, TPF 1D 4 21 0? (TEX) cane photo 
by Constance & Morrison 2269, 1150192 (MO)]; C Capsule 


nBrenc: ee (MO)I; D vet ee ae papillae on n stigma surface from Nichols B- ie Bh 92 (TEX) and 


Nichols B-4-21-92 (TEX)]; F-G.l f blade (F) detail showi 


t 
A(C\ with an Ak ! 


Aorrison n 2269 1150192 (MO)] 


citation in Yucca is often ns as: eed upiuipelct ton in seen Rep. U.S. & Mex. 
2:222.1859. Torrey noted “i lled 


i ay be called Y. i aaa ee 
house and Strother (in see consi ‘ 


ubl Li ished 
as of 1859, but validly published in Ives’“ Report upon the Colorado River of fas ae (Torrey 
1861). Baker (1892) suggested that the species should be recognized within Hesperoyucca. 
Trelease (1893) formally recognized Hesperoyucca as a genus, and Hesperoyucca whippleiasa 


BRIT.ORG/SIDA 19(4) 


Agave striata 
Nuevo Leon, MEX 


8 Hesperaloé parviflora 
ValVerde Co., TX 


chilies eel 
uevo Leon, M 


Hesperaloé nocturna 
Sonora, MEX 


Hesperoyucca peninsularis 
81% Baja Calif. Norte, MEX 

Hesperoyucca oo 
San Diego Co. 


eae whipplei 
Sonora, MEX 


Hesperoyucca newberryi 
Mohave Co., AZ 


Yucca L. (45 species) 
U.S.A & MEXICO 


94% 


kamen strict consensus tree of the 10,777 parsimonious 467 step found by PAUP (Cl=0.89, HI=0.527, 
ges Step ch i bove branches, with bootstrap p tag 50 % written below (Clary 1997). 
hipplei (S , Mexico) are from 


LA lath UW pact ft) Uy | Page Sper J u 


Ino uW * Laie Lb hbinnlat/C n: fal; 
a 


Bogler (1994, 1996) ples of Hes; , Hesperoyucca | pero) PI 


fornia) 


é é é 


species. The combination listed above, can be shortened to Hesperoyucca whipplei (Torr) Trel. 
following ICBN Art. 46.4 (Greuter et al. 2000). Torrey’s (1859) paper cited a single specimen, 
“a yucca found by Mr. Schott on rocks near San Pasqual, southern California.” Torrey (1861) in 
the Ives Report cited three collections, “Mouth of Diamond River, 3 April, growing in tufts, on 
rocks,” another by Dr. Bigelow in Cajon Pass oa in C W ha expedition and by 
Mr. Schott near San Pasqual. The latter is | ectoty 


Hesperoyucca whipplei var. Ballas Trel., Ann. ns Missouri Bot. Gard. 4:215. 1893, based on: 
Yucca graminifolia A.\ od, Proc. Acad. Nat. Sci. Philadelphia 20:167. 1868, non Zucc. 1837. 
TYPE: U.S.A. CAL sae : OS ANGE LEs CO. Mountains 12 mi E of Los Angeles, 3 Mar 1866, 
A. Wood s.n. sensed oe oe notes a specimen at GH. a Coreen liause and prEorner (in pres 

Veranltciee 


note that Wood’s existen c 
Zucc. The name was eee by Trelease’s oe in 1893. 

Yucca whipplei var. parishii ME. ee Contr. W. Bot. 15:59. 1929. Yucca whipplei subsp. parishii 
(M.F. Jones) A.L. Haines, Madrono 6:44. 1941. TYPE: No pear were cited by Jones (1929). 
Jones (p. 59) described this taxon as “th form at low elevations on the Pacific slope, 
seldom if ever found on the desert side.” Haines (1941) gave the location of the type (California: 

San Bernardino Co. above Cajon Pass, M.E. Jones s.n.) but selected n spe. Jones collected 
several specimens from the Cajon Pass (POM, photocopy!), but none ee besnides g 
a type. With further study, a type may be designated for this taxon. 


CLARY, 845 


ve whipplei itosa M_E. Jones, Contr. W. Bot. 15:59. 1929. Yucca whipplei subsp. caespitosa 
M.E Jones) A.L. Haines: Madrono 6:43. 1941. Type: U.S.A. CALIFORNIA. SAN BERNARDINO CO.: 
cae pat in Neen eae 12 ae 1926, Jones s.n. (HOLOTYPE: POM!; ISOTYPE: CAS). 

nterme s, Madrono 6:43. 1941. Yucca whipplei var. intermedia 

(ALL. ines. M. ee Yuccas sore 34.1953. TyPE: U.S.A. CALIFORNIA. Los ANGELES 

Co: Malibu Lake, Santa Monica Mountains, l Jun 1940, Haines s.n.[HOLOTYPE: LA (photocopy!)l. 


Yucca whi bsp. percursa A.L.Haines. Madrono 6:43:1941. Yucca whipplei var. percursa (A.L. 
Gainey M Webber. “HUSeaS Southw. 35. 1953. TYPE: U.S.A. CALIFORNIA. SANTA BARBARA CO.: 
Cachumah Rafael Mountains, Haines s.n.(not found). A search at LA has found 


no authentic material. A neotype may be designated after further study. 
Distribution.—U.S.A. California: San Diego, Orange, Riverside, San Bernardino, 
Los Angeles, Ventura, Santa Barbara, Kern, San Luis Obispo, Tulare, Monterey, 
San Benito counties; México. Baja California Norte: Municipio Ensenada, 
Mexicali; Sonora. 


Hesperoyucca oe (McKelvey) Clary, comb. nov. BasionyM: Yucca newberryi 
McKelvey, Yuccas Southw. U.S. 2:49. 1947. Yucca whipplei subsp. newberryi (McKelvey) 
Hochstatter, nee (Netherlands) 79:39. 2000. TYPE: U.S.A. ARIZONA. MOHAVE Co.: Be- 
low rim of S wall of Colorado River, at New Water Point, 29 Apr 1934, McKelvey 4087 (HOLO- 
TYPE: A!) 


Distribution.—US.A. Arizona: Mohave County. 


a ae peninsularis (McKelvey) Clary, comb. nov. BasionyM: Yucca peninsularis 
Kelvey, Yuccas Southw. U.S. 2:52. 1947. TYPE: MEXICO. BAJA CALIFORNIA NORTE: Canyon 10 
mi - of El Rosario, 8 Feb 1935, LL. Wiggins 7559 (HOLOTYPE: DS!). 


Yucca whi | Epling & A.L.Haines, Brittonia 9:172. 1957. Type: Epling and Haines 
peas a type een on Baja California Norte, from 13 mi SE of Rosario in cirio- 
cardon community, 15 Apr A.L. Haines 5759 (not found). A search at LA has found no 


authentic material. A neotype will be designated after further study. 

Distribution.—México. Baja California Norte. 

Matuda and Pina-Lujan (1980) consider Yucca whipplei subsp. eremica Epling 
& ALHaines to be a synonym of Y. peninsularis. The original collections of 
each species are from the same area, 10 E of, and 13 miles SE of El Rosario. Com- 
parisons of habit and leaf morphology in the original descriptions (McKelvey 
1947; Epling & Haines 1957; Matuda & Pifia-Lujan 1980) and of both live and 
vouchered specimens at TEX indicate that both belong to the same species. 


ACKNOWLEDGMENTS 
I thank James Zarucchi (MO), Victoria Hollowell (MO), William Hess (MOR), 
Beryl Simpson, James Henrickson and Tom Wendt (TEX), John Strother (UC) 
for helpful comments on the manuscript. Photocopies of specimens were pro- 
vided by Lee Lenz (POM) and Barry Prigge (LA). I thank the curators of UC, 
POM, DS, GH for specimen loans. David Bogler (MO) provided photographs and 
was an advisor for the illustration (Fig. 1) drawn by Bee Gunn (MO) and Yevonn 


846 BRIT.ORG/SIDA 19(4) 


Wilson Ramsey (MO). Molecular and morphological research was supported 
by the National Science Foundation Doctoral Dissertation Improvement Award 
#9410882. 

REFERENCES 

Baker, J. 1876. New garden plants. Gard. Chron. n.s.6:196-197. 

Baker, J.G. 1892. CCXXIIl.—Agaves and arborescent Liliaceae on the Riviera. Bull. Misc. In- 
form. Kew 4(61):1-10. 

Boater, D. 1994. Taxonomy and phylogeny Dasylirion (Nolinaceae). Ph.D. Dissertation. The 
University of Texas, Austin. 

Boater, D.and B.B. Simpson. 1995.A chloroplast DNA study of the Agavaceae. Syst. Bot.20:191- 
205. 

Boater, D. and B.B. Simpson. 1996. Phylogeny of Agavaceae based on ITS rDNA sequence 
variation. Amer. J. Bot. 83:1225-1235. 

GreuTer, W., R.K. Brummit, E. Farr, N. Kian, PM. Kirk, and PC. Sivva. 1993. Names in current use 
for extant plant genera. Reg. Veg. 129:522. 

C1ary, K. 1997. Phylogeny, character evolution, and biogeography of Yucca L. (Agavaceae) 
as inferred from plant morphology and sequences of the internal transcribed spacer 
(ITS) region of the nuclear ribosomal DNA. Ph.D. Dissertation. The University of Texas, 
Austin. 

Corrett, D.S. and M.C. Jounston. 1979. Manual of the vascular plants of Texas, Texas Re- 
search Foundation. Renner 

Davis, D. 1967. A revision of the moths of the subfamily Prodoxinae (Lepidoptera: 
INncurvariidae). U.S. Nat. Hist. Mus. Bull. 255:1—170. 

ENGLEMANN, G. 1871. Yucca and Hesperaloé. |n: S. Watson's Botany of the Fortieth Parallel. 
Report of the United States Geological Exploration of the Fortieth Parallel. V. Suppl, 
496-497, Washington. 

ENGLEMANN, G. 1873. Notes on the genus Yucca. Trans. St. Louis Acad. Sci. 3:17-54. 

ENcLemaNn, G. 1875. Notes on the genus Yucca. No. 2. Trans. St. Louis Acad. Sci. 3:210-214. 

Eptina, C.and A.L. Haines. 1957. A subspecies of Yucca whipplei Torr. Brittonia 9:171-172. 

Gentry, H.S. 1972. The Agave Family in Sonora.Agric. Handbook No. 399. U.S.D.A.,Washing- 
ton, D.C. 

GreenHouse, J.A., and J.L. StrotHER. 2001. Hesperoyucca whipplei and Yucca whipplei 
(Agavaceae). Madrono (in press). 

Greuter, W., McNett, J., Barrie, F.R., Buroet, H.M., Demoutin, V, FILGUEIRAS, T.S., NICOLSON, D.H., SiLvA, 
P.C., Skoa, J.E., TREHANE, P., TURLAND, NJ, and HawkswortH DLL. (eds.). International code of 
botanical nomenclature (Saint Louis Code). Adopted by the Sixteenth International 
Botanical Congress St. Louis, Missouri, July-August 1999. Regnum Veg. 138:1-474. 

Haines, L.1941.Variation in Yucca whipplei. Madrono 6:33-45. 

Hanson, M. 1993. Dispersed unidirectional introgression from Yucca schidigera into Y. 
baccata (Agavaceae). Ph.D. Dissertation. Claremont Graduate School, California. 


CLARY, 847 


HOcHSTATTER, F. 2000. Het geslacht Yucca (Agavaceae). Succulenta (Netherlands) 79:32-44. 

Jones, M.E. 1929. Notes and new species of United States plants. Contr. W. Bot. 15:46-75. 

Matupa, E.and |. Pifa-Lujan. 1980.Las plantas Mexicanas del genero Yucca. Serie Fernando 
de Alva Ixtlilxochitl. Coleccion Miscelanea Estado de México. 

McKetvey, S. 1947. Yuccas of the southwestern United States, Part 2. The Arnold Arboretum 
of Harvard University, Jamaica Plain, MA. 

McKinney, K.and J. Hickman. 1993. Yucca. In: The Jepson manual: higher plants of California 
(J.C. Hickman, ed.). Univ. California Press, Berkeley. 

Munz, PA., 1968. A flora of California with supplement. Univ. California Press. Berkeley. 

Petimyr, O. 1999. Systematic revision of the yucca moths in the Tegeticula yuccasella com- 
plex (Lepidoptera: Prodoxidae) north of Mexico. Syst. Entomology 24:243-271. 

Powett, J. and R. Mackie. 1966. Biological interrelationships of moths and Yucca whipplei 
(Lepidoptera: Gelechiidae, Blastobasidae, Prodoxidae). Univ. Calif. Publ. Entomology 42. 
Univ. Calif. Press, Berkeley, CA. 

Ritey, C. 1892. The yucca moths and yucca pollination. Annual Rep. Missouri Bot. Gard. 
3:99-158. 

Secraves, K.and O. Petimyr. 2001. Phylogeography of the yucca moth Tegeticula maculata: 
the role of historical biogeography in reconciling high genetic structure with limited 
speciation. Molec. Ecol. 10:1247-1253. 

Starr, G. 1997. A revision of the genus Hesperaloe (Agavaceae). Madrono 44:293-294,. 

Torrey, J. 1859 [1858]. Yucca whipplei Torr.,|n:W.H. Emory, Report on the United States and 
Mexican boundary survey. 2:222. 

Torrey, J. 1861. In: J.C. lves, Report upon the Colorado River. Catalogue of the plants col- 
lected upon the expedition. 4:[1]-30. 

Trecease, W. 1893. Further studies of Yucca and their pollination. Annual Rep. Missouri Bot. 
Gard. 4:181-225. 

Treease, W. 1902. The Yuccae. Annual Rep. Missouri Bot. Gard. 13:27-133. 

Wesper, J. 1953.Yuccas of the Southw. Agric. Monogr. No. 17, U.S.D.A.,Washington, D.C 


848 BRIT.ORG/SIDA 19(4) 
Book REVIEW 
PauL MInNNis (editor). 2000. Ethnobotany, a Reader. (ISBN 0-8061-3180-2, pbk.). 
University of Oklahoma Press, 4100 28th Avenue N.W, Norman, OK 73069- 
8218, U.S.A. (Orders: wwwouedu/oupress, 405-325-2291, 405-364-5978 fax). 
$18.95, 384 pp, 22 b&w illustrations, 42 line drawings, 10 maps, 53 tables, 
Cle xo. 


Table of Contents: 


art One: Ethnoecology 
Ethnoecology: An Introduction. Catherine S. Fowler 
1) Factors Influencing Botanical Resource Perception Among the Huastec: Suggestions for Fu- 
ture SEMuunagnei ign yans B Alcorn 
2) California I g 
Anderson 
3) Papago (O'odham) Influences on Habitat and Biotic Diversity: Quitovac Oasis Fthnoecology. 
Gary P. Nabhan etal 
Part Two: Folk Classification 
Folk Classification: An Introduction. Cecil H. Brown 
4) The Pragmatics of Folk Classification. Brian Mor 
5) General Plant Categories in ee ee and L ‘looet, Two Interior Salish Languages of British 
Columbia. Nancy J. Turn 
6) Alternatives to ne Hierarchy: The Sahaptin Case. Eugene S. Hunnand David H. French 
Part Three: Foods and Medicines 
Foods and Medicines: An Introduction. Timothy Johns 
7) Candomble Ethnobotany: African Medicinal Plant Classification in Brazil. Robert A. Voeks 
8) Virtuous Herbs: Plants in Chumash Medicine. Jan Timbrook. 
9) On the Relative Contribution of Men and Women to Subsistence Among the Hunters-Gath- 
of the Columbian Plateau. Eugene S. Hunn 
10) Quelites- Ethnoecology of Edible Greens- Past, Present, and Future. Robert Bye 
11) Famine Foods of the North American Desert Borderland in Historical Context. Paul E. Minnis 


d Use of Redbud by the Southern Sierra Miwok. 


a 


— 


Part Four: Agriculture 
Agriculture: An Introduction. Richard I. Ford 
12) Devil’s Claw Domestication: Evidence from Southwestern Fiel - oe P Nabhanetal 
an Potat lrur Stephen B. Brush 


13) Eth a Biodiversity, and M 
14) Choice of fuel for Bagaco Still Helps Maintain eo Diversity in Traditional Portuguese 
ees System. eg F. Estabrook. 
List of Contact ee and Index 
This is an edition - at most, if not all, enpelany students will enjoy. Its broad scope gives a holis 
I so commonly perceived as trekking through the Amazonian jungles for 


tic flay t SO 
the cure to cancer. iewoues there isa relre hingly applic d approac h to most of these articles, dem- 
i ing human-plant interactions. I 


onstrating how ethnobotany can answer greate1 
highly recommend this book to anyone reterested in learning more about ethnobotany. It 

work well in undergraduate or graduate courses in ethnobiology.—Kevin D.Janni, Botanical 
Research Institute of Texas, Fort Worth, TX 76102-4060, U.S.A,.kjanni@britorg. 


would 


SIDA 19(4): 848. 2001 


NEW COMBINATIONS IN CHIONOLAENA 
(ASTERACEAE: GNAPHALIEAE) 


G.L.Nesom 
Botanical Research Institute of Texas 
Fort Worth, TX 76102-4060, U.S.A. 


ABSTRACT 


Five nomenclatural binati plete the transfer of Mexican and Central American species 
from pisceaaanian to ene Chionolaena costaricensis, Chionolaena cryptocephala, 
Chionolaena durangensis, Chionolaena macdonaldii, and Chionolaena salicifolia. In this view, 
Chionolaena is a genus s ae across northeastern and northern South America, Central America, 
and south-central to southwestern Mexico 


RESUMEN 
Cinco combinaciones nomenclaturales completan la transferencia de especies mexicanas y 
Centroamericanas de Gnaphaliothamnus a Chionolaena: Chionolaena costaricensis, Chionolaena 
cryptocephala, Chionolaena durangensis, Chionolaena macdonaldii, y Chionolaena salicifolia. Desde 
este punto de vista, Chionolaena es un género distribuido por el noreste y norte de Sur América, 
América Central, y del sur-centro al suroeste de Méxi 


Ten species of Mexico and Central America were treated within 
Gnaphaliothamnus Kirpiczn. (Nesom 1990a, 1990b, 1994), but a case has been 
made for regarding these species as members of Chionolaena DC. (Anderberg 
& Freire 1989; Anderberg 1991; Freire 1993), which otherwise includes species 
from northeastern and northern South America. Emphasizing similarities in 
habit and microcharacters among all of these species, as well as their apparent 
geographical continuity, the view that all arose from an immediate common 
ancestor is ble and all will be recognized as members of Chionolaena in 
an upcoming treatment of Mexican Gnaphalieae (Nesom in prep.). 

In this view, Chionolaena is a genus spread across South America, Central 
America, and south-central and southwestern Mexico. The Mexican and Cen- 
tral American species are characterized by a low, woody habit, revolute leaf 
margins, mostly heterogamous heads, phyllaries with white-opaque, spread- 
ing tips, reddish corollas, fertile achenes ellipsoid to obovoid 1.2-2 mm long with 
minute, somewhat elongated, duplex hairs, and central florets functionally 
staminate, with apically swollen pappus bristles and narrowly lanceolate style 
branches with collecting hairs along their whole length. They occur primarily 
in areas of high elevation. Distinctive vegetative glandularity and features of 
achenial vestiture of the Mexican and Central American species of Chionolaena, 
however, indicate that they probably constitute a phyletically coherent northern 


SIDA 19(4): 849 — 852. 2001 


850 BRIT.ORG/SIDA 19(4) 


segment of the genus, rather than being cladistically interspersed among the 
South American species, as postulated by Freire (1993) (see Nesom 1994 for fur- 
ther commentar y). 

Five of the Mexican and Central American species have valid names within 
the genus Chionolaena, as summarized here. 


Chionolaena aecidiocephala (Grierson) Anderb. & Freire, Notes Roy. Bot. Gard. 
Edinburgh 46:40. 1989. Anaphalis aecidiocephala Grierson, Notes Roy. Bot. Gard. 
Edinburgh 31:389. 1972. Gnaphaliothamnus aecidiocephala (Grierson) Nesom, Phytologia 
68:373. 1990. 

Chionolaena concinna (A. Gray) Anderb. & Freire, Ann. Missouri Bot. Gard. 
80:415. 1993. Gnaphalium concinnum A. Gray, Proc. Amer. Acad. Arts 15:34. 1879. 
Gnaphaliothamnus concinnus (A. Gray) Nesom, ele a a 1990. Chionolaena 
mexicana Freire [nom. nov], Ann. Missouri Bot. Garden 80:427. | 

Chionolaena eleagnoides Klatt, Leopoldina 23:88. 1887. Gnaphaliothamnus eleagnoides 
Klatt) Nesom, Phytologia 68:376. 1990 

Chionolaena lavandulifolia (Kunth) Benth. & Hook. f. ex B.D. Jackson, Index 
Kew. 1:516. 1893. Helichrysum lavandulifolium Kunth, Nov. Gen. & Sp. 4[foliok68. 1818 [as 
Elychrysum lavandulaefolium]. Gnaphaliothamnus lavandulifolius (Kunth) Nesom, 
Pp} tol 1 


7 8%, 


Ghisudlaeas sartorii Klatt, ieee 23:89. 1887. Gnaphaliothamnus sartorii (Klatt) 
Nesom, Phytologia 68:379. 199 


Another five species are brought into Chionolaena with combinations provided 

here. The first four were not accounted for in the revision of Chionolaena by 

Freire (1993); the taxonomic disposition of the fifth (C. salicifolia) is discussed. 

Chionolaena costaricensis Secu Nesom, comb. nov. Gnaphaliothamnus costaricensis 
Nesom, Phytologia 68:374. 1990. 

Chionolaena ee (Nesom) Nesom, comb. nov. Gnaphaliothamnus 
cryptocephalus Nesom, Phytologia 68:375. 1990. 

Chionolaena durangensis (Nesom) Nesom, comb. nov. Gnaphaliothamnus durangensis 
Nesom, Phytologia 69:1. 1990 


Chionolaena macdonaldii (Nesom) Nesom, comb. nov. Gnaphaliothamnus macdonalc 
Nesom, Phytologia 68:378. 1990 


. 


Chionolaena salicifolia (Bertol.) Nesom, comb. nov. Helichrysum salicifolium Bertol, 
Nov. Comm. Acad. Sci. Bonon. 4:433. 1840. Gnaphaliothamnus salicifolius (Bertol.) Nesom, 
Phytologia 68:378. 19¢ 


Gnaphalium rhodanthum Schultz-Bip. in Seemann, Bot. Voy. Herald, 310.1856. Gnaphaliothamnus 
rhodanthus (Schultz-Bip.) Kirpiczn., Trudy Bot. Inst. Akad. Nauk SSSR, Ser. 1, FL. Sist. Vyss. 
Rast. 9:33. 1950. [the type of Gnaphaliot ain 

Gnaphalium seemannii Schultz-Bip. in Seemann, Bot. Voy. Herald, 309. 1856. Chionolaena 
seemannii (Schultz-Bip.) Freire, Ann. Missouri Bot. Gard. 80:432. 1993 

Chionolaena corymbosa Hemsley, Diagn. Pl. Nov. 2:32: 1879 


NESOM, NEW COMBINATIONS IN CHIONOLAENA 851 


Taste 1.Pappus features of Mexican and Central American Chionolaena. 


pappus bristles: pappus bristles pappus bristles 
SPECIES basal coherence basal persistence apical cell shape 
C. aecidiocephala separate persistent clavate 
C.concinna separate to slightly persistent clavate 

connate 
C. costaricens!s separate caducous clavate 
C.cryptocephala separate caducous linear to weakly clavate 
C. durangensis slightly connate persistent clavate 
C. eleagnoides slightly connate persistent clavate 
C.lavandulifolia separate persistent linear to weakly clavate 
C macdonaldii (not seen) (not seen) clavate 
C. salicifolia slightly connate caducous linear to weakly clavate 
C. sartorii (not seen) (not seen) (not seen) 


Freire (1993) noted that the genus Gnaphaliothamnus, restricted in her con- 
cept to a single species, Gnaphaliothamnus salicifolius, is the sister group to 
Chionolaena, differing from all other species of Chionolaena in its “free pappus 
bristles” [vs. pappus bristles “fused at base into a ring”) “with linear apical cells” 
[vs. “clavate” apical cells]. In contrast, she incorporated the same single species 
into Chionolaena, using one of its synonyms for the nomenclatural combination 
(Chionolaena seemannii = Gnaphaliothamnus rhodanthus = Gnaphaliotham- 
{, 


icifolius = Chionolaena salicifolia (see Nesom 1994 for further commen- 


J 


— 


nus Sd 


tary on this synonymy). 

My own observations regarding the pappus features that distinguish 
Gnaphaliothamnus sensu stricto (in Freire’s view) further indicate that its sepa- 
ration at generic rank from similar Mexican and Central American species is 
artificial. Table 1 compares pappus features of these species—C. salicifolia does 
not stand apart. Chionolaena salicifolia was hypothesized to be most closely 
similar and related to C. eleagnoides by Nesom (1994). 


ACKNOWLEDGMENTS 


1 am grateful to John Strother and Debra Trock for their comments on the 
manuscript. 


REFERENCES 


AnoerserG, A.A. 1991. Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). 
Opera Bot. 104:1-195. 

ANpeRBERG, A.A. and S.E. Freire. 1989. Transfer of two species of Anaphalis to Chionolaena. 
Notes Roy. Bot. Gard. Edinb. 46:3 7-41. 

Freire, S.E. 1993. A revision of Chionolaena (Compositae, Gnaphalieae). Ann. Missouri Bot. 
Gard. 80:397-438. 


852 BRIT.ORG/SIDA 19(4) 


Nesom, G.L. 1990a. Taxonomy of Gnaphaliothamnus (Asteraceae: Inuleae). Phytologia 
68:366-381. 

Nesom, G.L. 1990b. An additional species of Gnaphaliothamnus (Astereae: Inuleae) and 
further evidence for the integrity of the genus. Phytologia 69:1-3. 

Nesom, G.L. 1994. Comments on Gnaphaliothamnus (Asteraceae: Inuleae). Phytologia 
76:185-191. 


SYSTEMATICS OF THE CAREX JAMESII COMPLEX 
(CYPERACEAE: SECT. PHYLLOSTACH YAE) 


Robert F.C. Naczi Bruce A. Ford 


Claude E. Phillips Herbarium Department of Botan 
Dept. of Agriculture & Natural Resources University of Manitoba 
Delaware State University Winnipeg, Manitoba 
Dover, DE 19901-2277, U.S.A. R3T 2N2, CANADA 
rnaczi@dsc.edu bford@cc.umanitoba.ca 
ABSTRACT 


nalyses of morphologic, chem and ecologic data indicate three species comprise the Carex 
Faeries -C jamesii, C junipero m,and C. timida, sp. nov. Carex jamesii has shoot bases lack- 


ing red-purple, relatively tall culms ane long staminate portions of spikes, and relatively long 


perigynium peas It is common and widespeeos in mesic deciduous forests in much of the eastern 
Ontario. Car rorum has red-purple shoot bases, short culms, short 


staminate oe of ee and short perigynium beaks. It is rare and has a disjunct distribution in 
mesic deciduous and deciduous-juniper forests and forest edges in the eastern U.S.A. and southeast- 
ern Ontario. oe timida has me purple shoot bases, tall culms, short staminate portions of spikes, 
and short perigynium beaks. It is rare and has a rausimnct dismapun ont in mesic a ciduous pie of 


the eastern U.S.A. Phylogenetic analysis 0 
and Gea is the sister species of C. timida. The correct name and authorship of ‘the section 
to which the C. jamesii complex belongs is cetermuiisd to be Carex section Phyllostachyae Tuck. ex 

Kuk. Synonymies, typifications, descriptions, and citat f representative specimens are provided 


for each species, “lone with an identificarion key. 


RESUMEN 


Los analisis de datos morfolégicos, geograficos, y ecolégicos indican que el complejo Carex jamesii 
acne tres especies: Cjomestt, GC yamiperonam, Bf C. timida, a nov. SATA IAMESTE tiene er) dases 


enuevos que no son rojo-purpura 

eae largas, y picos del utriculo relativamente largos. Es comun y esta dienes en los 
bosques deciduos mesofilos en gran parte del este de U.S.A. y en el suroeste de Ontario. Carex 
juniperorum tiene las bases de los renuevos cata aa cortos, parte estaminada de las 
espigas corta, y picos del utriculo cortos. Es raro y tiene una distribucion disyunta en los bosques 
deciduos mesofilos y enebrales, y bordes de bosques en a este de U.S.A. y sureste de Ontario. Carex 
timida tiene las bases de los renuevos rojo-purpura, ctilmenes altos, aa te estaminada de = pies 
corta, y picos del poi cortos. Es raro y tiene una eon disyu 
mesofilos del este de U.S.A. El analisis morfoldgicos ers que al ae es 
monofilético y que C. j a - especie peonana de . timida, El nombre y autoria 
] Carex seccion Phyllostachyae 


de lasecci6na 


Tuck. ex Ktik. Se ofrecen sinonimias, S ipificacione® en y citaciones de especimenes 
representativos de cada especie, junto con una clave de identificacion. 


INTRODUCTION 


Carex section Phyllostachyae Tuck. ex Kuk. is a morphologically distinctive 
group of sedges endemic to North American forests and forest openings. The 


SIDA 19(4): 853 — 884. 2001 


854 BRIT.ORG/SIDA 19(4) 


large and often leaf-like pistillate scales are the most striking feature of these 
plants. Additional traits common to the members of this group are densely cae- 
spitose habit; winged culms and peduncles, both of which are dilated at their 
apices; androgynous spikes, with lateral ones usually arising from the plant 
base; and beaked perigynia with entire orifices. Various workers (e.g. Crins 1990; 
Starr & Ford 1995; Naczi et al. 1998) have used the unusual morphologic fea- 
tures of Carex sect. Phyllostachyae to suggest the section is monophyletic, a 
hypothesis recently supported by molecular data (Starr et al. 1999). Recently, 
several authors have focused on various aspects of the systematics of Carex sect. 
Phyllostachyae, including phylogeny (Crins 1990; Starr et al. 1997, 1999), tax- 
onomy (Catling et al. 1993; Naczi & Ford 1998; Naczi et al. 1998), genetic diver- 
sity (Ford et al. 1998a, 1998b, 1998c), chromosome numbers and arrangements 
(Naczi 1999), and anatomy and micromorphology (Starr & Ford 2001). 

Ten species belong to Carex section Phyllostachyae: C. backii Boott; C. 
basiantha Steud; C.jamesii Schwein.; C juniperorum Catling, Reznicek, & Crins; 
C. latebracteata Waterf.; C. saximontana Mack: C. superata Naczi, Reznicek, & 
B.A. Ford; C. willdenowii Willd.; C. sp. nov. Saarela & B.A. Ford; and one species 
described as new in this paper. The greatest species diversity occurs in the south- 
eastern United States, where as many as five of these species are sympatric in 
portions of Alabama, Arkansas, and Tennessee. 

One of the clades we have identified within Carex section Phyllostachyae is 
the C. jamesii complex (Naczi & Ford 1998; Starr et al. 1999). We hypothesize that 
this clade contains three species, C. jamesii, C.juniperorum, and an undescribed 
species here named C. timida. In this paper, we test this hypothesis by ones 
the systematics of the C._jamesii complex. This is the first of two paper 
ing the systematics and genetic diversity among taxa in the C.jamesii commle 


= 


MATERIALS AND METHODS 


We studied the morphology, geography, and ecology of the C. jamesii complex 
in the field on numerous trips during 1986-2001. The field work included ob- 
servations at sites throughout most of the geographic range of this complex. 
These sites were in Ontario, Canada, and 13 states of the eastern United States: 
Alabama, Arkansas, Illinois, Kentucky, Maryland, Michigan, Mississippi, Mis- 
souri, Ohio, Oklahoma, Tennessee, Virginia, West Virginia. 

For this project, we studied over 1,950 specimens from the following 27 her- 
baria: BEREA, BLH, DHL, DOV, ERY, IBE, KNK, KY, MICH, MO, MT, MU, MUR, 
NA, NY, OS, PENN, PH, TENN, US, VDB, VPI, WIN, WIS, WKU, ctb, bas. Abbre- 
viations of herbaria are those of Holmgren et al. (1990) except for Charles T. 
Bryson herbarium (ctb) and Bruce A. Sorrie herbarium (bas). 


ch 


Morphology 
For the study of morphology, we selected 74 specimens as a representative sub- 


set to measure for univariate and multivariate statistical analyses. We chose 
mature, complete, and ample specimens that exhibited the full range of mor- 
phologic variation of the C_jamesii complex and that originated from through- 
out the geographic range of the complex. We measured 16 continuous and two 
discrete characters on each of these specimens (Table 1). Each specimen we 
measured is denoted by a superscript asterisk in the specimen citations below. 

To detect groups among the specimens and identify the cl hat best 
diagnose these groups, we used principal components analysis (PCA). Before 
conducting the analysis, we standardized all measurements so each variable 
would have a mean of Oanda standard deviation of 1. For the PCA, we included 
only continuous characters. To avoid weighting characters, we excluded characters 
that are probably genetically redundant. To detect such characters, we checked 
the Pearson correlation coefficient for all pairs of characters, and regarded values 
of the correlation coefficient 2 0.7 as indicating possibly genetically redundant 
characters. Exclusion of seven genetically redundant characters and two dis- 
crete characters resulted in the remaining nine characters being included in 
the PCA (Table 1). 

We assessed the degree to which the groups detected in the PCA differed 
among themselves, character by character, by analyzing each of the measured 
variables with one-way analysis of variance (ANOVA) with the Tukey Honestly 
Significantly Different test for multiple comparisons of means. Because the vari- 
ances of several variables are heterogeneous (as determined by the Bartlett chi- 
square test for homogeneity of group variances), we transformed these data with 
the common logarithm before conducting the ANOVA. We performed all statisti- 
calanalyses with SYSTAT version 5.1 (Wilkinson 1989) on an Apple® 

In order to apply the available names to the groups revealed amouen PCA 
and ANOVA, we studied type specimens. Isotypes of C. steudelii Kunth were 
unsuitable for measuring because of their immaturity. We ascertained the ap- 
plication of this name by comparison of its isotypes with the specimens of the 
PCA and ANOVA groups. We applied all other names by inclusion of type speci- 
mens in the PCA and ANOVA. 

Geography 

To determine the geographic range of each of the groups identified in the mor- 
phologic studies, we used herbarium specimen collection data to plot their dis- 
tributions on outline maps. Each point on these maps is based on at least one 
herbarium specimen. 


Ecology 

For each population encountered in the field, we noted vascular plant associ- 
ates. Closely associated vascular plant species are those that grew within 10 m 
of a plant of the C. jamesii complex at more than one-third of all sites investi- 
gated for this study. We also surveyed all localities for additional members of 


856 BRIT.ORG/SIDA 19(4) 


Taste 1. Morphologic characters, with their abbreviations, measured on herbarium specimens of 
the Carex jamesii con \plex. The 9 characters marked with asterisks are those included in the princi- 
pal components analysis. 


Continuous Characters 


1. Height of tallest culm, including terminal spike (CLMHT)* 

2. Height of shoot bearing tallest culm (SHTHT)* 

3. Width of widest leaf blade (WLFW) 

4. Length of longest (per specimen) staminate portion of terminal spike (MSPL) 

5. Length of peduncle of longest (per specimen) staminate portion of terminal spike (MPDCLL)* 
6. Length of proximalmost staminate sce oF teinpiia) spike (MSCL)* 


7. Length of longest (per speci perigynium of terminal spike 
(LFSCL)* 
8, Width of hyaline margin of scale subtending distalmost perigynium of terminal spike (MARGW)* 
9. Length of perigynium (PERIGL)* 
. Width of perigynium (PERIGW)* 
1. Length of perigynium a (PERIGBKL) 
. Length of achene (AC 
13. Width of achene peter 
14. Height of culm/height of shoot (RCLM) 
15. Length of proximalmost staminate scale/length of longest (per specimen) staminate portion 
of terminal spike (RSC)* 
16. Length of ae beak/length of perigynium (RBK) 
Discrete sale 
17. er of ieee and bracts on shoot bearing tallest culm (LFNO 
18. te of perigynia in terminal spike (PERIGNO) 


— 
— 


the complex. We consider species of the complex found within 30 m of each 
other to be examples of syntopy. 

Phylogeny 

We compared members of the C. jamesii complex with each other and with other 
species in section Phyllostachyae in an attempt to discover morphologic char- 
acters useful for phylogenetic analysis. We polarized character states using mem- 
bers of sect. Phyllostachyae that constitute the “wide-scaled clade.” Four species 
comprise this clade: C. backii, C. latebracteata, C.saximontana,and an undescribed 
species similar to C. saximontana (C. sp. nov. Saarela & B.A. Ford). Previous analy- 
ses of morphologic and molecular data indicate this wide-scaled clade is sister 
to the clade that includes the C. jamesii complex (Starr & Ford 2001). 

To conduct the phylogenetic analysis, we used PA UP* version 4.0b (Swofford 
2000) on an Apple® computer. For this analysis, all characters were weighted 
equally. The most parsimonious tree was computed using an exhaustive search. 
We assessed the robustness of clades using decay and bootstrap analyses (10,000 
replicates for the bootstrap analysis). 


RESULTS 


Morphology 

A scatter plot of the scores of principal components 1 and 2 depicts three dis- 
tinct groups (Fig. 1). Component | provides separation of C. jamesii from C. 
juniperorum, while C. timida is separated on component 2. No subgroupings 
within these three groups are evident. Together, the first two components ex- 
plain 71% of the variance within the data set, with component | accounting for 
44% and component 2 accounting for 27%. The third principal component ac- 
counts for only 9.6% of the total variance and affords no separation of groups. 
All of the variables employed in the PCA have relatively high loadings (abso- 
lute values > 0.5) on at least one of the first two components (Table 2). The vari- 
ables with the highest loadings on component | are CLMHT, LFSCL, and PERIGL, 
in descending order (abbreviations as in Table 1). On component 2, RSC, MSCL, 
and MARGW had the highest loadings. A plot of the two variables with the 
highest loadings for each component, CLMHT vs. RSC, separates the three 
groups (Fig. 2), but not as well as in the PCA plot. 

Like the PCA, the ANOVA supports the existence of three groups in the C. 
jamesii complex. Though the measurements of many characters possess con- 
siderable variability and the ranges of the measurements overlap for many char- 
acters, at least two of the three groups have significantly different means for all 
18 measured variables (Table 3). For eight of these characters, all three means 
are significantly different from each other. The variables that are the best for 
distinguishing the members of the C. jamesii complex, based on high F values, 
are RCLM, RSC, and MSCL, in descending order. 

Each of the three species identified in the PCA and ANOVA is distinguished 
by several characters. Specimens with relatively high values for MSPL, MPDCLL, 
LFSCL, PERIGL, PERIGBKL, and RBK are referable to Carex jamesii, since its 
holotype falls within this group (Fig. 1). Plants with low values for CLMHT, 
MSCLL, MARGW, PERIGBKL, RCLM, and high values for LFNO and PERIGNO 
are C. juniperorum. A measured isotype of C.juniperorum is a member of this 
second group. The third group, with low values for MSPL, high for MSCL, and 
high for RSC, is C. timida. The specimen designated as holotype of C. timida is 
clearly in this third group. 

Several qualitative characters also distinguish members of the C. jamesii 
complex. Plants of C. juniperorum and C. timida have red-purple tinging on 
their dark brown shoot bases, whereas plants of C. jamesii lack red-purple col- 
oration. The cells of the cataphylls of specimens of C.juniperorumand C.timida 
are bulging (Fig. 3). However, in C_jamesii, the cataphylls have cell surfaces that 
are collapsed or flush with their anticlinal walls, at least in dried specimens. In 
living plants, C.timida is distinctive in the color of its foliage. Its leaves are lighter 


858 BRIT.ORG/SIDA 19(4) 
) T T T T 
juniperorum 
4 rN 
A 
WN | _ i Bes = A fal 
< = 
G . Bes ee 20” 90 ® °o 
©) 
> oo a 000 = 
= jamesii 
rS ‘@) 
O 
a 
© Le : "= 7 
2 os roy 
Oo timida ms 
c " . 
= 
al 2 - *, = 
= H 
a 
3 | I | | 
“2 “1 0 1 2 3 
Principal Component 1 
Fig. 1. aay plot ot I f principal comp t 2 vs. principal comp Piiom PCA of the measurements of 
jt fj lex. Circl i f C jamesii | juniperorum, and 
C timida.T ymbols adj ay" i Tr ked 


1 
isk Ly Wee? er ee i L tud £ * lt H £+h 


I 
C. jamesii com- 
J 7 i 


alex (Ford & Naczi 2001). 


green than those of C. jamesii and C. juniperorum. Also in living plants, the 
culms and leaves of C. timida are quite lax and spreading. Consequently, when 
perigynia are mature (and relatively heavy), most of the spikes are concealed 
by the foliage. The living culms of C. jamesii are more rigid, erect, and evident 
than those of C. timida. In C.juniperorum, the culms are erect and concealed by 
the greatly overtopping foliage (illustrated in Catling et al. 1993). The wider 
leaf blades of C juniperorum have hyaline margins, whereas those of C. jamesii 
and C. timida are green. The proximalmost staminate scale in each spike has 
connate margins in C. juniperorum and C. timida, but the margins are free in C. 
jamesii (Fig 4). In addition, the staminate scales are papillate in C. timida, but 
nonpapillate in C. jamesii and C. juniperorum (Fig 5). In C.timida, some of the 


Taste 2.Loadings for the first two principal components from PCA of specimens of the Carex jamesii 
complex. Character abbreviations correspond to those in Table 1. 


Character Component 1 Component 2 
CLMHT 0.89 -0.21 
SHTHT 0.76 0.087 
MPDCLL 0.72 0.089 
SCL 0.23 0.90 
LFSCL 0.83 0.11 
MARGW 0.58 0.68 
PERIGL 0.82 0.16 
PERIGW 0.57 0.44 
0.20 -0.93 


papillae are elongate and acute. Often these elongate papillae occur in pairs 
and resemble arthropod cerci. 

Judging from the amount of scatter of points on the PCA plot (Fig. 1), the 
morphologic variability of C. jamesii and of C. timida is similar. Each of these 
species has substantially more variation than C. juniperorum. A specimen from 
the population of C. timida that clusters with C. juniperorum based on 
allozymes (Ford & Naczi 2001) is nested firmly within the C. timida group on 
the PCA plot (Fig. 1). 


Geography 

Carex jamesii ranges widely, from northern New York and southern Ontario 
west to southeasternmost Minnesota and south to western South Carolina, 
northern Alabama, northern Mississippi, southern Arkansas, and 
southeasternmost Oklahoma (Fig. 6). It is very common in much of this region, 
but is rather rare east of Ohio, Kentucky, Tennessee, and Alabama. 

Carex juniperorum occurs in three disjunct areas: southeastern Ontario, 
southwestern Virginia, and southern Ohio and adjacent northeastern Kentucky 
(Fig. 7). Except for the populations in southeastern Ontario, all populations of 
C. juniperorum are sympatric with C. jamesii. The southern Ohio and north- 
eastern Kentucky populations of C.juniperorum are sympatric with C. timida. 
Carex juniperorum is local and quite rare, with populations known from only 
Six counties. 

Carex timida occurs disjunctly in three areas: the Ozark Mountains of 
southern Missouri and northern Arkansas, the Ouachita Mountains of south- 
western Arkansas and southeastern Oklahoma, and southwestern Ohio and 
Indiana south to northern Alabama (Fig. 7). Throughout its range, C. timida is 
sympatric with C. jamesii. Carex timida is local and rare, with populations 
known from only 20 counties. 


860 BRIT.ORG/SIDA 19(4) 


50 T I T 
jamesii 
= 40 + 4 
> ‘@) 
6 O 
we O O 7 timida 
ZO w+ 0 o “  s . d 
se 
= o- 5 2 1, a 
— fe) 
Co WW! 20 $0 a oa 
= oO & O | iW 
con O_O m 
(@)) Oo . 
wane @ a 
= 10 - . = _ 
Awd a A 
A as ; 
A juniperorum 
0 | l l l l l 


Length of Stam. Scale/Length of 
Stam. Portion of Term. Spike (RSC) 


Fic. 2. Scatter plot of measurements of the variable having the highest loadi | t 1 (height of 

tallest culm) vs. on componsit 2 [ratio Yi length of proximalmost staminate scale/length of ae (per specimen) 
f f terminal spike] for the members oft carex jamesit f . Circles represent specimens of C 

jamesii i gl I juniperorum 1 I timida 

Ecology 


Carex jamesii plants inhabit mesic deciduous woodlands. They often grow along 
streams, usually in uplands, though they occasionally grow on floodplains. 
Substrates for C. jamesii are loams that appear to be calcareous and nutrient- 
rich, judging from the diverse associated flora, including known calciphiles. 
Some close associates of C. jamesii are Acer saccharum Marshall, C.albursina E. 
Sheld., C.amphibola Steud.,C. blanda Dewey, C. laxiculmis Schwein. var. copulata 
(L.H. Bailey) Fernald, C. oligocarpa Willd., C. rosea Willd., Cercis canadensis L., 
Diplazium pycnocarpon (Spreng.) M. Broun, Hydrastis canadensis L., Jeffersonia 
diphylla (L.) Pers. Lindera benzoin(L.) Blume, Osmorhiza longistylis (Torr) DC., 
Podophyllum peltatum L., Polystichum acrostichoides (Michx.) Schott, and 


NAC7TI AND EQRN 861 


Tasie 3. Means + 1 SD and ranges for morphologic characters measured for Carex jamesii complex. 
Character abbreviations correspond to those in Table 1. All measurements are in millimeters, ex- 
cept characters 14-16 (ratios) and characters 17-18 (counts). N = sample size. Within a row, means 
with different superscripts differ significantly (ANOVA, P < 0.03). 


C. jamesii C. juniperorum C. timida ANOVA 
Character (N = 37) (N=17) (N = 20) F 
1. CLMHT 2399 + 68.7 61° + 16 2024 + 73 120 
(142-413) (32-91) (90-338) 
2. SATHT 385% + 91.6 271° + 90.2 314° + 747 13.2 
(248-626) (145-449) (219-453) 
3. WLFW 2.7? + 0.40 3.3°+0.61 3.0 + 0.73 7.0 
(1.8-3.5) (2.2-4.2) (1.7-4.3) 
4, MSPL 8.2° + 2.2 6.0? 41.5 47°+ 0.83 33 
(4.9-13.5) (3.6-8.4) (3.4-6.2) 
5. MPDCLL 1.99 + 0.7 0.7>+04 115+ 0.4 30 
(0.8-3.8) (0.1-1.4) (0.4-1.9) 
6. MSCL 1.72+0.21 1.17#0.22 2° 033 140 
(1.1-2.1) (0.9-1.6) (1.9-3.3) 
7. LFSCL 59% 22 24° + 6.9 355+ 19 38 
(31-119) (16-36) (17-81) 
8. MARGW 0.5? + 0.1 0.1° + 0.06 0.6 + 0.09 100 
(0.3-0.7) (0.05-0.3) (0.4-0.7) 
9. PERIGL 6.1°+0.75 4.6” + 0.38 5.0° + 0.46 44 
(4.8-7.6) (3.9-5.4) (4.0-6.0) 
10. PERIGW 2.1°+0.16 1.9°+ 0.14 1.99+0.15 19 
(1.8-2.5) (1.7-2.1) (1.6-2.1) 
11. PERIGBKL 2.9°+ 0.51 1.6? + 0.24 2.05 + 0.27 77 
(1.9-3.9) (1.2-2.2) (1.4-2.5) 
12, ACHNL 24° + 0.24 2.254 0.14 2.3% +012 5.2 
(1.9-2.8) (2.0-2.5) (2.1-2.5) 
13. ACHNW 2.0% O15 1.9°+ 0.13 1.86+0.14 19 
8-24) (1.6-2.1) (1.6-2.1) 
14, RCLM 0.62? + 0.095 0,23" + 0.050 0:63? 40.12 190 
45-0.86) (0.15-0.32) (0.39-0.83) 
15. RSC 0.21? + 0.045 0.20* + 0.048 0.54° + 0.10 160 
(0.13-0.35) (0.13-0.34) (0.35-0.77) 
16. RBK 0.47? + 0.038 0.35" + 0.029 0.396 + 0.029 85 
(0.39-0.53) (0.30-0.43) (0.34-0.44) 
17. LFNO 4° + 0.6 BP +1 6S + J 70 
(3-6) (6-11) (4-8) 
18. PERIGNO 3° + 0.7 6° + 1 37405 90 
1-4) (4-8) (2-4) 


862 BRIT.ORG/SIDA 19(4) 


na. eo ee 


- 


Fic. 3. Scanning electron micrographs of surfaces of 
+ hall £ | £4} . . oe 1 A 


C. jamesii (Naczi 4039 & Thieret, WIN), B. C. juniperorum 
(Naczi 3808 et al., WIN), C.C. timida (Ford 98145 & Naczi, 
WIN). 


Sdanguinaria canadensis L. Carex jamesii rarely grows with C. timida (Table 4, 
and see below). 

Plants of Carex juniperorum grow in mesic woodlands and edges of wood- 
land openings, usually with a mixture of deciduous trees and junipers 
QYuniperus virginiana L.). Especially in Ontario, C. jun iperorum occurs in juni- 
per-dominated woodlands with few deciduous trees, particularly surrounding 
alvar openings (Catling et al. 1993). In the southern part of its range, C. 
juniperorum can be found in woodlands that are devoid of juniper. The optimal 
habitat, judging from the fact that specimens that are the most robust and pro- 
duce the most perigynia grow in such places, appears to be mature, relatively 
open forests dominated by oaks and hickories, with few and scattered junipers. 
Usually, plants of C.juniperorum occur far from streams, particularly since they 
often grow on gentle slopes near hilltops. Its habitats are moist, at least during 
the early part of the growing season, often through groundwater seepage. Its 


Fic. 4. Carex timida (A—C) and C. jamesii (D—E). Carex timida: A. Habit, with enl f portion of cataphyll, B. Termi- 

nal spike, C. Staminate portion of terminal spike, with proximalmost scale in aba | view (left) and in n adaxial view 

(right). ex ome D. Eroxivoal region of staminate portion of terminal spike, with proximalmost scale in adaxial 
E. Terminal spike 


864 BRIT.ORG/SIDA 19(4) 


Fic 5 | . L £ £. fel 

proximalmost staminate scales in terminal spikes of 
I ft i] ij pl C.jamesii (For 

98116 & Naczi, WIN), B.C. juni, (Naczi 5524 | 


WIN), CC. timida (Ford 98120 & Naczi, WIN). 


substrates are clays that are calcareous and circumneutral (Catling et al. 1993) 
and usually are not rocky. Vascular pl closely associated with C.juniperorum 
include Calystegia spithamea (L.) Pursh, Carex laxiflora Lam., C. oligocarpa, C. 
umbellata Willd., Cercis canadensis, Houstonia canadensis Roem. & Schult. 
Juniperus virginiana, Lithospermum canescens (Michx.) Lehm.,, Lobelia spicata 
Lam., Rhusaromatica Aiton,and Viburnum prunifolium L. We | observed 
C. juniperorum growing with any other member of the C. jamesii complex. 
Though rare and local, C. juniperorum is often common where it does occur, 
The usual habitat of Carex timida is in relatively open (sometimes closed) 
mesic deciduous and deciduous-juniper woodlands. Populations often occur 
high on slopes and on hilltops, far from streams. The substrates are loams and 
clay-loams that are apparently calcareous. The substrates are of ten rocky, with 
limestone at or near the surface at several sites. Among the closely associated 
vascular plant species are Acer saccharum, Aristolochia serpentaria L., Carex 


NACZI AND FORD, 865 


° 
e 
‘© Ree 
= fe, ° | “Len Se ,_500km ; 
‘ e 
e® of — 
*38° Py , : ‘on 
e' . ore 
e e Ps 
“esr ge 
caw . . - 
‘4 Carex jamesii 
ain 
ee B0°W 


Fic. 6. Geographic distribution of Carex jamesii. 


blanda, C. cephalophora Willd., Carya spp., Cercis canadensis, Juniperus 
virginiana, Quercus spp.,and Ulmus spp. Of the nine sites for C. timida we have 
studied in the field, four of them also hosted C.jamesti (Table 4). When C. timida 
and C. jamesii are syntopic, C. timida usually grows higher on slopes in slightly 
drier microsites than C. jamesii. 
Phylogeny 
We discovered 13 morphologic characters useful in the phylogenetic analysis 
of the C. jamesii complex (Table 5). Nine of the characters are reproductive and 
four are vegetative. Each of these characters has two states (Tables 5, 6). The 
analysis resulted in one most parsimonious tree that is 18 steps long (Fig. 8). 
The values for the consistency index and the retention index, excluding unin- 
formative characters, are 0.67 and 0.64, respectively. Of the 13 characters em- 
ployed in the phylogenetic analysis, 8 are nonhomoplasious, 2 are subject to 
parallelism between the C. jamesii complex and the outgroup (characters #4 
and 7), and 3 are subject to reversal (#8, 10, and 12). 

Five synapomorphies, three of which are nonhomoplasious, diagnose the 
C. jamesii complex and support its status as a monophyletic group (character 
numbers as in Table 5 in parentheses following the synapomorphy): relatively 


866 BRIT.ORG/SIDA 19(4) 


80°W 


© Carex juniperorum 
© Carex timida 
+ 31°N 


narrow staminate portions of the terminal spikes (6); relatively short 
proximalmost staminate scales (8), reversed in C. timida; truncate staminate 
scales (9); pistillate scales with hyaline margins relatively wide (12), reversed 
in C. juniperorum, and perigynia abruptly contracted to beaks (13). Two 
nonhomoplasious synapomorphies support the sister-group status of C. 
juniperorumand C.timida: shoot bases tinged with red-purple (1) and cataphyll 
cells bulging, Fig. 3 (3). Each of the three species in the C. jamesii complex is 
diagnosed by at least two autapomorphies. Carex jamesii has relatively long 
staminate portions of the spikes (5) and staminate scales with free margins, a 
reversal (10). Carex juniperorum is diagnosed by culms quite short relative to 
shoots (2), wider leaf blades with hyaline margins (4), a relatively high num- 
ber of perigynia in the terminal spikes (7), and pistillate scales with the hya- 
line margins very narrow, a reversal (12). Carex timida has relatively long 
proximalmost staminate scales, a reversal (8), and staminate scales bearing pa- 
pillae, some of which are elongate and acute (11). The clade of C. jamesii + C. 
juniperorum + C. timida has a decay value of 2 and has 78% bootstrap support, 
while the C. juniperorum + C. timida clade has a decay value of | and is sup- 
ported in 51% of the bootstrap replicates. 


\ACZI AND FORD, 867 


Taste 4. Examples of syntopy of Carex jamesii and C. timida. Numbers listed are Naczi’s collection 
numbers for voucher specimens. Full specimen citations are provided in the taxonomic treatment. 


Locality C. jamesii C. timida 

Arkansas: Howard Co. 1925 1918,1921 

Kentucky: Campbell Co. 5649 5650 

Kentucky: Monroe Co. 7236 7232 

Ohio: Montgomery Co. 7363 7357 

Taste 5. Characters, character states, and character state polarizations di | for use in the phy- 
logenetic analysis of the Carex jamesii complex. The plesiomorphic state is indicated by “(0)” and 


the apomorphic state by"(1).” Abbreviations for characters included in the statistical analyses are as 
in Table 1 


1. Shoot bases, color: lacking red-purple (0), tinged with red-purple (1). 
2. Culm, height of tallest per specimen/shoot bearing tallest culm, height (RCLM): = 0.39 (0), S 
0.32 (1). 

Cataphylls, appearance of periclinal walls of epidermal cells, in dried specimens (Fig. 3): sunken 

or flush with anticlinal walls (0), bulging (1). 

4. Leaves, color of margins of wider blades: green (0), hyaline (1). The apomorphic state is due to 
complete seer of the margins (Starr & Ford 2001). Carex latebracteata and C. saximontana 
have hyaline leaf blade ee but C. backii and C. sp. nov. lack them. Hyaline margins are 
scored as ade specimens in the likely sister groups to sect. Phy llostachyae, sects 
Filifoliae and Firmi ae ee et al. 1999), have green margins. 

. Staminate portion of terminal spikes, length of longest per specimen (MSPL): S 8 mm (0), = 6 

m (1). 

. Staminate portion of terminal spikes, width: = 0.8 mm (0), 0.8 mm (1). Carex latebracteata has 
the staminate portions of terminal spikes greater than 1 mm, but sometimes, C. backii, C 
saximontana, and C. sp. nov. have them as narrow as 0.5 mm. However, most specimens of C. 
backii,C. saximontana, and C. sp.nov. have the staminate portions at least 0.8 mm wide. Thus, the 
plesiomorphic state is scored as staminate portions = 0.8 mm wide. 

. Perigynia in terminal spike, number (PERIGNO): < 5 (0), = 5 (1). Carex latebracteata usually has 7 

or more perigynia in the terminal spike, which is likely a parallelism with the occurrence of this 

apomorphy in the C.jamesii complex since C. backii, C. saximontana, and C. sp. nov. usually have 

5 or fewer perigynia per spike. 

Proximalmost staminate scale in pale spikes, length (MSCL): = 2.0 mm (0), S 2.0 mm (1). 

Staminate scales, shape: ovate (0), truncate (1). 

. Proximalmost staminate scale in a spikes, degree of fusion of margins: margins free (0), 
margins connate in basal 30-80% (0). Carex sate acteata He etalminate scales free mar- 
ale but : ean C.saximontana, and C.sp.nov.h margin 

ae) ecause speci inthe fikelysi sister groups to sect. aia 
sects. Fill folie and Been iculmes (Starr et al. 1999), have scales with free ma 

. Staminate scales, appearance of periclinal ee of epidermal cells (Fig. 5): Sees ), papil- 
late, with some papillae elongate and acu 

. Pistillate scale margins, width of hyaline wet of scale subtending distalmost perigynium of 
terminal spike (MARGW): S 0.3 mm (0), = 0.3 mm (1). 

13. Perigynia,abruptness of taper from bodies to beaks: gradually tapered (0), 


Ww 


Sa) 


ion 


NI 


9 © 


oO 


= 


N 


ptly contracted (1). 


868 BRIT.ORG/SIDA 19(4) 


Taste 6. Character state assignments for members of the Carex jamesii complex. Characters and 
their states are as in Table 4. Carex backii, C. latebracteata, C. saximontana, and C.n.sp. constitute the 
outgroup. 


Character Number 


C. jamesil 
C juniperorum 
C. timida 


C. latebracteata 
C. saximontana 
C. sp. nov. 


Oooooao--oa 
aooooc 6.4 6 
oe a a ea oe a) 
o--00-0o0 
O72! O'S one 
ooaoao-=- = 
oOo-o0-0 
ao00o00 = -— 
Oo OO. = = 
—-—=3- O-+--=0o 
oO 
Sooo e aoe 
oO 


DISCUSSION 


The results of the PCA and ANOVA support the recognition of three distinct 
species in the Carex jamesii complex. The best diagnostic features are CLMHT, 
MSCL, RCLM, and RSC. Morphologically, the most divergent member of the C. 
jamesii complex is C. juniperorum. On both the PCA and bivariate plots, C. 
juniperorum is the most distinct of the three species (Figs. 1, 2). Of the 16 mea- 
sured characters with statistically significantly different means that are ex- 
tremes, 10 of them belong to C. juniperorum (Table 3). Measurements for 8 of 
these 10 characters represent low extremes. For measurements of only one char- 
acter (MSPL) is C. juniperorum intermediate between C. jamesii and C. timida. 
Carex juniperorum is the only species with very short culms (CLMHT), very 
short peduncles on the staminate portions of spikes (MPDCLL), very short 
staminate scales (MSCL), very short pistillate scales (LFSCL), very narrow hya- 
line margins on its distal pistillate scales (MARGW), very short perigynium 
beaks (PERIGBKL), culms greatly overtopped by the leaves (RCLM), very short 
perigynium beaks relative to the length of the perigynia (RBK), a high number 
of leaves and bracts on reproductive shoots (LFNO), and a high number of 
perigynia in the terminal spikes (PERIGNO). Carex juniperorum isalso the only 
species with hyaline margins on the wider leaves. 

Carex jamesii is distinctive mostly in its high values for measured features. 
Of the 16 measured characters with statistically different means that are ex- 
tremes, 10 of them belong to C. jamesii (Table 3). Measurements for 9 of these 10 
characters represent high extremes. For measurements of only two characters 
(MSCL, MARGW) is C. jamesii intermediate between C. juniperorum and C. 
timida. Carex jamesii is the only species with very tall shoots (SHTHT), very 
long staminate portions of terminal spikes (MSPL), very long peduncles on the 
staminate portions of spikes(MPDCLL), very long pistillate scales (LFSCL), very 
long perigynia (PERIGL), very wide perigynia (PERIGW), very long perigynium 


+++ jamesii 
5 10 
78%/2 
HHH} 
Seat 4H juniperorum 
51%/1 24 7 12 
| | 
a 
1 3 
i+ timida 
8" 11 
outgroup 
Fic. 8. The singl imonious t Iting from t i lysi ere 


jamesii sauiplee The outgroup is composed of the“wide- clad lade” of sect. Phyllostachyae (see Materials and Meth- 
ods). The character state changes are represented by vertical bars (character numbers as in Table 5), with 


by superscript “P” on the character numbers, and reversals indicated by superscript “R.” Bootstrap values and decay 


J 


beaks (PERIGBKL), very wide achenes (ACHNW), very long perigynium beaks 
relative to the length of the perigynia (RBK), and a low number of leaves and 
bracts on reproductive shoots (LFNO). Brown shoot bases, cataphylls with col- 
lapsed or flush cell surfaces, and staminate scales with free margins also dis- 
tinguish C. jamesii. 

The status of C. timidaasa distinct species is less obvious than for C. jamesii 
and C. juniperorum. In five quantitative features (MPDCLL, LFSCL, PERIGBKL, 
RBK, LFNO), C. timida is intermediate between C. jamesii and C. juniperorum. 
Of the 16 measured characters with statistically different means that are ex- 
tremes, only 4 of them belong to C. timida. Carex timida is unique in having 
very short staminate portions of terminal spikes (MSPL), very long staminate 
scales (MSCL), very wide margins on the distalmost pistillate scale (MARGW), 
and proximalmost staminate scales that occupy a very high proportion of the 
length of the staminate portion of spikes (RSC). Qualitative features unique to 
C.timida include its lax culms and leaves, its relatively light green foliage, and 
its papillate staminate scales. While the features that distinguish C. timida are 


870 BRIT.ORG/SIDA 19(4) 


less obvious than those of the other species in the complex, it does have several 
unique quantitative and qualitative morphologic features supporting its status 
as a species. We have chosen the epithet “timida” to highlight one of this spe- 
cies’ morphologic qualities (lax culms usually hidden among the foliage and 
thus difficult to detect), but also because the true identity of C.timida had been 
undetected for so long. 

The PCA and bivariate plots indicate that C.jamesii and C. timida are more 
phenetically similar to each other than either is to C. juniperorum (Figs. 1, 2). 
Yet, the phylogenetic analysis indicates that C. juniperorum and C. timida are 
sister species. The retained plesiomorphies of C. jamesii and C. timida, in com- 
bination with the apparently greater amount of anagenesis of C. juniperorum 
(as determined by it having twice as many autapomorphies as C. jamesii and C. 
timida), account for this discrepancy between phenetic and phylogenetic rela- 
tionships. Genetic analysis of the C. jamesii complex also supports the sister- 
group status of C. juniperorum and C. timida (Ford & Naczi 2001). 

The bootstrap and decay values indicate relatively weak support for the 
branches in the phylogenetic tree (Fig. 8). The consistency index (0.67) is rela- 
tively Low, too, due to homoplasy. Given the relatively few morphologic charac- 
ters available for phylogeny reconstruction in these reduced plants, the rela- 
tively weak support and moderate level of homoplasy are not unexpected. 
Despite relatively weak support for the tree generally, the monophyly of the C. 
jamesti complex is well-supported (5 character state changes, bootstrap value 
of 78%, and decay value of 2). 

The presence of a greater amount of morphologic variability in C. jamesii 
and C. timida than in C. juniperorum is congruent with the results of genetic 
analysis of the C. jamesii complex (Ford & Naczi 2001). The reduced level of 
variation of C. juniperorum might be the result of morphologic features that 
limit gene flow (Ford & Naczi 2001). Carex juniperorum does have several char- 
acteristics that would seem to limit outcrossing and seed dispersal, such as very 
short culms, proximal pistillate scales that completely conceal the perigynia, 
and very short stigmas. It is surprising that C. timida would have a level of mor- 
phologic variation similar to the much more common and wider-ranging C. 
jamesii. Still, the geographic range of C. timida is large, and perhaps its rela- 
tively high morphologic variability results from diverse selection pressures and 
ecologic influences across its range. 

One point of disagreement between the results of morphologic and genetic 
studies is in the relationships of plants from a population in Queen Wilhelmina 
State Park, Polk County, Arkansas. Ford & Naczi (2001) found these plants to 
group with C. juniperorum. Based on morphology, a specimen from this popu- 
lation is clearly C. timida (Fig. 1). 

Each member of the C._jamesii complex has a unique geographic distribution. 
The limits of the range of C.jamesii are greater in all directions than the range 


limits of the other two species, except the northeastern extreme. In this area, C. 
juniperorum ranges farther north than C. jamesii. Carex juniperorum is broadly 
sympatric with C. jamesii, though its range is disjunct. The range of C. timida is 
completely encompassed in the range of C.jamesii. Carex timida differs geographi- 
cally from C. jamesii in being limited to three disjunct regions, while C. jamesii 
is relatively continuous across its range. The pattern of disjunction of C. timida 
is similar to several other vascular plant species. These species occur in the Ozark 
Mountains, the Ouachita Mountains, and in areas east of the Mississippi 
Embayment, but not intervening regions. Taxa sharing this pattern of disjunction 
include C.ouachitana Kral, Manhart, & Bryson (Naczi & Bryson 1990; McNeilus 
1992), C. willdenowii Willd. (Nacziet al. 1998), Castanea ozarkensis Ashe Johnson 
1988; Nixon 1997), Cotinus obovatus Raf. (Little 1977), Leptopus phyllanthoides 
(Nutt.) G.L. Webster (Clark 1967; Smith 1988), Magnolia tipetala L. (Little 1977; 
Meyer 1997), Monarda virgata Raf. (Scora 1967; Smith 1988), and Trillium 
pusillum Michx. var. ozarkanum (Palmer & Steyerm.) Steyerm. (Cabe & Werth 
1995). In a study of fish lineages with similar distribution patterns, Strange © 
Burr (1997) found different mechanisms may account for the disjunctions, in- 
cluding fragmentation of formerly widespread ranges by Pleistocene glacia- 
tions, dispersal events, and events that predated Pleistocene glaciations. 

As with morphologic and geographic data, ecologic features distinguish 
the members of the C. jamesii complex. Ecologically, C.juniperorum is the most 
divergent member of the C.jamesii complex. It grows in the most open habitats 
with substrates having the highest clay content. It apparently does not occur 
with C. jamesiti or C. timida, probably because of the distinctiveness of its habi- 
tat. Carex jamesii occurs in relatively moist sites in closed forest. Carex timida 
usually grows in slightly drier sites in more open forests, though it often grows 
near C. jamesii. The syntopic occurrences of C.jamesii and C. timida are signifi- 
cant because the two species probably overlap in flowering period and appear 
to have the opportunity to interbreed when syntopic. Our morphologic and 
genetic analyses revealed no evidence of hybrids. The fact that plants of such 
closely related species co-occur and maintain their distinctions is additional 
evidence of their status as separate species. 

Carex jamesii is the most common member of the complex. In many parts 
of its range, it is one of the most common sedges of mesic deciduous forests. In 
contrast, C.juniperorum and C. timida are truly rare. Their rarity is reflected in 
the relative recency of their earliest collections. Whereas C.jamesii was collected 
at least as early as 1824 (the year of its description), the first known collection 
of C. juniperorum was in 1940 (originally identified as C. jamesii) and of C. 
timida in 1921 (again, originally identified as C. jamesii). Though additional 
collecting undoubtedly will reveal additional populations of these species, we 
recommend a range-wide review of their conservation status, with consider- 
ation for protection. 


872 BRIT.ORG/SIDA 19(4) 


Though the eastern North American flora is relatively well-studied, it does 
continue to harbor undescribed species. As with C. jamesii in this study, mor- 
phologically distinctive species described early from northeastern North 
America have been a rich source of previously unsuspected diversity (Naczi et 
al. 1998). These species are often so distinctive and easy to identify that they are 
thought to be without taxonomic complexity until critically examined. Our 
expectation is that future studies of such species will yield more novelties, in 
sedges as well as in other plant groups. 


TAXONOMIC TREATMENT 


Sectional Nomenclature 

Much confusion has surrounded determination of the correct name and au- 
thorship of the section to which the Carex jamesii complex belongs. Most mod- 
ern authors (e.g. Mackenzie 1935; Fernald 1950; Gleason & Cronquist 1991) have 
used Phyllostachyae Tuck., but recent authors have followed Catling et al. (1993) 
in using Phyllostachys J. Carey) L.H. Bailey. 

Tuckerman (1843) was the first to apply an infrageneric name to the C. 
jamesii complex and its closest relatives. He used “Phyllostachyae” for three spe- 
cies, C. backii, C.jamesii (as C. steudelii),and C. willdenowii. Though Tuckerman 
attributed the name to Gray, Gray apparently did not publish Phyllostachyae 
himself. Tuckerman did not provide a description for Phyllostachyae, nor did 
he make its rank explicit. Christ (1885) used the name Phyllostachyae explic- 
itly as a section for two European species, but again did not provide a descrip- 
tion. It was not until 1909 that Kiikenthal validly published the name 
Phyllostachyae at the sectional level. 

Carey (1848) was the first to use the infrageneric name “Phyllostachys,” for 
the same three species as Tuckerman, and attributed the name to Torrey and 
Gray. The attribution of the name to Torrey and Gray is probably because of 
their suggestion that C. willdenowii may belong toa genus separate from Carex, 
one “...to which the name Phyllostachys would be appropriate” (Torrey 1836: 404). 
While Carey described the infrageneric group he called Phyllostachys, he was 
not explicit about its rank. Apparently, Bailey (1885) was the first to explicitly 
assign the rank of section to the name Phyllostachys. He also listed the mem- 
bers of the section, designated its type, and divided the section into two “subor- 
dinate groups” (rank not explicit). Bailey placed six species in sect. Phyllostachys: 
C. backii, C. geyeri Boott, C. jamesii (as C. steudelii), C. multicaulis L.H. Bailey, C. 
phyllostachys C.A. Meyer, and C. willdenowii. He clearly designated C. 
phyllostachys the type of the section by stating, “..all but the type species exclu- 
sively American.” Furthermore, he regarded C. bachii, C. jamesii, and C. 
willdenowii as different enough from typical members of Phyllostachys to as- 
sign them to the “subordinate group” Bractoideae. 


We regard C. phyllostachys as belonging toa section separate from C. jamesii 
and its closest relatives. Carex phyllostachys lacks the apical dilation of culms 
that characterizes the C. jamesii complex and its allies. In addition, C. 
phyllostachys has multinerved and vetricose perigynia, unlike the unnerved (but 
2-ribbed), nonventricose perigynia in the C.jamesii complex and allies. Because 
Bailey used C. phyllostachys to typify sect. Phyllostachys, “Phyllostachys” cannot 
be used for the section that includes the C.jamesii complex. Instead, the correct 
name and authorship of the section that includes the C.jamesii complex is Carex 
sect. Phyllostachyae Tuck. ex Kuk. 


Carex section Phyllostachyae Tuck. ex Kiik., Pflanzenr. IV. 20 (Heft 38):642. 1909. 


LECTOTYPE, here designated: Carex jamesii Solingen, 


Dapedostachys Borner, Abh. Naturwiss. Vereine Bremen 21:265. 1913. TyPE: Carex steudelii Kunth. 
Non Carex section Phyllostachys (Torrey & A. Gray ex J. Carey) LH. Bailey, Bot. Gaz. 10:208. 1885. 
Type: Carex phyllostachys C.A. Meyer. 


Identification Key to Members of the Carex jamesii Complex 


Mature, Complete. pene am a specimens are necessary for correct identification of the members of 


. Tallest culm 9.0-41 cm ee 39-86% of plant height: sorter Spike ee 
ae wider leaves w 
m wide; perigynium beak 34-53% of perigynium length. 
2. a (per plant) staminate portion of terminal spike 3.4—5.6(-6.2) mm long; 
proximalmost staminate scale in terminal spike (1.9-)2.1—-3.3 mm long, (35-) 
44-77% of length of staminate portion of terminal spike; perigynium beaks 
1.4-2.3(-2.5) mm long, 34-44% of perigynium length; shoot bases tinged with 
red-purple Carex timida 
2. Longest (pe portion of terminal spike (4.9-)5.8-13.5 mm long; 
ey ee staminate Se in terminal spike 1.1-1.8(-2.1) mm long, 13-26(- 
35)% of length of staminate portion of terminal spike; perigynium beaks (1.9-) 
2.3-3.9 mm long, 39-53% of perigynium length; shoot bases lacking red-purple 
Carex jamesii 
1. Tallest culm 3.2-9.1 cm high, 15-32% of plant height; terminal spike with 4-8 
perigynia; wider leaves with hyaline margins 0.05-0.2 mm wide; hyaline margins of 
distal pistillate scales 0.05-0.3 mm wide; perigynium beak 30-38(—43)% of 
perigynium length Carex juniperorum 


Species Accounts 


Carex saan Schwein., Ann. oe Nae te New York Lot 1824. Type: U.S.A. 
NA. [no additional localit ided] James s.n. HOLOTYPE: PH*: 
PROBABLE ISOTYPE: NY). 


Carex steudelii Kunth, Enum. Pl. 2:480. 1837. D ] } lelii ( I x Fedde & 
C. Schuster, Just’s Bot. Jahresber. 41. IL9. 1918. TyPE: U.S.A. OHIO. [County unspecified, but in 
southwestern Ohio in the region drained by the Miami River (R.L. Stuckey, pers. comm.) 
Mia eine Frank s.n. (HOLOTYPE: B, n.v., presumably destroyed; lsoTyPEs: MO, NY). For his- 
cen background on this collection, see ei 74). 


874 BRIT.ORG/SIDA 19(4) 


Perennial herb, densely caespitose. Rhizomes very short, 0.1-0.4 mm long be- 
tween shoots or branches of the rhizomes, covered with cataphylls, with in- 
ternodes 0.1-0.2 mm long and 1.2-L9 mm thick. Shoot bases surrounded by 
cataphylls, dark brown or ferruginous to stramineous, lacking red-purple col- 
oration. Reproductive shoots 12-63 cm tall, spreading; culms 5.7-41 cm tall, the 
tallest 14-41 cm high, slightly to moderately overtopped by the leaves, (0.45-) 
0.52-0.72(-0.86) of shoot height, 0.4-0.7 mm wide at midheight, smooth proxi- 
mally and denticulate in distal half, acutely trigonous and three-winged, di- 
lated just basal to terminal spike, 0.7-1.5 mm wide just basal to terminal spike. 
Cataphylls glabrous, multicostate, with cells with their outer walls sunken or 
flush with their anticlinal walls. Leaves 3-5(-6), blades 1.1-59 cm long, 1.2-3.1 
(-3.5) mm wide, the widest (1.8-)2.3-3.1(-3.5) mm wide, deep green, flat to barely 
plicate, glabrous, adaxial surface smooth or sparsely antrorsely scaberulous in 
distal portion, abaxial surface smooth; margins green, smooth or antrorsely 
scaberulous in distal portion; leaf sheaths 1.8-9.8 cm long, tight, glabrous; 
adaxial face of sheaths with hyaline band, hyaline band with apex subtruncate; 
ligules subtruncate to depressed-lingulate, 0.6-1.4 mm long. Vegetative shoots 
13-57 cm tall; leaves like those of reproductive shoots; pseudoculms 3.3-9.9 cm 
tall, 1.1-2.6 mm wide at mid-height, 0.09-0.30 of shoot height. Infructescence a 
single terminal spike and (0-)I-3 lateral spikes. Spikes androgynous, simple, 
with staminate scales and perigynia spirally and densely imbricate. Terminal 
spike atop widely spreading to nodding culm, 6.4-15.7 mm long, 4.8-9.33 mm 
wide; staminate portion 2.4-10.3(-13.5) mm long, the longest (4.9-)6.0-10.3 
(-13.5)mm long, 0.4-0.7(-0.8) mm wide, 5-12-flowered, on peduncle 0.8-3.8 mm 
long; pistillate portion overlapping and usually exceed by staminate portion 
but sometimes slightly exceeding staminate portion, 6.4-11.3 mm long, 48-93 
mm wide, (1-)2-3(-4)-flowered. Lateral spikes on widely spreading to nodding 
peduncles arising from base of culm; peduncles 2.7-14 cm long, capillary, flat, 
narrowly two-winged, dilated just basal to spike; spikes similar to terminal 
spikes except staminate portion 2.0-9.5 mm long. Proximalmost staminate scale 
of each terminal spike 1.1-1.9(-2.1) mm long, 0.13-0.26(-0.35) of length of stami- 
nate portion of terminal spike, 0.7-14 mm wide, short-cylindric, truncate or 
subtruncate (very broadly ovate), longitudinally 1-veined in basal 0.7, with lon- 
gitudinal and narrow green band centered on vein, transversely brown-banded 
distal to apex of green band; margins free, overlapping base of adjacent distal 
staminate scale but not sheathing it, hyaline. Distalmost pistillate scale of each 
spike 3.8-21 mm long, 1.6-2.4 mm wide, ovate to lanceolate, acute and awnless 
but occasionally with awn to 5.9 mm long or longer scales leaf-like, center green 
and 4-9-veined, margins hyaline, whitish or whitish with brown band paral- 
leling margin, hyaline margins (0.3-)0.4-0.7 mm wide. Proximalmost pistillate 
scale of each spike with morphology dependent on spike position; in terminal 
spikes, proximalmost scale leaf-like; in lateral spikes, proximalmost scale ovate 


— 


NACZI AND FORD, 875 


and awned or awnless. Proximalmost pistillate scale of terminal spikes 12-119 
mm long, the longest 31-119 mm long; basal portions 1.6-2.9 mm wide, par- 
tially or completely concealing perigynia, green with hyaline margins, green 
portion 5-12-nerved, hyaline margins 0.05-0.3 mm wide; distal portions green, 
long-acuminate, with antrorsely scaberulous margins. Other pistillate scales, 
if present between proximalmost and distalmost scales, morphologically in- 
termediate between distalmost and proximalmost scales. Filaments laminar, 
0.20-0.30 mm wide, usually slightly wider than anthers. Anthers 3, 0.8-1.2 mm 
long. Styles jointed with summits of achenes, portions distal to achenes wither- 
ing with age. Stigmas 3, 1.7-2.1 mm long, withering with age. Perigynia (4.8-) 
5.4-6.8(-7.6) mm long, L.8-2.3(-2.5) mm wide, (2.3-)2.5-3.3(-3.6) times as long 
as wide, (2.1-)2.4-3.0 times as long as achenes, ascending to spreading, 
subrotund to very broadly trigonous in cross-section, 2-ribbed, nerveless, gla- 
brous, pale green to tan, narrowly ovoid to narrowly ellipsoid with obovoid body 
and compressed-deltoid beak; body tapered from widest point to subacute base, 
abruptly contracted to beak; beak (1.9-)2.3-3.4(-3.9) mm long, (0.39-)0.43-0.53 
of perigynium length, straight, scaberulous on angles, apex entire. Achenes 1.9- 
2.8 mm long, 18-2.4 mm wide, 0.9-14 times as long as wide, tightly enveloped 
by perigynia, subrotund to very broadly trigonous in cross section, brown wit 
3 longitudinal paler brown lines that intersect at style base, subglobose, ba- 
sally abruptly contracted to light tan stipe, beakless; stipes 0.3-0.4 mm long, 
0.8-1.1 mm wide (Fig. 3A, 4, 5A). 
imens. Speci 
Ses. CANADA. ONTARIO. Essex Co.: Pelee Island, Fish Point, 25 Jun 1981, Reznicek 6338 (MICH, MO, 
VPI, ctb). Kent Co.: Orford Township, Clear Creek, 22 May 1991, Oldham 12554 (MICH*). Middlesex 
Co.: 9 km ESE of Glencoe P.O.,N side of Thames River, 29 May 1987, Reznicek 7899 (MICH, MO, ctb”*). 
Welland Co.: 2.5 mi W of Port Colborne, near Rathfon Point, 1 Jul 1958, Calder 23864 (MICH*, MT, US). 
U.S.A. ALABAMA. Madison Co.: Huntsville, Monte Sano State Park, 8 May 1986, Naczi 1020 (DOV). 
Marshall Co.: near Grant, near bottom of Fletcher's Hollow, 13 May 1939, Harper 3728 (MICH*, US). 
Tuscaloosa Co.: along Warrior River, ca.0.25 mi above Lock 14, 10 May 1935, Harper 3366 (MO, PH, 
US). ARKANSAS. Benton Co.: N of Bella Vista, 17 May 1986, Castaner 9209 (MO, ctb). Carroll Co.: US 
highway 412 and Osage Creek, 24 May 1992, Hyatt 4386.08 (MICH, ctb*). Cross Co.: W of Birdeye, 20 
May 1990, Naczi 2449 (DOV, ctb). Franklin Co.: ca. 1 miN of Cecil, 12 May 1989, Naczi 2151 (DOV, ctb). 
Howard Co.: ca.8 mi W of Umpire, N of route 4, along W side of Cossatot River, 17 el 1988, Naczi 
1925 (MICH*). Marion Co.: Hand Valley at Ranchette (White River) Access Area, 1 May 1992, Hyatt 
4821.45 (MICH’).Montgomery Co.: ca.6 mi NW of Mount Ida, 20 May 1994, Naczi 3935 & Ford (DOV). 
Scott Co.: ca.2 mi N of Y City, 10 May 1989, Naczi 2128 (DOV, ctb). Stone Co.: ca.3 mi ENE of Fifty Six, 
21 May 1986, Naczi 1206 (DOV). DISTRICT OF COLUMBIA. Washington and vicinity, 15 May 1896, 
Steele s.n. (MU*). ILLINOIS. Cook Co.: Elk Grove, 28 May 1948, Chase 9497 (NY). Jackson Co.:5 mi SW 
of Murphysboro, 11 Apr 1976, Solomon 1453 (MO). Jo Daviess Co.: 7.5 mi NW of Stockton, Apple 
River Canyon, 16 Jun 1937, Hermann 8842 (NY). Peoria Co.: N of Princeville, 16 Jun 1897, Chase 36 
(MO, PH, US). Pulaski Co.: 2.1 mi SW of Olmsted, along E side of route 37, 26 May 1990, Naczi 2488 
(DOV*, ctb) Saline Co.: ca.7 mi ESE of Harrisburg, 20 May 1992, Hyatt 4356 (MICH*). Winnebago Co.: 
W of Rockford, 4 Jun 1954, Fell 54303 (US). INDIANA. Carroll Co.: 5 mi £ of Brookston, 8 Jun 1924, 
Deam 40663 (NY). Henry Co.:3 mi E of Mt. Summit, 24 May 1938, Kriebel 5284 (NA). Wabash Co.: ca. 


Ara | I iC me i 


— 


— 


876 BRIT.ORG/SIDA 19(4) 


6 mi SW of Wabash, Asher Wood Nature Preserve, 21 May 1991, Rothrock 2271 ee: IOWA. Hardin 
Co.: Near Eldora, along lowa River, 3 Jun 1950, Thorne 9619 (MT, US). Johnson Co.: McBride State 
Park, 25 May 1954, Thorne 14072 (DHL, on NY). Mills Co.: Wilson Timber, sate R41W, section 24, 29 
May 1992, Wilson 5175 (MICH’). Story Co.: Ames, 16 May 1938, Weber 616 (MO, OS). KANSAS. Linn 
Co.: La Cygne Lake Park, along road to campground, 28 May 1994, Reznicek 9819 & Reznicek (MICH*, 
ctb). Miami Co.: Miami County State Lake, 16 May 1964, Harms 2046 hae oo ada Co.: 
ca. 3 mi SSW of Haywood, Brigadoon State Nature Preserve, 14 Jun 1997, g (KNK*). 
Bath Co.: 4 mi N of Salt Lick, 12 May 1999, Naczi 7830 (DOV). eine Co.: ee W Fork 
Clarks River, 30 May 1973, Athey 2332 (MUR).Campbell Co.: ca. 2 mi S of Alexandria, along E side of 
route 27,5 Jun 1996, Naczi 5649 et al. (DOV*), the Co.: Lexington, 6 Jun 1942, McFarland 93 (MO, 
NA, PENN, PH, TENN, US). Graves Co.: Folsomdale, 24 Apr 1972, Athey 1608 (MUR, WKU). Henderson 
Co.: Henderson, Audubon State Nature Preserve, 14 Jun 1997, Naczi 6477 & Heeg (KNK*). Laurel Co.: 
ca.6 mi WNW of Bernstadt, along E side of Rockcastle River, 21 May 1998, Naczi 7139 & Ford (DOV*). 
Letcher Co:: S from Whitesburg, just S of crest of Pine Mountain, 17 May 1994, McKinney 6177 (DOV). 
Madison Co.: 5 mi SW of center of Richmond, 15 Jun 1998, Naczi 7373 & Reznicek (DOV*). Monroe 
Co.: ca.8 mi SE of Tompkinsville, S of route 216,23 May 1998, Naczi 7236 & Ford (DOV). Pendleton 
Co.: 2 mi ESE of Falmouth, 3 Jun 1994, Naczi 4039 & Thieret (DOV, WIN, ctb). MARYLAND. Cecil Co. 
1.0 mi SE of Conowingo,0.2 mi E of route 222 bridge, along Octoraro Creek,29 May 1999, Naczi 8051 
(DOV*). Kent Co.: N of Copeland, 6 Jun 2001, McAvoy 5112 (DOV).Montgomery Co.: Cropley, along 
Potomac River, 4 May 1929, Blake 10833 (MT, NA). MICHIGAN. Allegan Co.: 18 mi S of Grand Rapids, 
18 Jun 1942, Bazuin 4551 (KY).Ingham Co.: eee [no ae) /eu0 1891,C.F Wheeler s.n. (MICH). 
Washtenaw Co.: 4.75 mi S of Ann Arbor, 17 May 193 9419 (NA, US).Wayne Co.: ca. 2.5 mi 
E of Belleville, Lower Huron Metropark,N side ae River, 24 hay 1991, Reznicek 8759 etal. (MICH*, 
VPI, ctb). MINNESOTA. Fillmore Co.: Forestville State Park, adjacent to South Branch Root River, 19 
Jun 1984, G.A. Wheeler 8361 (ctb*). Houston Co.: Beaver Creek Valley State Park, 19 Jun 1984, Wheeler 
91 (ctb). MISSISSIPPI. Lowndes Co.: 2 mi S of Artesia, 15 May 1989, Bryson 8557 (ctb). Oktibbeha 
mi E of Mississippi State University, 15 May 1989, Bryson 8586 & Naczi (OS, ctb’). aa 
Co.: Yellow Creek Quad, Burns Hollow, 15 May 1979, Rogers 46688 (TENN). Tunica Co.: ca. 11 mi S of 
Tunica, 24 Apr 1992, Bryson 11400 & Newton (MO, ctb). MISSOURI. Boone Co.: Sane Grind- 
stone Park, 8 Jun 1996, Reznicek 10184 et al. (DOV, MICH). Callaway Co.: 147N, R8W, section 3, 24 Apr 
1991, Currier 91-004 (ctb*). Christian Co.: 0.5 mi S of end of road FF, 27 Apr 1985, Castaner 8265 
(DOV). Cooper Co.: 0.9 mi N of junction of routes AE & Z,21 May 1981, Castaner 6380 (ctb*). Jack- 
Co.: Independence, 2 Jun 1895, Bush 709 (DOV, NY). Schuyler Co.: 3 mi E of Livonia, along E side 
of Chariton River, 1 Jul 1933, Palmer 4106 1a & Steyermark (MO). Scott Co.: 2.4 mi S of Blodgett, 0.6 mi 
on main road into Fox Meadow Estates, 16 May 1992, Jones 8546 & Jones (MICH*). NEBRASKA. 
Richardson Co.: SE of Rulo, along Nemaha River, 22 Jun 1940, Reynolds 1450 (MO, NA). NEW JERSEY, 
Hunterdon Co.: Lambertville, foot of Goat Hill, near Delaware River, 31 May 1924, Long 30383 (PH). 
Mercer Co.: N of Moore, 29 Apr 1922, Mackenzie s.n. (NY). NEW YORK. Monroe Co.: Near Honeoye 
Falls, 15 Jun 1941, Matthews 4339 (KY, MICH, MO, MT, NA, PENN, TENN, US). Tompkins Co.: Ithaca, Six 
Mile Creek, 8 Jun 1919, Wiegand 11595 (MICH, MO, MT, PENN, PH, US). NORTH CAROLINA. Durham 
«3 or 4 mi N of Durham, ca. 0.25 mi E of Old Oxford Road, along S side of Eno River, 8 May 1972, 
ae a nes. SE beet MUR, MICH’, TENN, VPI). Lee Co.: 2 mi W of Moncure, near Deep River, 
3 Jun 1960, Radford (KY, TENN). OHIO. Butler Co.: £ of Miami University campus, E of Marcum 
Conference Center, 30 Te 991, Vincent 4539 (MU*). Lawrence Co.: 0.5 mi W of Burlington, 21 Apr 
1992, Cusick 30148 (MICH). Montgomery Co.: ca. 2 mi SW of Farmersville, ca. 0.2 mi E of Anthony 
Road, 0.3 mi SSW of junction of Anthony Road and Manning Road, 29 May 1998, Naczi 7363 & Ford 
(DOV*). OKLAHOMA. Cherokee Co.: 7.5 mi NNE of Tahlequah, 13 May 1987, Naczi 1591 (DOV). 
Delaware Co.: along N shore of Lake Eucha, 1.6 mi E of route 59/10, 25 vice 994, Reznicek eke 
Reznicek (MICH*, ctb). McCurtain Co.: ca. 9 mi NE of Broken Bow, 12 1987, Naczi 1576 (DO 
MICH). Muskogee Co., Braggs Hill Canyon, 6 May 1940, Bebb 5163 Ae BENNEV ANIA re 


— 


ND FORD, 877 


Co.: 2.25 mi WNW of Everett, 17 Jun 1944, Berkheimer 4920 (PENN, PH). Berks Co.: 0.75 mi W of 
Sinking Spring, 6 Jun 1943, Berkheimer 3668 (PENN, PH, TENN). Bucks Co.: E of Ottsville, by Tinicum 
Creek, 30 May 1925, Long 32469 (PENN, PH).Centre Co., Woodward, 6 Jun 1937, Wahl 2539 & Clausen 
(NA, US). Franklin Co.: ca. 1.5 mi WSW of Yeakle Mill, 7 May 1993, Kunsman 106174 (PH). Lancaster 
Co.: Shenk’s Ferry Ravine, 22 Jun 1940, Tanger 3885 (PENN, PH). Westmoreland Co.: Near Saunders, 
Turtle Creek, 27 May 1933, Bright 8838 (PENN). SOUTH CAROLINA. McCormick Co.: E of Savannah 
River, Steven's Creek Preserve, 17 Apr 1992, Sorrie 6257 et al. (bas). TENNESSEE. Cannon Co.: ca. 1.5 
mi NE of Sugar Tree Knob Church, 3.0 mi W of TN 146,N side of Short Mountain Road, 3 May 1989, 
Orzell & Bridges 9436 (MICH*). Cheatham Co.: S of I-40 and TN 239 junction, above large truck park, 
28 Apr 1993, Kral 8211 ae ctb). Davidson Co: Nashville, 30 May 1990, Naczi 2518A & Kral (DOV). 
Dekalb Co.: 0.4 miW on TN 141 from junction TN 96 at Center Hill Dam, 30 May 1988, Thompson 8&- 
617 (BEREA). Dyer Co.: NW be Dyersburg, 29 Apr 1949, Sharp 12231 et al. (TENN). Franklin Co.: Near 
Sherwood, 3 May 1941, Shanks 1305 (TENN). Obion Co.: ca. 3.5 mi E of Ridgely, 27 Jun 1995, Naczi 
4852 & Reznicek (DOV). Perry Co.: SE of Linden on old SR 13,E side of Buffalo River, 4 May 1993, Kral 
82279 (MICH*, ctb). Putnam Co.: ee Fork River, 27 Apr 1973, Kral 49694 (MO, PH).Rutherford Co.: 
W of Murfreesboro, off highway 96, Scales Mountain, 26 Apr 1987, McKinney 2420 ae eee 
3 mi N of Hartsville, 4 oe 1973, Kral 49794 (MO, PH). VIRGINIA. A ttox Co.: Just N of 
Beni eae 1 Jun 1967, Harvill 16519 (MO). Bath Co.: ca. 0.4 mi S of Healing Springs, 23 ne 1994, 
Naczi 4482 & Thieret (DOV). Fauquier Co.: 2.5 mi below Thorofare Gap, 13 May 1945, Allard 11243 
(MT, PENN, VPI), Lunenburg Co.: ca. 13 mi SE of Lunenburg, along Flat Rock Creek, 3 Jun 1986, Wieboldt 
5982 (MICH, MO, VPI, ctb). Prince George Co.: By James River, Indian Point, 6 May 1940, Fernald & 
Long 11768 (MO, PH, US). Rockbridge Co.: Natural Bridge, nee ial 30 May 1891, ee . 
(MO). Surry Co.: Along James River, Sati 20 May 193 ld & Long 9862 (PH 
Wythe Co.: Jackson's Ferry, 22 J 940, Hermann 10649 a WEST VIRGINIA. Nee oe 
Guyandotte, 13 Apr 1995, Cusick er es MO, MU, VPI, ctb). Fayette Co.: ca.8 mi NE of Beckley, 
New River Gorge, 20 May 1985, Wieboldt 5523 (MICH, VPI, ctb). Mason Co.: 0.5 mi down 16 Mile 
Creek Road (78),W of route 35,22 May 1991, Vincent 4707 & Hickey (MU*). Pendleton Co.: ca. 3.25 mi 
WNW of Cherry Grove, 23 May 1988, Reznicek 8140 & Reznicek (MICH, MO,VPI,ctb). Wayne Co.: Mouth 
of Big Creek, 6 May 1938, MacFarland 4353 (NA). WISCONSIN. Grant Co.: T4N, R6W, sect. 15, 26 May 
1979, Tans 1920 (WIS). Green Co.: On “K" near Oakley store, 2 Jun 1958, Fel! 58-176 (WIS). LaCrosse 
Co.: Branch of Coon Creek in Bohemian Valley, 19 Jun 1959, Hartley 6774 (WIS). 


Carex juniperorum Catling, Reznicek, & Crins, Syst. Bot. 18:497. Type: CANADA. 

ARIO. HASTINGS Co. 7 km NE of Shannonville, S side of hwy. 401, 9 Jun 1991, Catling 

9100 (HOLOTYPE: DAO, n.v; ISOTYPES: KNK*, MICH). 

Perennial herb, densely caespitose. Rhizomes very short, 0.1-2.1 mm long be- 
tween shoots or branches of the rhizomes, covered with cataphylls, with inter- 
nodes 0.1-1.6 mm long and 1.3-1.9 mm thick. Shoot bases surrounded by 
cataphylls, dark red-purple to ferruginous, usually dark brown tinged with red- 
purple; red-purple coloration extending 17-68 mm above base of plant. Repro- 
ductive shoots 8.2-45 cm tall, spreading; culms 1.9-9.1 cm tall, the tallest 3.2-9.1 
cm high, greatly overtopped by the leaves, 0.15-0.32 of shoot height, 0.3-0.7 mm 
wide at midheight, smooth except denticulate just basal to terminal spike, 
acutely trigonous and three-winged or occasionally four- or five-angled and 
four- or five-winged, dilated just basal to terminal spike, 0.8-1.3 mm wide just 
basal to terminal spike. Cataphylls glabrous, multicostate, with cells with their 
outer walls bulging. Leaves 6-11, blades 11-43 cm long, 0.7-4.2 mm wide, the 


878 BRIT.ORG/SIDA 19(4) 


widest 2.2-4.2 mm wide, deep green, flat to barely plicate, glabrous, adaxial 
surface smooth or minutely papillate or sparsely antrorsely scaberulous in dis- 
tal portion, abaxial surface smooth or minutely papillate; margins of wider 
blades hyaline, hyaline portions 0.05-0.2 mm wide; margins of narrower blades 
green; margins of all blades smooth or antrorsely scaberulous in distal portion; 
leaf sheaths 1.4-8.2 cm long, tight, glabrous; adaxial face of sheaths with hya- 
line band, hyaline band with apex truncate; ligules truncate to depressed- 
lingulate, O-0.7 mm long. Vegetative shoots 9.8-45 cm tall: leaves like those of 
reproductive shoots; pseudoculms 1.3-5.6 cm tall, 1.1-3.7 mm wide at mid-height, 
0.11-0.16 of shoot height. Infructescence a single terminal spike and (0-)1-3 lat- 
eral spikes. Spikes androgynous, simple, with staminate scales and perigynia 
spirally and densely imbricate. Terminal spike atop erect or oo culm, 
5.4-13.1 mm long, 4.9-8.8 mm wide; staminate portion 3.1-8.4 mm long, the long- 
est 3.6-8.4 mm long, 0.4-0.9 mm wide, 8-17-flowered, on peduncle 0.1-1.4 mm 
long; pistillate portion overlapping and exceeding staminate portion or slightly 
exceeded by staminate portion, 5.4-9.4 mm lone, 4.9-8.8 mm wide, 4-8-flow- 
ered. Lateral spikes on ascending to spreading peduncles arising from base of 
culm; peduncles 0.7-6.3 cm long, capillary, flat, narrowly two-winged, dilated 
just basal to spike; spikes similar to terminal spikes except staminate portion 
L.7-2.9 mm long, pistillate portion (2-)4-6-flowered. Proximalmost staminate 
scale of each terminal spike 0.9-1.6 mm long, 0.13-0.25(-0.34) of length of stami- 
nate portion of terminal spike, 1.1-1.6 mm wide, short-cylindric, truncate or 
subtruncate (very broadly ovate), longitudinally 1-veined in basal 0.7-0.8, with 
longitudinal and narrow green band centered on vein, with green band nar- 
rowly bordered by dark brown; margins connate in basal 0.3-0.8, free apically, 
tightly sheathing adjacent distal staminate scales, hyaline. Distalmost pistil- 
late scale of each spike 3.0-10.1 mm long, L.1-1.5 mm wide, lanceolate, leaf-like, 
acute, awnless, green and 1-5-veined except for margins, margins hyaline, whit- 
ish, hyaline margins 0.05-0.2(-0.3) mm wide. Proximalmost pistillate scale of 
each spike leaf-like. Proximalmost pistillate scale of terminal spikes 12-36 mm 
long, the longest 16-36 mm long; basal portions 1.6-3.4 mm wide, completely 
concealing perigynia, green with hyaline margins, green portion 3-9-nerved, 
hyaline margins 0.05-0.2 mm wide; distal portions green, long-acuminate, with 
antrorsely scaberulous margins. Other pistillate scales morphologically inter- 
mediate between distalmost and proximalmost scales. Filaments laminar, 0.20- 
0.30 mm wide, usually slightly wider than anthers. Anthers 3, L1-1.7 mm long. 
Styles jointed with summits of achenes, portions distal to achenes withering 
with age. Stigmas 3, 0.8-1.2 mm long, withering with age. Perigynia (3.9-)4.2- 
5.0(-5.4) mm long, 1.7-2.1 mm wide, 2.0-2.6(-3.0) times as long as wide, 2.0-2.3 
times as long as achenes, ascending to spreading, subrotund to very broadly 
trigonous in cross-section, 2-ribbed, nerveless, glabrous, pale green to tan, nar- 
rowly ovoid to narrowly ellipsoid with obovoid body and compressed-deltoid 


RCTICANREADT 879 


beak; body tapered from widest point to subacute base, abruptly contracted to 
beak; beak (1.2-)1.4-1.8(-2.2) mm long, 0.30-0.38(-0.43) of perigynium length, 
straight, smooth or barely scaberulous on angles, apex entire. Achenes 2.0-2.5 
mmm long, 1.6-2.1 mm wide, 1.0-14 times as long as wide, tightly enveloped by 
perigynia, subrotund to very broadly trigonous in cross section, brown with 3 
longitudinal paler brown lines that intersect at style base, subglobose, basally 
abruptly contracted to light tan stipe, beakless; stipes 0.3 mm long, 0.7-0.8 mm 
wide gay SB. 5B). 


+ i 


Ses. | CANADA. ONTARIO: cna: Co.: 5.5 km NE of Shannonville, W of Salmon River, 11 Jun 1991, 
Catling 9102 (MICH*); 2 km SSW of Lonsdale, 13 Jun 1994, Catling 20285 & Norris (MICH*). U.S.A. 
KENTUCKY. Bath Co.: Salt Lick, 12 May 1940, McFarland 4799 (NA*); 1 mi W of Polksville on US 60, 
“Blue Clays” on Fearing Road, 31 May 1991, Campbell s.n. (MICH*); ca.5 mi ESE of Owingsville, ca. 0.5 
mi S of route 60,0.3 mi E of Ore Mine Road, 16 May 1994, Naczi 3890 (DOV*, MICH, WIN, ctb). Fleming 
Co.: ca.4 mi E of Hillsboro, along N side of route 1013, 28 May 1998, Naczi 7321 & Ford (DOV*, MICH). 
Lewis Co.: 3.2 mi E of routes 10 & 57 junction in Tollesboro, S of route 10,5 May 1994, Naczi 3802 et 

al. (DOV*, VDB, WIN, ctb); ca. 3.5 mi ESE of Trinity, Hughes Knob, 5 May 1994, Naczi 3808 et al. (APSC, 
DOV*, VDB, WIN, ctb); ca. 3.3 mi SW of Concord, N side of Crooked Creek Road, 30 May 1996, Naczi 
5524 et al. (DOV*, WIN); ca. 1 mi E of Concord, N side of route 8, 30 May 1996, Naczi 5538 & Trauth 
(APSC, DOV*, MICH, VDB, VPI, WIN, ctb). OHIO. Adams Co.: 1.5 mi N of West Union on S side of Adams 
Lake, Adams Lake State Park, 6 May 1991, Reznicek 8742 et al. (MICH*, OS); 3.5 mi NW of West Union, 
Chaparral Prairie Preserve, Hawk Hill Rd.,6 May 1991, Reznicek 8744 et al. (MICH*); 3/5 mi S of Lynx, 
Lynx Prairie,6 May 1991, Reznicek 8748 et al. (DOV*, MICH, OS); 3 mi SW of Lynx,“Hanging Prairie” N of 
Black Run Rd.,6 May 1991 |Rezni ek 8750 - (MICH*); ca. 3 mi NE of Peebles, S side of route 32, 16 
May 1994, Naczi 3878 (DOV*, WIN, ctb). VIRGINIA. Montgomery Co.: Radford, Wildwood Park, W- 
facing slope above Connelly’s Run, 9 Jun 1999, Wieboldt 10214 (MICH*, VPI). 


Carex timida Naczi & B.A. Ford, sp. nov. Type: U.S.A. KENTUCKY. ROWAN Co: ca. 6.5 mi § 
of center of Morehead, ca. 0.25 mi downslope from W side of route 1274, upslope from Sugar 
Camp Branch, ca. 1.3 road mi N of junction of routes 1274 and 801, | Jun 1996, Naczi 5598 & 
Trauth (AOLOTYPE: DOV*; IsOTYPES: BRIT, CAN, DAO, EKY, FE GA, GH, IBE, KNK, KY, MDKY, 
MICH, MO, MU, NCU, NY, OS, PH, TENN, UARK, UNA, US, USCH, VDB, VPI, WIN, ctb). 
A Carex jamesii basibus surculorum purpureis, rostris perigyniorum brevioribus differt; a Carex 
juniperorum culmis altioribus, laminis foliorum latorum viridimarginatis, spicis terminalibus 2- 
3(-4) perigyniis instructis, partibus hyalinis squamarum pistillatarum distalibus 0.4-0.7 mm latis 
differt; a speciebus ambabus partibus staminatis spicarum brevioribus, squamis staminatis 
longioribus differt. 
Perennial herb, densely caespitose. Rhizomes very short, 0.1-0.5 mm long between 
shoots or branches of the rhizomes, covered with cataphylls, with internodes 
0.1-0.2 mm long and 1.4-1.8 mm thick. Shoot bases surrounded by cataphylls, 
dark red-purple to ferruginous, usually dark brown tinged with red-purple; red- 
purple coloration extending 34-47 mm above base of plant. Reproductive shoots 
17-45 cm tall, spreading; culms 4.6-34 cm tall, the tallest 9.0-34 cm high, 
slightly to moderately overtopped by the leaves, (0.39-)0.59-0.83 of shoot height, 
0.4-L0 mm wide at midheight, smooth except denticulate just basal to termi- 
nal spike, acutely trigonous and three-winged, dilated just basal to terminal 


880 BRIT.ORG/SIDA 19(4) 


spike, 0.7-1.3 mm wide just basal to terminal spike. Cataphylls glabrous, 
multicostate, with cells with their outer walls bulging. Leaves +-6(-8), blades 
4.1-35 cm long, 1.0-4.3 mm wide, the widest (1.6-)2.1-4.3 mm wide, medium 
green, flat to eaey plicate, glabrous, adaxial sur face emo or Sparsely 
antrorsely scaberulous in distal portion, abaxial surface smooth; margins green 
smooth or antrorsely scaberulous in distal portion; leaf sheaths L 3-6. Sem 1 long, 
tight, glabrous; adaxial face of sheaths with hyaline band, hyaline band with 
apex subtruncate to slightly convex; ligules subtruncate to depressed-lingulate, 
0.2-1.8 mim long. Vegetative shoots 18-45 cm tall; leaves like those of reproduc- 
tive shoots; pseudoculms 3.2-7.6 cm tall, 1.5-2.9 mm wide at mid-height, 0.14- 
0.20 of shoot height. Infructescence a single terminal spike and 1-3 lateral spikes. 
drogynous, simple, with staminate scales and perigynia spirally and 
densely imbricate. Terminal spike atop widely spreading to nodding culm, 6.8- 
8.8 mm long, 4.6-6.8 mm wide; staminate portion 2.4-5.6(-6.2) mm long, the 
longest 3.4-5.6(-6.2) mm long, 0.6-0.8(-1.0) mm wide, 2-12-flowered, on pe- 
duncle 0.4-2.3 mm long; pistillate portion overlapping and exceeding stami- 

nate portion or slightly exceeded by staminate portion, 6.8-8.5 mm long, 4.6- 
6.8 mm wide, 2-3(-4)-flowered. Lateral spikes on widely spreading to nodding 
peduncles arising from base of culm; peduncles 1.4-21 cm long, capillary, flat, 
narrowly two-winged, dilated just basal to spike; spikes similar to terminal 
spikes except staminate portion 1.4-4.2 mm long. Proximalmost staminate scale 
of each terminal spike (1.9-)2.1-3.3 mm long, (0.35-)0.44-0.65(-0.77) of length 
of staminate portion of terminal spike, 0.9-1.7 mm wide, short-cylindric, trun- 
cate or subtruncate (very broadly ovate), longitudinally l-veined in basal 0.7, 
with longitudinal and narrow green band centered on vein, transversely brown- 
banded distal to apex of green band or with brown patch completely surround- 
ing green band; margins connate in basal 0.3-0.8, free apically, tightly sheath- 
ing adjacent distal staminate scales, hyaline. Distalmost pistillate scale of each 
spike 3.1-8.6 mm long, l.6-2.4 mm wide, ovate, usually acute and awnless but 
occasionally with awn to 2.3 mm long, center green and 3-7-veined, margins 
hyaline, whitish with brown band paralleling margin, hyaline margins 0.4-0.7 
mm wide. Proximalmost pistillate scale of each spike with morphology depen- 
dentons pike position; in terminal spikes, proximalmost scale usually leaf-like, 
rarely like distalmost scale; in lateral spikes, proximalmost scale like distalmost 
scale. Proximalmost pistillate scale of terminal spikes 11-42(-81) mm long, the 
longest 17-42(-81) mm long; basal portions 1.6-2.1 mm wide, partially conceal- 
ing perigynia, green with hyaline margins, green portion 5-12-nerved, hyaline 
margins 0.05-0.3 mm wide; distal portions green, long-acuminate, with 
antrorsely scaberulous margins. Other pistillate scales, if present between 
proximalmost and distalmost scales, morphologically intermediate between 
distalmost and proximalmost scales. Filaments laminar, 0.15-0.20 mm wide, 
usually slightly wider than anthers. Anthers 3, 0.4-1.5 mm long. Styles jointed 


— 


Spikes 


NACZI AND FORD, 881 


with summits of achenes, portions distal to achenes withering with age. Stig- 
mas 3, 1.4-1.6 mm long, withering with age. Perigynia (4.0-)4.4-5.6(-6.0) mm 
long, 1.6-2.1 mm wide, (2.2-)2.5-3.1 times as long as wide, 1.9-2.4 times as long 
as achenes, ascending to spreading, subrotund to very broadly trigonous in 
cross-section, 2-ribbed, nerveless, glabrous, pale green to tan, narrowly ovoid to 
narrowly ellipsoid with obovoid body and compressed-deltoid beak; body ta- 
pered from widest point to subacute base, abruptly contracted to beak; beak 
(1.4-)1.7-2.3(-2.5) mm long, 0.34-0.44 of perigynium length, straight, 
scaberulous on angles, apex entire. Achenes 2.1-2.5 mm long, 1.6-2.1 mm wide, 
1.0-1.4 times as long as wide, tightly enveloped by perigynia, subrotund to very 
broadly trigonous in cross section, brown with 3 longitudinal paler brown lines 
that intersect at style base, subglobose, basally abruptly contracted to light tan 
stipe, beakless; stipes 0.3-0.5 mm long, 0.7-0.9 mm wide (Figs. 3C, 4, 5C). 


Additional Specimens Examined. Specimens marked with asterisks are those measured for statisti- 
cal analyses. U.S.A. ALABAMA. Colbert Co.: ca.4 mi S of Tuscumbia, N slope of Little Mountain, 31 
Mar 1935, Harper 3325 (MO,NY*, PH, US). Jackson Co.: 9 mi S of Huntland [Tennessee], by Ala.65, 28 
Apr 1972, Kral 45800 (MO, ctb*). Madison Co.: Near anaes W dace epMione te Sano Mountain, 14 
Apr 1935, Harper 3331 (MO,NY,PH,US);E Huntsville, Mt.A e San khead Parkway, 2 Ma 
1974, Bryson 492 (ctb); E side of Huntsville, summit of Monte Sano Mountain, along entrance drive 
to Monte Sano State Park, 25 May 1998, Naczi 7264 & Ford (DOV", MICH, VDB, WIN, ctb). ARKANSAS. 
Baxter Co.: Clifty Canyon Botanical Area, Ozark N | Forest, T17N,R12W,NE 1/4 of section 33,14 
Apr 1992, Hyatt 4528.03 (MICH*),T17N, R12W, NW 1/4 of section 32,30 Apr 1992, Hyatt 4815.03 (VDB). 
Howard Co.: To NE of Cossatot River and Arkansas highway 4 crossing, 10 May 1986, Bryson 4333 
(MICH, VDB, ctb); ca. 8 mi W of Umpire, N of route 4, along W side of Cossatot River, 17 May 1988, 
Naczi 1918 (MICH*), 17 May 1988, Naczi 1921 (MICH). Montgomery Co.: Ouachita oerie aie 
ca. 0.2 mi W of Little Missouri Trail’s W end, along Blaylock Bee T4S, R27W, SE 1/4 of S of 
section 29,15 Apr 1996, Hyatt 6937 (MICH*). Pike Co.: Ouachita National Forest, along Little ane 
River, SW of Raven Creek, 75S, R27W,N edge of section 5,19 Apr 1997, Hyatt 7343 (DOV, MICH*). Polk 
Co.: Blackfork Mountain, in ravine NE of Littlke Cemetery, TIN, R32W, W1/2 of SE1/4 of section 26,4 
May 1982, Rettig 503 (VDB), 4 May 1982, Rettig 504 (VDB); Along Cossatot River above route 246 
bridge, 3 May 1983, Rettig 810 (VDB), Rettig 825B (VDB), 10 May 1986, Bryson 4323 (IBE, MICH, MO, 
TENN, VDB, ctb), 12 May 1987, Naczi 1579 (DOV), 14 May 1987, Castaner 9681 (MO, ctb), 13 May 1990, 
Jones 4648 & Jones (MICH, MO, VDB, ctb), 18 May 1993, Jones 10139 & Jones (MICH, VPI*), 20 May 1994, 
Naczi 3949 & Ford (DOV); Queen Wilhelmina State Park, below and to N of visitor center and Arkan- 
sas highway 88,115, R32W, section 11,10 May 1986, Bryson 4287 (MICH, MO, TENN, VDB, ctb), 11 May 
1989, Naczi 2137 (DOV*, ctb), 20 May 1994, Naczi 3940 & Ford (DOV). Stone Co.: ca. 2 mi NW of Fifty- 
six, Ozark National Forest, Sylamore Ranger District, T16N, R12W, section 27, 28 Jun 1993, Hyatt 5653 
(VDB). INDIANA. Jay Co.: 2 mi SE of Boundary, 14 May 1921, Deam 33879 (NY, PH*). Lawrence Co.: 
Bedford, Wilson Park, along Spider Creek, 5 Jun 1934, Kriebel 1768 (PH*). KENTUCKY. Campbell Co..: 
ca.2 mi S of Alexandria, along E side of route 27,5 Jun 1996, Naczi 5650 et al. (DOV*).Monroe Co.: ca. 
8 mi SE of Tompkinsville, S of route 216, 23 May 1998, Naczi 7232 & Ford (DOV*, MICH, MU, TENN, 
USCH, VDB, VPI, WIN, ctb), Ford 98700 & Naczi (WIN). Rowan Co.: Type locality, 28 May 1998, Naczi 
7313 & Ford (DOV,WIN), Ford 98145 & Naczi (WIN). MISSOURI. Taney Co.: 6 mi SE of Protem,4 mi S of 
Ocie, in Big Cedar Hollow, just W of Taney Co. line, section 18, 30 Apr 1938, Steyermark 5303 (MO, 
NA*). OHIO. Montgomery Co.: £, Anthony Road, 0.3 mi SSW, junction of Manning Road, NE1/4, 
section 5,German Twp., 14 May 1985, Cusick 25229 (MU*), ca. 2 mi SW of Farmersville, ca.0.2 mi E of 
Anthony Road, 0.3 mi SSW of junction of Anthony Road and Manning Road, 29 May 1998, Naczi 


WS 


882 BRIT.ORG/SIDA 19(4) 


/357 & Ford (DOV, MICH, WIN, ctb), Ford 98153 & Naczi (WIN). [Preble or Butler Co.]: N of Oxford, E 
Hueston’s woods, 20 May 1934, Belk s.n. (MU*). OKLAHOMA. LeFlore Co.: NW and below State Line 
Historical Site, N of Oklahoma highway 1,10 May 1986, Bryson 4307 (MICH*, MO, VDB, ctb). TENNES- 
SEE. Franklin Co.: Huntland, 3 May 1939, Svenson 10036 (TENN); 2.6 mi S of Huntland, 28 Apr 1972, 
Kral 45837 (MO,VDB), 24 May 1998, Naczi 7242 & Ford (DOV*™, MICH, TENN, USCH, VDB, VPI, WIN, ctb), 
Ford 98108 & Naczi (WIN). Wilson Co.: Cedars of Lebanon State Park, 28 May 1989, McNeilus 89-287 
(IBE, MICH*, MO, TENN, VDB, ctb). 


ACKNOWLEDGMENTS 


We thank Charles T. Bryson and Patrick J. Calie for valuable comments on the 
manuscript through their reviews. We also thank Charles T. Bryson, Anton A, 
Reznicek, Amy Trauth-Nare, Robert Kral, and John W. Thieret for assistance 
with field work. William A. McAvoy, Bruce A. Sorrie, and Thomas F Wieboldt 
generously allowed us to publish information on their noteworthy collections. 
Curators of BEREA, BLH, DHL, DOV, EKY, IBE, KNK, KY, MICH, MO, MT, MU, 
MUR, NA, NY, OS, PENN, PH, TENN, US, VDB, VPI, WIN, WKU; and Charles T. 
Bryson and Bruce A. Sorrie loaned specimens or assisted us during our visits. 
Theodore S. Cochrane supplied information on specimens of Carex jamesii at 
WIS. Thomas J. Reaume drew Carex timida and C jamesii. The Kentucky State 
Nature Preserves Commission permitted collection at some of its preserves. 
National Science Foundation Doctoral Dissertation Improvement Grant BSR- 
9001260 (to RFCN), Natural Sciences and Engineering Research Council of 
Canada operating and equipment grants (to BAF), grants from the Horace H. 
Rackham School of Graduate Studies of the University of Michigan (to RFCN), 
and a small grant from the Kentucky State Nature Preserves Commission (to 
RECN) provided financial support for portions of this work. 


REFERENCES 


Baiey, LH. 1885. Notes on Carex—lll. Bot. Gaz. 10:203-208. 

Case, PR. and C. WertH. 1995. The Trillium pusillum Michaux (Liliaceae) complex in Virginia. 
Il. lsozyme evidence. Castanea 60:15—29. 

Carey, J. 1848. Carex L. In: Gray, A., A Manual of the Botany of the Northern United States, 
from New England to Wisconsin and South to Ohio and Pennsylvania Inclusive. James 
Munroe & Co., Boston. Pp. 535-567. 

CATLING, P.M., A.A. Reznicex, and W.J. Crins. 1993. Carex juniperorum (Cyperaceae), a new spe- 
cies from northeastern North America, with a key to Carex sect. Phyllostachys. Syst. Bot 
18:496-501. 

Curist, H. 1885. Nouveau catalogue des Carex D'Europe. Bull. Soc. Roy. Bot. Belgique 24. 
l:10-20. 

Crark, R.C. 1967. Andrachne phyllanthoides (Nuttall) Muell.on the Cumberland Plateau of 
Alabama. Castanea 32:/3-74. 

Crins, W.J. 1990. Phylogenetic considerations below the sectional level in Carex. Canad. J. 
Bot. 68:1433-1440. 


Fernatp, M.L. 1950. Gray’s manual of botany, 8th ed. American Book Co., New York, NY. 

Foro, B.A., D.A.R. McQueen, R.F.C. Naczi, and A.A. Reznicek. 1998a. Allozyme variation and ge- 
netic relationships among species in the Carex willdenowii complex (Cyperaceae). Amer. 
J. Bot. 85:546-552. 

Foro, B.A., D.A.R. McQueen, J.R. Starr, and R.F.C. Naczi. 1998b. The impact of species-specific 
traits and phylogenetic related on allozyme diversity in Carex section Phyllostachys 

Cyperaceae). Pl. Syst. Evol. 212:13-29. 

Foro, B.A.and R.F.C.Naczi.2001.Genetic eek in i Carex jamesii complex (Cyperaceae: 
sect. Phyllostachyae), with insights into t tion and origin of the newly described 
species Carex timida. Sida 19885-8907, 

Foro, B.A.,J.R. Starr, D.A.R. McQueen, and R.F.C. Naczi. 1998c. Relationships among species in 
Carex section Phyllostachys (Cyperaceae) based on allozyme divergence. PI. Syst. Evol. 
212:31-51, 

Gieason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada, 2d ed. New York Botanical Garden, Bronx, NY. 

Houmoren, P.K., N.H. Houmeren, and L.C. Barnett [eds.]. 1990. Index herbariorum part |. The 
herbaria of the world, 8th ed. New York Botanical Garden, Bronx, NY. 

JOHNSON, G.P. 1988. Revision of Castanea sect. Balanocastanon (Fagaceae). Journal of the 
Arnold Arboretum 69:25-49. 

Lirtte, E.L. 1977. Atlas of United States trees, vol. 4. United States Government Printing 
Office, Washington, DC. 

McNetus, V.B. 1992. Carex ouachitana (Cyperaceae) new to Tennessee. Sida 15:154—155. 

Mackenzie, K.K. 1935. Cyperaceae - Cariceae. N. Amer. Fl. 18:169-478. 

Meyer, F.G. 1997. Magnolia. In: Flora of North America cha Committee, eds. Flora of 
North America North of Mexico, Vol.3. Magnoliophyta: Magnoliidae and Hamamelidae. 
Oxford Univ. Press, New York. Pp. 4-10. 

Naczi, R.A.C. 1999. Chromosome numbers of some eastern North American species of 
Carex and Eleocharis (Cyperaceae). Contr. Univ. Michigan Herb. 22:105-119. 

Naczi, R.F.C. and C.T. Bryson. 1990. Noteworthy records of Carex (Cyperaceae) from the 
southeastern United States. Bartonia 56:49-58 

Naczi, R.F.C. and B.A. Foro. 1998. Systematics of the Carex jamesii complex (section 
Phyllostachys, Cyperaceae). Suppl. to Amer. J. Bot. 85:147. 

Naczi, R.F.C., A.A. Reznicek, and B.A. Foro. 1998. Morphological, geographical, and ecological 
differentiation in the Carex willdenowii complex (Cyperaceae). Amer. J. Bot. 85:434- 
447 

Nixon, K.C. 1997. Castanea. In: Flora of North America Editorial Committee, eds. Flora of 
North America North of Mexico,Vol.3. Magnoliophyta: Magnoliidae and Hamamelidae. 
Oxford Univ. Press, New York. Pp. 439-442. 

Scora, R.W. 1967. Interspecific relationships in the genus Monarda (Labiatae). Univ. Calif. 
Publ. Bot.41:1-59. 

Smity, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, 2nd ed. By 
the author, Fayetteville, Ark. 


—= 


884 BRIT.ORG/SIDA 19(4) 


Starr, J.R., RJ. Bayer, and B.A. Foro. 1999. The phylogenetic position of Carex section 
Phyllostachys and its implications for phylogeny and subgeneric circumscription in 
Carex (Cyperaceae). Amer. J. Bot. 86:563-577. 

Starr, J.R. and B.A. Foro. 1995. Phylogeny and character evolution in Carex section 
Phyllostachys. Suppl. to Amer. J. Bot. 82:164-165. 

Starr, J.R.and B.A.Forb. 2001.The taxonomic and phylogenetic utility of vegetative anatomy 
and fruit epidermal silica bodies in Carex section Phyllostachys (Cyperaceae). Canad. J. 
Bot. 79:362-379. 

Starr, J.R., B.A. Foro, and RJ. Bayer. 1997. Testing phylogenetic hypotheses in Carex 
(Cyperaceae) using sequences from the internal transcribed spacer (ITS) region of 
nrDNA. Suppl. to Amer. J. Bot. 84:235. 

STRANGE, R.M. and B.M. Burr. 1997. Intraspecific phylogeography of North American high- 
land fishes: a test of the Pleistocene vicariance hypothesis. Evolution 51:885-897. 

Stuckey, R.L. 1974. Dr. Joseph C. Frank’s botanical work in Ohio, with a list of his 1835 type 
specimens of vascular plants in American herbaria. Castanea. 39:263-272. 

SworrorD, D.L. 2000. PAUP*: Phylogenetic Analysis Using Parsimony (* and Other Meth- 
ods), version 4.0. Sinauer, Sunderland, Massachusetts 


Torrey, J. 1836. Monograph of North American Cyperaceae. Ann. Lyceum Nat. Hist. New 
York 3:239-448, 

TUCKERMAN, E. 1843. Enumeratio methodica caricum quarundam. Schenectady, New York. 

Wickinson, L. 1989. SYSTAT: the system of statistics. SYSTAT, Evanston, IL. 


GENETIC DIVERSITY IN THE CAREX JAMESII COMPLEX 
(CYPERACEAE: SECT. PHYLLOSTACH YAE) WITH INSIGHTS 
INTO THE EVOLUTION AND ORIGIN OF THE 
NEWLY DESCRIBED SPECIES CAREX TIMIDA 


Bruce A. Ford Robert F.C. Naczi 


Pepantinent of botany Claude E. Phillips Herbarium 
j it Dept. of Agriculture & Natural Resources 
“Winnibed Manltoba- Delaware State University 
R3T 2N2, CANADA Dover, DE 19901-2277, U.S.A. 
bford@cc.umanitoba.ca raczi@dsc.edu 
ABSTRACT 
The Carex jamesii complex is composed of three species: C. jamesii, C. juniperorum, ang e timida. 
Despite the morphological similarity of C.jamesii and C.timida, tl linked 
ina cluster analysis using las igen uty, values. Rather, a ena) is Serouped with C.juniperorum. 
The close genetic similarity between a previous phyloge- 
netic study that showed that C. juniperorum aaa C. timida a are sister eae Carex timida popula- 
tions from the Ouachita Mountains of Arkansas are g nique and blur the distinctiveness 


between the latter species and C. juniperorum. aes a finding pat be a sampling artifact. Alterna- 
tively, the plants found in the Ouachita Mountains may be closest to the ancestor that gave rise to C. 
juniperorum and C. timida. A comparison of Sous Paylogenctic ane phenétic relationships pro- 


vides insights into the delineation of p j g C.timida 
at the species level. 
ABSTRACT 
El complejo Carex jamesii esta P s especies: C. jamesii, C.juniperorum, y C. allies A 
pesar de - semeanes mor koe ae Cjamesity y C timida, 
usando d genética. En su lugar G cna esta ea ae con 
f 4 iii q bt] 4 } 
eC unipeonn I g 
trod C timid ies | s poblaciones 


ty 
de eyes oid de | a romans eer de ila son n geneticamente singles y enturbia la 
cistncien entre la uliana especie y C.ju neu Tal hall pued 


I las montanas Ouachita pueden ser mas cercanas 
a ac. Pee ¥C. pea Una comparacion de las relaciones genéticas, 
ae y fenéticas proporcionan una mejor comprension de la delineacion de los taxa 
infraespecificos y la justificacion para el reconocimiento de C. timida a nivel especifico. 


INTRODUCTION 
Carex L. section Phyllostachyae Tuck. ex Kiik. is a well-defined group of 10 species 
(C. backii Boott; C. basiantha Steud.; C_jamesii Schwein.; C_juniperorum Catling, 
Reznicek, & Crins; C. latebracteata Waterfall: C. saximontana Mack. C. superata 


SIDA 19(4): 885 — 897. 2001 


886 BRIT.ORG/SIDA 19(4 


= 


Naczi, Reznicek, & B.A. Ford; C. timida Naczi & B.A. Ford; C. willdenowii Willd: 
C. sp. nov. Saarela & B.A. Ford) that is endemic to forested and semi-forested 
habitats in North America. This section has been the subject of considerable 
systematic research (Catling et al. 1993; Ford et al. 1998a, 1998b, 1998c; Naczi et 
al. 1998; Starr et al. 1999: Naczi & Ford 2001: Saarela and Ford in press) with 
new insights having been gained not only into the taxonomy, phylogeny, and 
genetic structure of this section, but the genus Carex as a whole. One of the 
most unexpected findings of our research has been the discovery of a propor- 
tionately large number of undetected or undescribed species. These new spe- 
cies turn out to be members of widespread species complexes that are masquer- 
ading undera single name. Undetected species usually become apparent when 
plants from the northeastern and central United States and adjacent Canada 
are compared with specimens from the southeastern United States or western 
North America. This trend was evident in our earlier study of the C. willdenowii 
complex (C. willdenowiis. str; C. basiantha; C. superata) (Ford et al. 1998c; Naczi 
et al. 1998) and most recently in our investigation of C. jamesiis. lat. (C. jamesii 
s. str; C.juniperorum, C. timida) (Naczi & Ford 2001) and C. backii (C. backiis. 
str; C. saximontana; C. sp. nov.) (Saarela & Ford in press). 

The C.jamesii complex isa well-defined, monophyletic assemblage of taxa 
(Ford et al. 1998b; Starr et al. 1999) distinguished from other species in section 
Phyllostachyae by a combination of filiform stigmas, pistillate scales that do 
not conceal the perigynium, and perigynium bodies that are abruptly con- 
tracted into a beak. Molecular studies indicate that all three species share iden- 
tical ITS sequences (Starr et al. 1999; Starr pers. comm.). Morphologically, C. 
juniperorumis the most divergent member of this complex being distinguished 
by its numerous perigynia (4-9 per inflorescence), short culms (<1/3 the length 
of the leaves), and lack of hyaline margins on the pistillate scales. Carex jamesii 
and C. timida are characterized by their relatively few perigynia (1-3 per inflo- 
rescence), elongate culms (subequal to the length of the leaves), and pistillate 
scales with conspicuously hyaline margins. Both species are superficially simi- 
lar but can be readily distinguished by differences in cataphyll epidermal cell 
morphology, sheath color, and proximal staminate scale length. 

Isozyme analysis has provided important ancillary data for systematic 
studies of Carex and in particular for recent taxonomic investigations of spe- 
cies complexes in section Phyllostachyae (Ford et al. 1998a, 1998b, 1998c). The 
intent of this study was to: 1) assess the taxonomic status of C. timida using 
isozyme data; 2) determine the degree of genetic divergence within and between 
species in the C. jamesii complex; and 3) compare the phylogenetic inferences 
from isozyme data to the evolutionary hypothesis proposed by Naczi and Ford 
(2001). 


FORD AND NACZI, GENETIC 887 


MATERIALS AND METHODS 


A total of 649 individuals were collected from 26 populations (16 of C. jamesii, 
4 of C juniperorum, and 6 of C.timida) (Table 1). Our study included all C.jamesii 
and C. juniperorum populations examined by Ford et al. (1998a, 1998b) plus 
additional populations of C. jamesii and C. timida that were collected after the 
completion of these studies. The methodology for field sampling and enzyme 
analysis follows that of Ford et al. (1998c). Eleven enzymes coded by 15 inter- 
pretable, putative loci were included in this study. The 15 loci and their associ- 
ated alleles were: aspartate aminotransferase, AAT 1 (a); diaphorase DIA-1 (a to 
e), DIA-2 (a); glucose-6-phosphate isomerase, GPI-2 (a to f); leucine aminopep- 
tidase, LAP-1 (a to c); malate dehydrogenase, MDH-1 (a to b), MDH-2 (a to c); 
menadione reductase, MDR (a to b); peroxidase, PER-2 (a); phosphoglucomu- 
tase, PGM-1 (a toe), PGM-2 (a toe); shikimate dehydrogenase, SKD (a toc, allele 
cisanullallele observed in population 20 of C. timida); superoxide dismuta 
SOD (a to b); and triose-phosphate isomerase, TPI-l (a to d), TPI-2(a toc). Allele 
frequencies, Nei’s unbiased genetic identities (I) (Nei 1978), and an UPGMA 
(unweighted pair-group method) phenogram were calculated using BIOSYS-1 
(Swofford & Selander 1981). Total genetic diversity for each species (H7), aver- 
age diversity within (Hs) and among populations (Dsr), and the coefficient of 
genetic differentiation (Gsr) were calculated using Nei and Chesser’s (1983) 
procedure, unbiased for sample size, using GENESTATPC v. 2.1 (Lewis & 
Whitkus 1989). These analyses included both monomorphic and polymorphic 
loci in their calculations. 


RESULTS 


A total of 15 putative loci were surveyed in this study with all loci, expect AAF1, 
DIA-2,and PER-2 being polymorphic in one or more populations. Carex jamesii 
had 12 polymorphic loci, while 9 variable loci were found in C.juniperorum and 
C. timida. Carex jamesii also had the greatest allelic diversity with 42 alleles 
identified, while 30 and 31 different alleles were observed in C. juniperorum 
and C. timida, respectively. With the exception of unique alleles for PGM-1, the 
allozymes found in C. timida and C. juniperorum were a subset of those found 
in C. jamesii (Table 2). 

Genetic variability statistics fell within the range previously reported for 
C.jamesii and C.juniperorum (Ford et al. 1998a) (Table 3). The mean number of 
alleles per locus (K) ranged from 1.2 in C. juniperorum and C. timida (popula- 
tion 52, Hastings Co., Ontario and population 53, Monroe Co., Kentucky, respec- 
tively) to 2.1 in the Campbell Co., Kentucky population of C. jamesii (popula- 
tion 2). The percentage of polymorphic loci (P) was variable and ranged from 
20.0 in the Monroe Co,, Kentucky population of C. timida (population 53) to 


888 BRIT.ORG/SIDA 19(4) 


Tasie 1.Collection data for populations of the Carex jamesii complex. Population codes are referred 
to parenthetically following each citation. Vouchers are deposited in WIN except where noted. 


Carex jamesii Schweinitz 


CANADA. ONTARIO. Essex Co.: Anderdon Twp.,5 km NE of Amherstburg, 22 May 1994, Ball 940526 
(22). Niagara Regional Mun.: Louth Twp., Twenty Mile Creek, Jordan, 13 Jun 1979, Ball 79039 (PWB 
in TRTE (31). Waterloo Co.: Wilmot Twp.,8 km W of New Dundee on the Nith River, 3 Jun 1982 Bal 
82074 (PWB in TRTE) (32). U.S.A. ARKANSAS. Franklin Co.: ca. 1 mi N of Cecil, Citadel Bluff Army 
Corps of Engineers Park, 19 May 1994, Naczi 3923 & Ford (21).Newton Co.: ca. 3 mi NE of Boxley, Lost 
Valley Recreation Area of Buffalo National River, 19 May 1994, Naczi 3917 & Ford (13). Scott Co.: ca. 2 
mi N of ¥ City, W of route 71 and S of Fourche La Fave River, 20 May 1994, Naczi 3939 & Ford (18). 
INDIANA. Grant Co.: Taylor University Arboretum, SW edge of Upland, 17 May 1994, Rothrock 3255 
(9); Stellers Road, 1.3 mi N of Matthews, 17 May 1994, Rothrock 3254 (8). KENTUCKY. Boone Co.:3 air 

mi S of Petersburg, ca.0.3 mi W of route 20 along S side of Woolper Creek, 12 Jun 1994, Naczi 4096 
(39), Campbell Co.: Highland Heights, 10 May 1994, Naczi 3826 (2); Silver Grove, N of route 8, flood- 
plain of Ohio River, opposite St. Anne’s Convent, 12 May 1995, Naczi 4575 & Ganss (51). Mason Co.: 
ca. 2 air mi W of Dover, along S side of route 8,29 May 1994, Naczi 4027 & Flynn (33); ca. 2 air mi W of 
Dover, along S side of route 8, 29 May 1994, Naczi 4028 & Flynn (34). MISSISSIPPI. DeSoto Co.: ca. 2 
mi N of Walls, along E side of route 61,25 May 1994, Naczi 4026 et al. (24). OHIO. Montgomery Co., 
SW of Farmersville, E side of Anthony Road, 0.3 mi SW of its junction with Manning Road, 29 May 
1998, Ford 98152 & Naczi (57). VIRGINIA. Bath Co.: ca.0.4 mi S of Healing Springs, along W side of 
route 220, 23 Jun 1994, Naczi 4482 & Thieret (41). 


Carex juniperorum Catling, Reznicek, & Crins 


CANADA. ONTARIO. Hastings Co.: Tyendinaga Twp.,E side of Salmon River,ca. 15 km W of Napanee, 
24 Jul 1995, Ford 9566 et al. (52).U.S.A. KENTUCKY. Bath Co.: ca.5 air mi ESE of Owingsville, 16 May 
1994, Naczi 3890 (5). Lewis Co.: ca. 3.5 air mi ESE of Trinity, 5 May 1994, Naczi 3808 et al. (1). OHIO, 
Adams Co.: ca. 3 air mi NE of Peebles, 16 May 1994, Naczi 3878 (7) 


Carex timida Naczi & B.A. Ford 


U.S.A. ARKANSAS. Polk Co.: SW of town of Rich Mountain on summit of Rich Mountain, Queen 
Wilhelmina State Park, along N side of route 88, in vicinity of trailhead of Spring Trail, 20 May 1994, 
Naczi 3940 & Ford (20);ca.8 mi E of Vandervoort, N of route 246 and E of Cossatot River, 20 May 1994, 
Naczi 3949 & Ford (16). KENTUCKY. Monroe Co.: SE of Tomkinsville, along the W side of route 216,6 
road mi E of its junction with route 163, along McFarland Creek, 23 May 1998, Ford 98100 & Naczi 
(53). Rowan Co.: ca.6.5 air mi S of center of Morehead, ca.0.25 mi down slope from W side of route 
1274, upslope from Sugar Camp Branch, ca. 1.3 road mi N of junction of routes 1274 and 801, 28 
May 1998, Ford 98145 & Naczi (55). OHIO. Montgomery Co.: SW of Farmersville, E side of Anthony 
Road, 0.3 mi SW of its junction with Manning Road, 29 May 1998, Ford 98153 & Naczi (56). TENNES- 
SEE. Franklin Co.: S of Huntland, along E side of route 97, 2.6 road mi S of its junction with route 
122, 24 May 1998, Ford 98108 & Naczi (54). 


73.3 in the Grant Co., Indiana population of C. jamesii (population 8). The aver- 
age observed heterozygosity within populations (Hops) ranged from 0.137 in the 
Bath Co, Virginia population of C.jamesii (population 41) to 0.336 in the Grant 
Co, Indiana population of this same species (population 8). The expected het- 
erozygosity in each population based upon Hardy-Weinberg expectations 
(Hexp) was less than that observed in each population with values ranging from 


FORD AND NACZI, GENETIC 889 


Taste 2. Allozyme frequencies for polymorphic loci in Carex jamesii, C. juniperorum, and C. timida as 


averages for each species (except where noted). N = number of individuals used in the calculation 
of averages 
Locus Allele C. jamesii Cjuniperorum  C.timida C. timida C. timida 
(N = 383) (N= 112) Pop.53-56 Pop. 16 Pop. 20 
(N = 98) (N = 24) (N = 32) 
DIA-1 A 0.360 0.509 0.500 0.500 0.484 
B 0.026 - - - 0.016 
C 0.391 0.246 0.398 0.500 0.484 
D 0.001 0.009 - - - 
E 0.221 0.237 0.102 - 0.016 
GPI-2 A 0.297 = - - 
B 0.124 0.500 0.500 0.500 0.500 
Ss 0.159 = = = - 
D 0.409 = = 
E 0.009 0.500 0.372 0.500 0.500 
F ~ - 0.128 
LAP-1 A 0.008 0.019 
B 0.987 0.972 1.000 1.000 1.000 
C 0.005 0.009 
MDH-1 A 0.537 0.960 .000 0.979 0.984 
B 0.463 0.040 0.021 0.0.16 
MDH-2 A 0.026 0.054 .000 000 0.938 
B 0.026 0.009 - - 0.03 
é 0.948 0.938 - ~ 0.031 
MDR A 0.057 - = 
B 0.943 1.000 1.000 000 000 
PGM-1 A 0.020 - - - - 
B 0.095 - - = 
C 0.885 - 1.000 - 
D - 1.000 
E - 1.000 = 1.000 
PGM-2 A 0.054 0.063 - = _ 
B 0.534 = 
C 0.003 0.938 = 0.625 1.000 
D 0.363 
E 0.047 - 1.000 0.375 7 
SOD A 0.979 0.977 1.000 1.000 000 
B 0.021 0.023 - 
SKD A 0.005 0.194 1.000 - 
B 0.995 .000 0.806 = 
C 


(null allele) — - - - 1.000 


890 BRIT.ORG/SIDA 19(4) 


Taste 2. cont. 


Locus Allele C. jamesii Cjuniperorum  C. timida C. timida C. timida 
(N = 383) (N = 112) Pop. 53-56 Pop. 16 Pop. 20 
(N = 98) (N= 24) (N = 32) 
TPI-1 A 0.009 0.004 0.372 0.167 0.469 
B 0.337 0.991 0.628 0.833 0.469 
C 0.631 0.004 - ; - 
D 0.023 - - - - 
TPI-2 A 0.004 0.504 0.372 0.500 0.484 
B 0.449 0.004 0.128 = 0.016 
C 0.547 0.491 0.500 0.500 0.500 


0.082 in the Bath Co,, Virginia population of C.jamesii (population 41) to 0.261 
in the Mason Co,, Kentucky population of this same species (population 34). 

Gene diversity statistics (Table 4) indicated that C. juniperorum had the 
lowest total gene diversity (Hy) (0.135), while that for C. jamesii (0.265) and C. 
timida (0.247) was almost twi great. A similar trend was seen within-popu- 
lations (Hs), with the lowest value found in C_juniperorum while much higher 
values were observed in C. jamesii (0.188) and C. timida (0.141). 

The gene diversity among populations (Dsr) and coefficient of genetic dif- 
ferentiation (Gs7) were also variable with extremely low values found in C. 
juniperorum (Dst = 0.007, Gsr = 0.049), while relatively high values were asso- 
ciated with C jamesii (Dst= 0.077, Gsy = 0.290) and C. timida (Dst = 0.106, Gsr 
= 0.429). Taken together these numbers indicate that species within the C. 
jamesii complex harbor as little as 57.1% (C. timida) to as much as 95.1% (C. 
juniperorum) of their genetic diversity within populations. 

Intra-specific genetic identity values were variable, and in some instances 
surprisingly low (Table 5). Average values ranged from 0.990 for C.juniperorum 
to 0.852 in C. timida. The low value found in this latter species was largely the 
result of the presence of unique alleles for the isozymes PGM-l, PGM-2, and 
SKD in the Arkansas populations (populations 16 and 20) of this taxon (Table 
5). No activity was observed for SKD in population 20 suggesting that these 
plants may have lost the ability to express this enzyme phenot 

Inter-specific ic identity values were lower than those observed within 
species and ranged from 0.655(C._jamesiiand C.timida) to 0.760(C.juniperorum 
and C. timida) (Table 5). These lower values can be attributed to the presence of 
diagnostic or high frequency alleles in all species. Alleles for the isozymes MDH- 
2, PGM-1,and PGM-2 helped to differentiate C._jamesii from C. timida and/or C. 
juniperorum (Table 2). The high genetic identity between C. juniperorum and 
C.timida was the result of the presence of similar alleles for PGM-l and/or PGM- 
2 in Arkansas populations of C. timida and populations of C. juniperorum. All 


FORD AND NACZI, GENETIC 891 


Tas.e 3. Genetic variability in 26 populations of the Carex jamesii complex: sample size (N), mean 
number of alleles per locus + SE (k), percentage of polymorphic loci + SE (P) (a locus is considered 
olymorphic if the frequency of the most common allele does not exceed 0.99), observed het- 


erozygosity + SE (H, 


bs 


), expected heterozygosity + SE (H.,_) (Unbiased estimate Nei [1978]). 


Kp 


Population # N k P Hoos Hexp 
Carex jamesil 
2 38 2.1+0.3 66.7 0.211 + 0.095 0.195 + 0.061 
8 31 1.9+0.2 733 0.336 + 0.123 0.250 + 0.069 
9 30 1.9+0.3 53.3 0.329 +0.121 0.253 + 0.073 
13 30 1.7 +40.2 533 0.264 + 0.116 0.168 + 0.058 
18 26 15+0.2 33:3 0.262 + 0.116 0.149 + 0.060 
21 34 1.5+0.2 40.0 0.259 + 0.114 0.147 + 0.060 
22 13 1.34011 3312 0.333 + 0.126 0.173 + 0.066 
24 31 1.9+0.3 46,7 0.252 +0.111 0.181 + 0.065 
31 24 1.9+40.3 53:3 0.331 40.125 0.198 + 0.066 
32 19 1.7+0.2 60.0 0.270 + 0.114 0.163 + 0.058 
33 20 1.7 +0.2 53,3 0.333 + 0.124 0.217 + 0.064 
34 9 18+ 0.2 60.0 0.341 40.118 0.261 + 0.069 
39 18 1440.2 33.3 0.333 + 0.126 0.179 + 0.068 
4] 34 1440.2 26.7 0.137 + 0.090 0.082 + 0.082 
51 12 14+02 33:3 0.333 + 0.126 0.177 + 0.067 
57 14 16+0.2 40.0 0.310+0.118 0.216 + 0.072 
Carex juniperorum 
] 34 1.7+0.2 46.7 0.214 + 0.106 0.164 + 0.058 
b) 27 1.7+0.2 53.3 0.208 + 0.103 0.131 + 0.052 
7 29 14+0.2 33.3 0.205 + 0.105 0.115 + 0.054 
52 22 12+0.1 20.0 0.200 + 0.107 0.102 + 0.055 
Carex timida 
16 24 1.4+0.1 40.0 0.231 +0.105 0.156 + 0.059 
20 32 17403 40.0 0.265 + 0.116 0.153 + 0.062 
53 25 1.2+0.1 20.0 0.200 + 0.107 0.102 + 0.055 
54 20 1.3+0.1 26,7 0.267 + 0.118 0.137 + 0.061 
55 26 1.3081 26./ 0.267 + 0.118 0.136 + 0.060 
56 27 lat 0.1 33.3 0.267 + 0.118 0.164 + 0.062 


populations of C. timida could be distinguished from C. juniperorum by the 
presence of the allele MDH-2a (Table 2). 

A cluster analysis of populations using Nei’s (1978) unbiased genetic iden- 
tity values indicated the presence of two distinct groups, one corresponding to 
C. jamesii and a second to C. juniperorum/timida. Syntopic populations of C. 
jamesiiand C. timida (population 56 and 57) were separated with no interme- 
diates being detected. Within the second cluster, C. juniperorum and C. timida 
formed indistinct groups owing to the allelic similarity of the two Arkansas 
populations of C. timida with C. juniperorum (Fig. D). 


892 BRIT.ORG/SIDA 19(4) 


Taste 4. Gene diversity statistics for the Carex jamesii complex. H, = total gene diversity, H.= within 
population gene diversity, D., = gene diversity among populations, G. = coefficient of genetic dif- 
ferentiation 


Species Hy Hs Dsy Gsr 

Carex jamesii 0.265 0.188 0.077 0.290 
Carex juniperorum 0.135 0.128 0.007 0.049 
Carex timida 0.247 0.141 0.106 0.429 


Taste 5. Matrix of genetic identity coefficients (range) for all pairwise comparisons of sampled popu- 
lations (N) of the Carex jamesii complex 


Species N C jamesii C juniperorum C.timida 

Carex jamesil 16 0.901 
ie a 1.000) 

Carex juniperorum 4 0.990 
A an 0.834) (0.985-1.000) 

Carex timida 6 0.655 0.760 0.852 
(0.545-0.779) (0.695-0.845) (0.717-0.984) 


DISCUSSION 


Taxonomic and Phylogenetic Implications 

Our isozyme study provides allelic data that supports the recognition of three 
species in the C. jamesii complex (cf. Naczi & Ford 2001). Each species is distin- 
guished by at least one unique or high frequency allele, and with one exception 
(see below), each forms a distinctive group in the cluster analysis of genetic 
identity values. Despite the age coene pallanity, of ame and C. timida, 
these two species were separated in the cluster analysis. Even when 
these species occur in mixed i pe et (e.g., populations 56 and 57), no inter- 
mediates were detected. 

Phylogenetic relationships within section Phyllostachyae have been ex- 
plored in a number of papers, with Naczi and Ford (2001) having investigated 
the relationship of the newly described C. timida to other members of the sec- 
tion. Despite the morphological similarity of C.jamesii and C.timida, this study 
suggests that C. timida and C. juniperorumare sister species and that C. jamesii 
is basal to this clade. Our genetic distance analysis of isozyme data substanti- 
ates this hypothesis. 

he discovery of genetically unique populations of C. timida from the 
Ouachita Mountains of Arkansas, which blur the distinctiveness between the 
latter species and C juniperorum, was surprising, especially since morphologi- 
cally these plants have been shown to be C. timida (Naczi and Ford 2001). There 


FORD AND NACZI, GENETIC 893 


nsauel 


mA. ats 
em. 
ma ee 
2. 
wrmoleduntl 


20 - Ark 
16 - Ark 
53 - Ky S| 
54 - Tenn o 
55 - Ky” 
56 - Ohio 

.60 .70 .80 .90 1.00 

Genetic Identity 
Fic. 1. Phenog f 26 populations of the Carex jamesii complex using Nei’s (1978) unbiased g ty values 
and UPGMA cluster analysis. * = type locality for C. timida. Copt i lati fficient = 0.915 


are, however, a number of possible explanations for this finding. One possibil- 
ity is that the lack of distinct groups is a sampling artifact. If more loci had 
been surveyed, other diagnostic alleles, such as MDH-2a, might have been found 


894 BRIT.ORG/SIDA 19(4) 


resulting in the formation of more distinctive clusters. Another possibility is that 
the C. timida populations sampled from the Ouachita Mountains (especially 
population 20) are closest to the ancestor that gave rise to C.juniperorum and 
C.timida, accounting for their unique positions in the cluster analysis. Alterna- 
tively, C. juniperorum could have arisen from a population, or populations, of C. 
timida similar to those found in western Arkansas. The Ouachita Mountains 
are a recognized glacial refugium and a center of endemism (Robinson & Allen 
1995), a number of narrowly distributed plant and animal species occur in this 
region (Kral & Bates 1991; Robinson & Allen 1995). Many species that found 
refuge here are thought to have moved northward following the events of Pleis- 
tocene glaciation (Robinson & Allen 1995). In some instances, these more north- 
erly populations became isolated from populations in the south resulting in 
the formation of new species (Ross & Ricker 1971; Robinson & Allen 1995). This 
phenomenon could account for the origin of C juniperorum and the occurrence 
of genetically unique populations of C. timida in the Ouachita Mountains. 


Genetic Variability and Diversity 

Table 3 and 4 indicate that C.juniperorum possesses about half the genetic vari- 
ability and diversity found in C.timida and C. jamesii. This pattern is similar to 
that found in our study of C. willdenowii s. lat., where C. superata was thought 
to possess half of the variation/diversity found in the other species of this com- 
plex due to its very short culms (could restrict pollen and seed movement) and 
restricted distribution (limited gene flow, selection due to environmental ho- 
mogeneity (Ford et al. 1998c). Like C. superata, C. juniperorum has extremely 
short culms, with inflorescences that are crowed in the base of the plant. Fur- 
thermore, this species is rare and occurs in disjunct regions in southern Ontario, 
Kentucky, Ohio, and Virginia (Naczi and Ford 2001). The combined evidence 
suggests that factors similar to those operating in C. superata may be influenc- 
ing the genetic structure of C. juniperorum. 

At the opposite end of the spectrum is C. jamesii. This is the widest rang- 
ing species in the C. jamesii complex and is sympatric with both C. timida and 
C. juniperorum. Carex jamesii has the highest number of polymorphic loci (12 
out of 15), the greatest number of alleles, and the highest genetic variability 
and diversity values (Hy and Hs) for any species in this clade. In addition, with 
the exception of unique alleles for PGM-1, the allozymes found in C. timida and 
C. juniperorumare a subset of those found in C. jamesii. The widespread nature 
of C. jamesii, and its adaptation to a variety of climatic and ecological condi- 
tions, may be factors contributing to the high levels of genetic diversity and 
variability found in this species. Being only one node away from the ancestor 
that gave rise to the C. jamesii clade (cf. Naczi & Ford 2001), it is also possible 
that C. jamesii retains much of the variation found in the ancestor to this group 
accounting for this species extensive allelic diversity. 


FORD AND NACZI, GENETIC 895 


Insights into the Delineation of Infraspecific Taxa and Justification for Recog- 
nizing C. timida as a Distinct Species 

A comparison of the evolutionary hypothesis proposed in this paper with the 
superficial similarity found between C. timida and C. jamesii allows us to explore 
issues surrounding the recognition of infraspecific taxa and justification for 
recognizing C. timida at the species level. Infraspecific categories are frequently 
used by taxonomists as a means of recognizing poorly differentiated taxa or 
taxa distinguished by seemingly minor morphological differences. Current phy- 
logenetic methods may not be appropriate for determining relationships at this 
level since these relationships are not necessarily hierarchical and the charac- 
ters used to define taxa are not always discrete. This fact, along with problems 
associated with outgroup selection, has led most caricologists to develop in- 
fraspecific classifications using phenetic methods (e.g., Murray 1969; Reznicek 
& Ball 1980; Crins & Ball 1983; Reznicek 1987; Standley 1985; Crins & Ball 1989a, 
1989b; Ball & Zoladz 1994; Dunlop & Crow 1999). One might wish to consider 
C.timida as a subspecies of C. jamesii because of the close morphological simi- 
larity of these two taxa. Indeed, an evaluation of the results of our phenetic 
study might have made this a tenable conclusion (cf. Naczi & Ford 2001). How- 
ever, when the results of our phylogenetic and genetic research are considered 
it is clear that the recognition of C. timida as a subspecies of C. jamesii would 
have created a clade composed entirely of artificial taxa. While infraspecific 
relationships are not necessarily hierarchical, infraspecific classifications do 
represent explicit phylogenetic hypotheses. This study shows that genetic and 
phylogenetic divergence is not necessarily correlated with striking morphologi- 
cal differences: morphologically similar taxa are not necessarily closely related. 
We submit that infraspecific classifications based entirely on grouping mor- 
phologically similar taxa can lead to the recognition of artificial species. Fur- 
ther, we suggest that if taxa possess clear-cut differences, no matter how nar- 
row, it is best to recognize these taxa as distinct species. Infraspecific taxa should 
exhibit some degree of intergradation thus making the identification of a sig- 
nificant number of individuals imy le even under the most perfect circum- 
stances. Using this criterion, C. timida is best recognized as a distinct species. 


ACKNOWLEDGMENTS 


We greatly appreciate the assistance of D.A. Ross McQueen and Julian R. Starr 
in our earlier studies of Carex section Phyllostachyae. A special thanks to Anton 
A. Reznicek for sharing his thoughts on the recognition of infraspecific taxa. We 
are also grateful to Patrick Calie and Charles Bryson for their careful review of 
the manuscript. Peter Ball, Wasyl Bakowsky, Charles Bryson, Mike Oldham, Paul 
Rothrock, and Donald Sutherland supplied key population samples and/or 
assisted with various aspects of our fieldwork. Piggly Wiggly provided victuals 


896 BRIT.ORG/SIDA 19(4) 


and inspiration. This research was supported by a Natural Sciences and Engi- 
neering Research Council of Canada operating and equipment grants to B.A.F 


REFERENCES 


Batt, PW.and M. Zotapz. 1994. The taxonomy of Carex petricosa (Cyperaceae) and related 
species in North America. Rhodora 96:295-310 

CATLING, P.M. A.A. Reznicek, and W.J.CrINS. 1993. Carex juniperorum (Cyperaceae), a new spe- 
cies from northeastern North America, with a key to Carex sect. Phyllostachys. Syst. Bot 
18:496-501, 

Crins, W.J. and P.W. Batt. 1983 The taxonomy of the Carex pensylvanica complex 
(Cyperaceae) in North America. Canad. J. Bot.61:1692-1717. 

Crins, W.J.and PW. Bait. 1989a. Taxonomy of the Carex flava complex (Cyperaceae) in North 
America and northern Eurasia. |. Numerical ta~onomy and character analysis. Canad. J. 
Bot. 67:1023-1047 

Crins, W.J.and PW. Bait. 1989b. Taxonomy of the Carex flava complex (Cyperaceae) in North 
America and northern Eurasia. Il. Taxonomic treatment. Canad. J. Bot.67:1048-1065 

Duntor, D.A.and G.E. Crow. 1999. The taxonomy of Carex section Scirpinae (Cyperaceae). 
Rhodora 101:163-199. 

Foro, B.A., D.A.R. McQueen, J.R., Starr, and R.F.C. Naczi. 1998a. The impact of species-specific 
traits and phylogenetic relatedness on allozyme diversity in Carex sect. Phyllostachys 
(Cyperaceae). Pl. Syst. & Evol. 212:13-29. 

Foro, B.A., J.R., Starr, D.A.R. McQueen, and R.F.C. Naczi. 1998b. Relationships ee species 
in Carex sect. Phyllostachys (Cyperaceae) based on allozyme diverg PI. Syst.& Evol. 
212:31-51 

Foro, B.A., D.A.R. McQueen, R.FC. Naczi, and A.A. Reznicek. 1998c. Allozyme variation and ge- 
netic relationships among species in the Carex willdenowii complex (Cyperaceae). Amer. 
J.Bot.85:546-552. 

Krat, R.and V. Bates. 1991.A new species of Hydrophyllum from the Ouachita Mountains of 
Arkansas. Novon 1:60-66. 

Lewis, P.and R.Wuirtkus. 1989. GENESTAT-PC V. 2.1. Department of Botany, Ohio State Uni- 
versity, Columbus. 

Murray, D.F. 1969. Taxonomy of Carex sect. Atratae (Cyperaceae) in the southern Rocky 
Mountains. Brittonia 21:55-76 


Naczi, RFC, A.A. Reznicek, and B.A. Forp. 1998. Morphological, geographical, and ecological 
differentiation in the Carex willdenowii complex (Cyperaceae). Amer. J. Bot. 85:434- 
447. 

Naczi, R.EC. and B.A. Forp. 2001. Systematics of the Carex jamesii complex (Cyperaceae: 
sect. Phyllostachyae). Sida 19:853-884. 

Ne, M. 1978. Estimation of average heterozygosity and genetic distance from a small 
number of individuals. Genetics 89:583-590,. 

Ne, M. and R.K. CuHesser. 1983. Estimation of fixation indices and gene diversities. Ann. 
Human Genet. 47:253-259. 


FORD AND NACZI, GENETIC 897 


Reznicek, A.A.and PW. Batt. 1980.The taxonomy of Carex section Stellulatae in North America 
north of Mexico. Contr. Univ. Michigan Herb. 14:153-203. 

Reznicek, A.A. 1987. Carex sect. Stellulatae (Cyperaceae) in the neotropics.Contr. Univ. Michi- 
gan Herb. 16:201-210. 

Rosinson, H.W. and R.T. Auten. 1995. Only in Arkansas: a study of the endemic plants and 
animals of the state. The University of Arkansas Press, Fayetteville. 

Ross, H.H. and W.E. Ricker. 1971. The classification, evolution, and dispersal of the winter 
stonefly genus Allocapnia. Illinois Biol. Monogr 45:1—-166. 

Saarela, JM. and B.A. Foro. In press. Taxonomy of the Carex backii complex (section 
Phyllostachyae, Cyperaceae). Syst. Bot 

Stanb.ey, L.A. 1985. Systematics of the Acutae group of Carex (Cyperaceae) in the Pacific 
Northwest. Syst. Bot. Monogr. 7:1-106. 

Starr, J.R., R.J. Bayer, and B.A. Foro. 1999. The phylogenetic position of Carex section 
Phyllostachys and its implications for phylogeny and subgeneric circumscription in 
Carex (Cyperaceae). Amer. J. Bot. 86:563-577 

Sworroro, D.L.and R.B. SeLanver. 1981.Biosys-1:computer program for the analysis of allelic 
variation in genetics. User's manual. University of Illinois, Urbana. 


a 


898 BRIT.ORG/SIDA 19(4) 


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essays detailing the history o I John James Audubon 


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The book is an exceptional tribute to Audubon. Pl Ginsburg, Botanical Research Institute of 
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SIDA 19(4): 898. 2001 


STENOTIS (RUBIACEAE), A NEW SEGREGATE GENUS 
FROM BAJA CALIFORNIA, MEXICO 


Edward E. Terrell! 


arch Collabora 
Department of Systematic hae Botany 


Smithsonian Institution 


L 
Natural PHOTO 


Wee Dc 20560, U.S.A. 


ABSTRACT 
seven former Hedyotis (Rubiaceae) species native to Baja Cantor, pMERICO, are Soles and placed 
nanew genus, Stenotis, on the basis of seed and other hara 
oi and recent DNA evidence. Keys descriptions aaeibacinine mae ided. Two of 


r 
the species are annual herbs (Stenotis arenaria, S. asperuloides), and five are perennial woody herbs 
or shrublets (S. australis, S. brevipes, S. gracilenta, S. mucronata, and S. peninsularis). Two varieties are 


recognized in S. asperuloides. 


RESUMEN 


Se revisan siete pepe de Hedolts ee) Semis de Baja Caltornis, MEE. bi se colocan 


Stenotis. I y enotros 
caracteres aorislorices enel numero de eroniecanias y en evidencias recientes derivadas del ADN. 
Se presentan claves taxonomicas, descripciones, distribuciones y sinonimos. Dos de estas especies 
son hierbas anuales (Stenotis arenaria y S.asperuloides), y cinco son plantas perennes de tallo lenoso 
0 pequenios arbustos (S. australis, S. brevipes, S.gracilenta, S. mucronata, y S. peninsularis). Se reconocen 
dos variedades dentro de la especie S. asperuloides 


INTRODUCTION 


Early explorations in Baja California, Mexico, by TS. Brandegee, I'M. Johnston, 
and others from 1844 to 1924 turned up eight new species assigned to Houstonia 
or Hedyotis (Hedyotideae; Rubiaceae). Seven of these species (excluding Hedyotis 
vegrandis WH. Lewis pCO prostrata Brandegee] of uncertain affinity), 
sometimes termed the F | wo annual herbs and 
five perennial herbs or shrublets. An additional species, Hed greenei A. Gray, 
occurs in Arizona and is closely related to H. arenaria, but is excluded from the 
present study pending further study of its relationships. These species differ in 
pera) from the approximately six so-called Hedyotis species occurring 
in Mexi utside of Baja California, whose relationships require further study. 

Previous work on Houstonia (Terrell 1996) and other related genera showed 
that two taxonomically meaningful characters are seed morphology and chro- 
mosome number. The seeds (Fig. 1) of the seven Bajan species are generally el- 
lipsoid with a more or less centric punctiform hilum, but some species have a 


lyot ronatd group, i 


‘Address for correspondence: 14001 Wildwood Dr, Silver Spring, MD 20905, U.S.A. 
SIDA 19(4): 899 — 911. 2001 


900 BRIT.ORG/SIDA 19(4) 


Fic. 1 Cc 7 £o. ~ . i. y 4 g I i Cc | L | fa let L J cn 

i howi ichil A. Stenoti , Wiggins 14424 (TEX).B. Sena usta Carter et al. 2337 
(GH), C. Stenotis peu, Carter 200e bee D. Stenois arenaria, Brandegee s.n., 10/14/1893 (NY, US). E. Stenotis 
arenaria, same collectio Stenotis arenaria, Lewis 5341 i 


conspicuous ventral hilar ridge or vary somewhat in being irregularly and ob- 
tusely angulate, with a punctiform hilum more or less centered on the ventral 
ridge. Generally, these seeds exhibit morphology distinctive among Mexican 


and American species of Hedyotiss.l. Five of these species are reported to have a 


TERRELL, STENOTIS 901 


chromosome number of x=13 (Lewis 1962), a unique number in the Hedyotideae 
(Terrell 1996). A recent DNA study (Church 2001, unpublished) has shown that 
four of the species form a clade distinct from species of Houstonia (Terrell 1996) 
and Stenaria (Terrell 2001). 

The heterogeneous genus Hedyotis was discussed by Terrell (1996) and com- 
pared with Houstoniaand Oldenlandia species. More recently, DNA results from 
Bremer and Manen (2000) found Hedyotis to be paraphyletic. All North Ameri- 
can species of Hedyotis are distinct from the type species, H. fruticosa, which is 
representative of a group of Sri Lankan and other Asian species. One species of 
the Asian group has a chromosome number of 2n=90-160 (Kiehn 1986), and 
the seeds of these species also differ from the North American species (Terrell 
1996). 

In consideration of the uniqueness of the Asian species, the Baja Califor- 
nian species heretofore treated as Hedyotis need a new generic name. The name 
chosen here, Stenotis, is derived in two parts: first, “Sten-,” from Stenaria,a recently 
named genus (Terrell 2001), whose name was based on Houstonia subgenus 
Stenaria Raf. second, from the root “-otis,” meaning “ear,” which terminates the 
name Hedyotis. Also, the Greek word, “stenos,” meaning “narrow, is descriptive 
of the narrow leaves of the plants in the new genus. 


SYSTEMATIC TREATMENT 
Stenotis [errell, gen. NOV. TYPE SPECIES: Stenotis mucronata (Benth.) Terrell. 
Herbae annuae vel perennes lignosae; folia 3-50 mm longa, 0.3-6.0 mm lata, plerumque linearia vel 
eon elliptica; inflorescentia cymosa, floribus heterostylis; corollae 2-18 mm longae 
fundibuliformes vel Lean albae vel roseolae; capsulae 1.3-5.0 mm longae 1.3-4.0 mm 
- atae subglobosae vel turbinatae; semina 0.3-1.0 mm longa 0.2-0.7 mm lata paulo compressa plus 


— 


minusve ellipsoidea pagina ventralis convexa vel rotundata hilo ee chromosomatum nu- 
merus x=13. 

Annual, soft-stemmed, or perennial woody-stemmed herbs or shrublets. Stems 
3-100 cm tall, terete or angular, branched. Leaves 3-50 mm long, 0.3-6.0 mm 
wide, linear, filiform, or subterete to narrowly elliptic or narrowly oblanceolate, 
thin or thickish. Stipules to 1.5 mm long, to 2 mm wide, whitish, scarious, 
toothed, lobed, or entire. Inflorescence cymose, terminal or axillary, earliest flow- 
ers sometimes sessile, later flowers pedicelled. Flowers heterostylous. Hy- 
panthium (calyx cup) hemispherical or cup-shaped; calyx lobes 0.5-2.5 mm 
long, usually linear or lanceolate. Corollas 2-18 mm long, salverform or funnel- 
form, white, pink, or rose, 8-nerved, tube longer than lobes. Pin flowers with 
stigma lobes 0.3-18 mm long. Thrum flowers with anthers 0.5-1.7 mm long. 
Capsules 1.3-5.0 mm long, 1.3-4.0 mm wide, subglobose or turbinate, 2/3 to 7/ 
8 inferior, with 8 dark nerves, dehiscing loculicidally then septicidally; placenta 
attached ca.l/3 to 2/5 of distance from base to top of septum. Seeds to ca. 47 per 
capsule, 0.3-1.0 mm long, 0.2-0.7 mm wide, brown or black, slightly or some- 


902 BRIT.ORG/SIDA 19(4) 


what dorsiventrally compressed, elliptic, oblong or irregularly obtusely angled 
in outline, dorsal face flat or convex, ventral face flat or convex to rounded with 
large hilar ridge, hilum punctiform, centric, testa reticulate. Chromosome num- 
ber x=13 for five species, others unknown. 

Distribution.—Baja California, Mexico. 

The following key is partly derived from Johnston (1924) and Wiggins 
(1980). 


1. Plants annual, soft-stemmed. 


2. Leaves 0.5-6.0 mm wide; oldest flowers mostly sessile; corollas white 1.S.arenaria 
2. Leaves 0.3-2.0(—3.5) mm wide; fl all or mostly pedicelled; corollas pink, rose, 
white 2. S.asperuloides 


1. Plants perennial, stems woody at least at base. 
3. Plants densely canescent or densely puberulent; rare species of southern Baja 
California Sur 7.S. peninsularis 
3. Plants glabrous or glabrate. 
4. Stems angular in cross section; leaves often fascicled, to 18 mm long. 


5. Plants stout, bushy; leaves numerous, crowded 6.S.mucronata 
5. Plants slender,not bushy;leaves not numerous, not crowded: known mainly 
from San Diego Island 5. S.gracilenta 


4. Stems terete; leaves not fascicled, to 50 mm long. 
6. Corollas (4—)5—10(-11) mm long 3.S. australis 
6. Corollas (8-)10-18 mm long 4.S. brevipes 


1. Stenotis arenaria (Rose) Terrell, comb.nov. Houstonia arenaria Rose, in Vasey, G. and 
J.N. Rose, Contr. U.S. Natl. Herb. 1:70.1890. Hedyotis arenaria (Rose) WH. Lewis, Rhodora 
63:221.1961. TYPE: MEXICO. BAJA CALIFORNIA: La Paz, 20 Jan-5 Feb 1890, Palmer 28 (LECTO- 
TYPE, here designated: US! ISOLECTOTYPES: F! GH! NY!) 


Small annual herb. Stems 3-30 cm tall, slender, quadrangulate to terete, erect or 
spreading, glabrous to scaberulous, with slender branches 2-20 cm long from 
any or all nodes. Leaves 5-40 mm long, 0.5-6.0 mm wide, sessile or lower leaves 
with short petioles, thin, l-nerved, narrowly oblanceolate, narrowly elliptic, or 
linear, glabrous. Stipules to ca. 1mm long and wide, rounded, with several mar- 
ginal teeth, some gland-tipped. Inflorescence with earliest flowers sessile in axils 
of peduncles or branches, surpassed by later erect peduncles or pedicels to 16 
mm long, (sometimes appearing secund), buds obovate. Hypanthium glabrous; 
calyx lobes 0.5-2.0 mm long, 0.2-0.4(-0.7) mm wide, lanceolate or narrowly 
lanceolate. Corollas 2-6 mm long, funnelform, white; tube 1-3.5 mm long, 0.6-3 
mim wide at throat, sometimes abruptly flared at throat, glabrous within; lobes 
1-3 mm long, 0.5-2 mm wide, ovate. Pin flowers with stigma lobes 0.3-1.0 mm 
long, linear, exserted to ca. 1 mm beyond throat, anthers at or just below corolla 
sinuses. Thrum flowers with anthers 0.5-1,.0 mm long, oblong, subsessile or on 
filaments to 1 mm long, exserted to ca.l mm beyond throat, stigmas near mid- 
point of tube. Capsules 2-3(-4) mm long and wide, 3/4-7/8 inferior, subglobose 
to subturbinate, tan or straw-colored with 8 darker nerves, thin-walled. Seeds 
0.4-0.9 mm long, 0.25-0.60 mm wide, in outline oblong or elliptic or irregu- 


TERRELL, STENOTIS 903 


larly obtusely angled, dorsal face flat or convex, ventral face with large rounded 
hilar ridge, testa reticulate. Chromosome number n=13 (Lewis 1962). 

Phenology.—Flowering August-September to April. 

Distribution.—Sandy places, granitic talus, and similar habitats at low el- 

evations; Mexico, Baja California Sur in Cape region from La Paz south to end of 
peninsula. 
Selected representative specimens examined: MEXICO. Baja California: 1 km S of Caduano, Lewis 
5341 (MO, SMU, TEX, US); 11 km N of Santa Anita, Moran 69179 (ARIZ, GH, K, MEXU, MICH, TEX); San José 
del Cabo, Purpus s.n., Jan-Feb 1901 (F, MO, NY, US); Arroyo de San Bartolo, 1.5 mi NW of village of San 
Bartolo, Wiggins 14751 (ARIZ, GH, K). 


An Arizona species, Hedyotis greenei A. Gray, is being studied to determine 
whether it is conspecific with S. arenaria. 

I have examined 26 collections of S. arenaria. This species appears to inter- 
grade slightly with S. asperuloides. 


2. Stenotis asperuloides (Benth.) Terrell, comb. nov. } Hedyot es Benth., Bot. 
Voy. Sulphur 19, t. 13. 1844. Houstonia asperuloides (Benth.) A ‘Gia Bre caer Acad. Art 
5:158. 1861. TYPE: MEXICO. BAJA CALIFORNIA: C ape San Lucas, Hinds s.n., 1841 Pear Rs ee 
designated: K!, not found at BM). 


Small annual herb. Stems to 2.8 dm tall (28 cm), very slender, terete or slightly 
angulate, ascending or decumbent, spreading, glabrous or puberulent-scabrous 
near base, diffusely much-branched from base or all nodes, internodes longer 
than leaves. Leaves 3-30 mm long, 0.3-2.0(-3.5) mm wide, sessile, linear or fili- 
form, glabrous or scabrous above, glabrous beneath, obtuse or acutish. Stipules 
tol mm long, to2 mm wide, rounded or truncate, margin with several linear or 
lanceolate teeth, some gland-tipped. Inflorescence spreading, flowers often 
numerous, earliest flowers sessile or shortly pedicellate, later flowers on fili- 
form pedicels to ca. 30 mm long, buds obovate. Hypanthium glabrous to pu- 
bescent; calyx lobes 0.6-2.3 mm long, linear or lanceolate, obtuse or acute. Co- 
rollas 3-11 mm long with lobes extended, funnelform or subsalverform, pink, 
rose, or white, sometimes with blue or green central nerves on lobes and red- 
dish spots at throat and with black spots externally near midpoint of tube, gla- 
brous or puberulent externally; tube 2-7 mm long, 1-4 mm wide at throat, some- 
times abruptly widened distally, glandular-puberulent or puberulent within 
especially near throat; lobes 1.5-5.0 mm long, 1-4 mm wide, ovate. Pin flowers 
with stigma lobes 1.0-1.4 mm long, linear, exserted to 1 mm long, anthers sessile 
near midpoint of corolla tubes. Thrum flowers with anthers 1.0-1.3 mm long, 
linear, sessile or subsessile, exserted just beyond corolla sinuses, stigmas in- 
cluded. Capsules 1.3-5.0 mm long, 1.3-2.0 mm wide, about 7/8 inferior, turbi- 
nate, oblong, or elliptic, brown, pale brown, or whitish with ca. 8 conspicuous 
dark nerves, thin-walled. Seeds 0.3-0.6 mm long, 0.2-0.4 mm wide, dorsal face 
flat or convex, shape quite variable, ventral face with conspicuous angulate hi- 


904 BRIT.ORG/SIDA 19(4) 


lar ridge or irregularly angulate or elliptic or oblong in outline with more 
rounded hilar ridge, with testa shallowly reticulate. Figure 2 (from Bentham’s 
protologue, 1844). 

Phenology.—Flowering October to May. 

The species is quite variable, especially in flower shape. Certain collections 
appear to resemble H. arenaria to some extent. Number of collections exam- 
ined: 26 

Two varieties have been recognized, as follows. 


KEY TO VARIETIES 


. Calyx lobes 0.8-2.3 mm long, often acute; corollas 6-11 mm long; capsules 2-5 


mm long, usually longer than wide var. asperuloides 
1. Calyx lobes 0.6-1.3 mm long, often obtuse; corollas 3-6 mm long; capsules 1.3-2.0 
long, usually equally long and wide var. brandegeana 


2a. Stenotis asperuloides var. asperuloides 

Calyx lobes 0.8-2.3 mm long, often acute. Corollas 6-11 mm long; tubes 3.5-6.5 
mm long; lobes 2-5 mm long, 1-4 mm wide. Capsules 2-5 mm long, 1.5-2.0 mm 
wide, narrowly turbinate, oblong, or elliptic. Chromosome number: n=13 (Lewis 


Distribution.—Sandy or gravelly flats or slopes, dunes, dry banks, stream 
beds, and similar habitats at low elevations near ocean and inland. Mexico: Baja 
California Sur, Cape region from La Paz area to southern end of peninsula. 


Selected representative specimens examined: MEXICO. Baja California: cae E of La Paz, along 
road to Punta Coyote, Carter 2606 (GH, K, LL, MO, US); Pacific coast N of Arroyo de Candelaria, NE 
Cabo San Lucas, Constance 3187 (F, GH, K, LL, MICH, MO, NY, US); 1 km a of El Triunfo off hwy 1 
Lewis 5337 (MO, SMU, TEX, US); 2.5 km NE of Cabo San Lucas, Moran 7044 (ARIZ, GH, K, MEXU, Ve 
TEX); 2 mi E of Buena Vista, Reeder & Reeder 6754 (ARIZ, ENCB); 8 mi N of Todos Santos, Shreve 7216 
(ARIZ, MICH, US). 


2b. Stenotis eiccconraes var. brandegeana (Rose) Terrell, comb. nov. Houstonia 
brandegeana Rose, in Vasey, G. and J.N. Rose, Cent: USS. Natl. Herb. 1:70. 189 0. ehnieees 
asperuloides f. brandegeana (Rose) WH. Lewis, Rh 63:221. 1961. H 
var. eee eres, oe in Shreve, F &@ LL. Wiggins, Veg. & Fl. Sonoran Desert 21400. 
-dyotis asperuloides var. brandegeana (Rose) Terrell, Phytologia 71:224. 1991. TYPE: 
= 0. Baja Gai: La Bae 20 Jan-5 Feb 1890, Syntypes: Palmer 31 and 24a (LECTOTYPE, 
here designated: at 31 (US); ISOLECTOTYPES: F!, GH!, K!, MEXUI). Rose in the protologue 
cited Palmer 31 and 24a together, which theref types. Palmer 31 is the better and 
most widely distributed collection. 


Calyx lobes 0.6-1.3 mm long, often obtuse. Corollas (3-)3.5-6.0 mm long, tubes 
2-3mm long, lobes 1.5-3.3 mm long, 1.0-2.2 mm wide. Capsules 1.3-2.0 mm long 
and wide, shortly turbinate to elliptic. Chromosome number: n=13 (Lewis 1962). 

Distribution.—Mexico. Baja California. Same distribution as var. 
asperuloides. In the protologue Rose noted that this taxon occurred in the same 
habitats as H. asperuloides. 


TERRELL, STENOTIS 905 


Fis. 2. Stenotis asperuloides. Drawing of holotype, Plate XIII, Bentham’s protologue, 1844. Left to right (all magnified): 


' , 7 


tati ined: MEXICO. Baja California: La Paz, Palmer 24a, 20 Jan- 
Feb 1890. (US); near hwy 1 SE of San Bartolo, 28.2 mi SE of El Triunfo, Daniel 2522 (ASU); valley 4.8 
km SW of Santiago, Carter et al. 2182 (ARIZ, F, GH, K, MO, US);4 km SE of San Pedro, Lewis 5336 (MO, 
SMU, TEX, US); Punta Frailes, Dawson 1773 (MICH) 


906 BRIT.ORG/SIDA 19(4 


This variety intergrades somewhat with var. asperuloides, however, | here fol- 
low Wiggins (1964) in maintaining it as a variety, although Johnston (1924) 
noted that it “can scarcely be distinguished”. In herbarium specimens | found 
certain collections with conspicuously smaller capsules and somewhat smaller 
flowers. Without having seen these plants in the field or having available other 
evidence | prefer to retain var. brandegeana. Number of collections seen: 8. 


3. Stenotis australis (1.M. Johnst.) Terrell, comb. nov. Ho | : LM. Johnst., 
Univ. Calif. Publ. Bot. 7:446. 1922. Hedyotis saxatilis WH. Lewis, Rhodora 63:222. 1961, non 
Hedyotis australis WH. Lewis & D.M. Moore, Southw. Naturalist 3:208. 1959. a MEXICO. 
BAJA CALIFORNIA: Binorama, 27 Sep 1899, T.S. Brandegee s.n. (HOLOTYPE: UC-201101)). 

Perennial woody herb. Stems to ca. 6 dm tall, woody toward base, slender, 

subterete, ascending or spreading, glabrous or minutely puberulent in inflo- 

rescence, epidermis brownish or grayish, often shredding. Leaves to 40 mm long, 
to 2mm wide, sessile, linear or filiform, glabrous or minutely puberulent, ob- 
tuse or acute and somewhat indurate at apex. Stipules 0.5-1.5 mm long and wide, 
truncate to deltoid, with l-few marginal teeth to ca. 1.5 mm long, sometimes 
gland-tipped. Inflorescence cymose, terminal. Earliest flowers sessile or 
subsessile, later flowers on pedicels to 10 mm long. Hypanthium glabrous or 
densely puberulent; calyx lobes 1-2 mm long, lanceolate or linear-lanceolate, 
acute. Corolla (4-)5-10(-11) mm long, narrowly funnelform, pink, light pink, or 
lavender-pink, throat sometimes yellow, glabrous or densely whitish-puberu- 
lent externally; tube 3-7 mm long, somewhat widened distally, glabrous or 
puberulent distally within; lobes 1.5-3.5 mm long, |-2 mm wide, ovate. Pin flow- 
ers with stigma lobes 1-15 mm long, linear, exserted ca.1 mm beyond throat, 
anthers included just below throat. Thrum flowers with anthers ca.1 mm long, 

elliptic or oblong, sometimes curved, purplish, subsessile or filaments to 0.4 

mm long, partly or fully exserted. Capsules (1.5-)2-2.5(-3) mm long, equally 

wide, 3/4-4/5 inferior, subglobose, often whitish with dark ribs. Seeds 0.45- 

0.8 mm long, 0.3-0.6 mm wide, usually longer than wide, rather shiny, in out- 

line oblong or irregular, dorsal and ventral faces flat or slightly convex, testa 

minutely and shallowly reticulate. Chromosome number: 2n = 26 (Lewis 1962). 

Phenology.—Flowering and fruiting September to April. 

Distribution.—Granite outcrops, rocky cliffs, canyon walls, among granite 
boulders in oak forest; altitudes 250-2000 m (750-6000 feet); Mexico. Baja Cali- 
fornia: Cape Region from San Bartolo (south of La Paz) south to tip of peninsula 
in the Sierra de la Laguna and Sierra de la Victoria. The type locality (“Binorama” 

or Vinorama) is apparently southeast of La Paz. Number of collections seen: 13. 


Selected representative specimens examined. MEXICO, Basa CALIFORNIA: Ca. 2.4 km SE of Rancho San 
Bernado, Carter 2693 (LL, US);La Laguna, Sierra de la Laguna, E of Todos Santos, Carter et al, 2337 (GH, 
US);Sierra de la Laguna, Hammerly 392 (CAS, GH, US); near canyon mouth, El Chorro, Moran 7300 (GH, 
MEXU, US) 


TERRELL. STENOTIS 907 


4. Stenotis brevipes (Rose) Terrell, comb. nov. Houstonia brevipes Rose, in Vasey, G. &J.N. 
Rose, Contr. U.S. Natl. Herb. 1:83. 1890. Hedyotis brevipes (Rose) WH. Lewis, Rhodora 63:221. 
1961. TyPE: MEXICO. BAJA CALIFORNIA: protologue stated “only a single specimen collected 
near Santa Rosalia, in a canon, 24 Feb-3 Mar, 1890, Palmer 202 (LECTOTYPE, here designated: 
US! ISOLECTOTYPES: F! GH! NY! USI). See explanation below. 


Perennial herb or shrublet. Stems toca. 1 m tall, herbaceous or becoming woody 
toward base, crown or base to 1 cm thick, terete, ascending or spreading, gla- 
brous, epidermis gray, sometimes shedding. Leaves 5-30(-50) mm long, 0.3-1 
(-2) mm wide, sessile, linear or filiform, glabrous, acute or mucronulate at apex. 
Stipules less than 2 mm long and wide, subtriangular, apices truncate or 
rounded, with marginal teeth. Inflorescences cymose, terminal, few-flowered. 
Earliest flowers short-pedicelled or subsessile, later flowers with pedicels to ca. 
12 mm long. Hypanthium Seer calyx lobes 0.5-2 mm long, 0.2-0.8 mm 
wide, (in fruit equalling or slightly su le), triangular to lanceolate, 
acute or obtuse. Corolla (8-)10-18 mm 1 long, salverform, pink, lavender, rose, or 
white, glabrous externally; tube (6-)8-13 mm long, usually about 3-4 times 
longer than lobes, slender, abruptly widened around anthers in pin flowers, 
glabrate or puberulent distally within; lobes 1.5-4(-5) mm long, 1-2.5(-3) mm 
wide, ovate, glabrous or minutely densely puberulent. Pin flowers with stigma 
lobes 0.7-1.8 mm long, linear, exserted 1-2 mm beyond throat, anthers included, 
located near 3/4-point of tube. Thrum flowers with anthers 1-1.5 mm long, ver- 
satile, elliptic or oblong, subsessile or on filaments to 0.5 mm long, exserted 
partly or fully at corolla lobe sinuses. Capsules 1.5-3.5 mm long, equally wide, 
subglobose, 2/3-4/5 inferior, light tan with slightly darker nerves. Seeds 0.6-1 
mm long, 0.3-0.7 mm wide, in outline elliptic or oblong, ventral face flat or 
slightly concave, testa shallowly reticulate. Chromosome number: 2n=26 (Lewis 


Phenology.—Flowering October to June. 

Distribution—In sandy, rocky, or gravelly places, canyon walls and floors, 
talus slopes, granitic outcrops, bay margins, at low altitudes; Mexico: Baja Cali- 
fornia, central and east side of Baja California peninsula and on various off- 
shore islands in ius California, from ca. 24° to 29° N latitude. Number of 
collections examine 


Selected representative specimens examined. MEXICO. BAJA CALIFORNIA. oe de! mello, E ot La Paz, 
ee (K,LL, US); Isla Coronado, Carter 4337 (ENCB, MICH); Arroyo Culebriad 
Carter & Ferris 4064 (ARIZ, MICH, TEX, US);4.4 mi NW of El Triunfo, Danie! 2488 nee Coyote Bay, Bahia 
Concepcion, Gentry 4063 (GH, MO, NA); San Nicholas Bay, Johnston 3723 (F, GH, K, MO, NY, US); Arroyo 
del Salto, Cape Region, Moran 7140 (ARIZ, GH, TEX); NW side of San Marcos Island, Bay of California, 
Moran 8945 (KANU, LL, MO, NY, PH); Carmen Island, Palmer 836 (F, GH, K, MICH, NY, US); Isla Ceralvo, 
Gulf of California, Wiggins 17754 (ENCB, MEXU, MICH, US); Santispaquis, Bahia de la Concepcion, 
Wiggins & Wiggins 17975 (BM, MEXU, MICH, TEX, US) 


Vasey and Rose (1890) provided a detailed explanation of Palmer’s collecting 
activities, but data on the type labels are confusing. The collections were made 


908 BRIT.ORG/SIDA 19(4) 


in 1890, although the US and GH specimens are labeled 1889. Palmer collected 
at Santa Rosalia from February 20 to March 3, 1890, then departed, but returned 
on March 15. Of the type specimens cited, the lectotype and GH isotype are 
labeled as “February 24 to March 3,” but the US isotype is labeled as just “1889”. 
The Fand NY isotypes also bear the collection number 202 and look like pieces 
of the same plant as the other types, but are labeled “March 15, 1890”. As only 
one specimen was collected originally, the US and GH specimens must be parts 
of the same plant. The Fand NY specimens apparently collected on March 
15 on Palmer's return. These are cited here as isotypes; possibly they should be 
considered paratypes. 

This species and H. mucronata are the two most wide-ranging species of 
Stenotis. 


5. Stenotis gracilenta (1.M. Johnst.) Terrell, comb. nov. Houstonia gracilenta LM. Johnst., 
Proc. Calif. Acad. Sci. ser. 4, 12:1174. 1924. sa eosin: M. Johnst.) WH. Lewis, Rhodora 
63:222. 1961. TyPE: MEXICO. BAJA CALIFORNIA: rocky slope facing sea, San Diego Island, 27 
May 1921, LM. Johnston 3927 (HOLOTYPE: CAS-1306h ISOTYPES: -GHI K! US). Usotypes labeled 
“egracillima”). 


Limited description based on holotype and two isotypes with damaged or im- 
mature flowers and description in Johnston (1924) and Wiggins (1964). 

Perennial shrublet with thick woody base. Stems to 20cm or more tall, slen- 
der, angulate, woody, decumbent, glabrous or slightly glandular, much- 
branched, epidermis on old stems gray or brown, shredding. Leaves to 15 mm 
long, tol mm wide, sessile, frequently fasciculate, filiform, coriaceous, acute or 
mucronulate. Stipules less than 1 mm long, subtriangular or lobed, sometimes 
with short, glandular teeth. Hypanthium glabrous; calyx lobes to 2.5 mm long, 
linear-lanceolate. Corollas 10-12 mm long with lobes extended, salverform, pink 
with darker rose lines extending to throat: tube 5-6 long; lobes ca. 2mm or more 
long. Capsules 1.5-2.5 mm long and wide, 2/3-3/4 inferior subglobose or slightly 
longer than wide. Seeds (2 seen) ca. 0.8 mm long. Flowering May. Chromosome 
number unknown. 

Distribution —Mexico: Type collection from San Diego Island, Gulf of Cali- 
fornia, Baja California. This small island is 75 miles NNW of La Paz and just 
south of San José Island. Another collection from San Francisco Island (Wig- 
gins et al. 377, DS) agrees with the protologue. 

Johnston (1924) noted that the type collection resembled both H. brevipes 
and H. mucronata, particularly the latter; consequently, it seems likely that H. 
gracilenta isa hybrid of these two species, and my study tends to confirm this. 
Hedyotisgracilenta needs field study and is here provisionally treated as a spe- 
cies. The taxonomic treatment by Wiggins (1980) stated that it occurs on sev- 
eral islands in the Gulf of California between Loreto and La Paz, but I have seen 
only the two collections cited. 


TERRELL, STENOTIS 909 


Wiggins 17812 (MICH) from San Diego Island and Wiggins et al. 378, (DS, 
MEXU, MICH, US) from San Francisco Island were labeled by Wiggins as H. 
gracilenta; however, the former is H. mucronata, the latter H. brevipes. Houstonia 
mucronata was collected on San Diego Island by Moran (9592, MICH). 


6. Stenotis mucronata (Benth.) Terrell, comb. nov. Hedyotis mucronata Benth., Bot. Voy. 
Sulphur 19. 1844. Houstonia mucronata (Benth.) B.L. Rob., Proc. Amer. Acad. Arts 45:401. 1910. 
TYPE: MEXICO. BAJA CALIFORNIA: Bay of Magdalena, Oct-Nov 1839, G.W. Barclay 3093 (LEC- 
TOTYPE, here designated: BM!; ISOLECTOTYPE: k!). 

Houstonia fruticosa Rose, Contr. U.S. Natl. Herb. 1:132. 1892. TyPE: MEXICO. BAJA CALIFORNIA: 
Beach, Carmen Island, 1-7 Nov 1890, Palmer 885 (LECTOTYPE, here designated: US-47250); 
ISOLECTOTYPES: CASI, Fl, GH!, K!, MO!, NY!, US); photo F! of US type. 

Perennial shrublet or herb. Stems 2-9 dm tall, stout, to 1.5 cm thick at base, gla- 
brous, with gray, tan, or yellowish shredding epidermis, densely leafy, much- 
branched, branches erect or ascending. Leaves 3-18 mm long, to 18 mm wide, 
sessile, fascicled, linear or subterete, thickened, straight or falcate, glabrous or 
scabridulous, mucronulate. Stipules to 1 mm long and wide, deltoid or ovate, 
entire or lobed, with brownish marginal glands. Inflorescences terminal, cymose, 
few-flowered. Flowers subsessile, or with pedicels to ca. 10 mm long. Hy- 
panthium glabrous; calyx lobes 13-3 mm long, 0.5-1.2 mm wide, lanceolate, 
ovate, or obovate, thickened, acute, sometimes with a stipitate gland on sinus 
margin. Corolla 8-14 mm long, salverform, white or tinged or lined with pink 
or rose; tube 6-10 mm long, about 1 mm wide at base, only slightly widened 
distally, glabrate within; lobes 1.5-4 mm long, 1-3 mm wide, ovate. Pin flowers 
with stigma lobes 1-15 mm long, linear, exserted 1-2 mm beyond corolla throat, 
anthers included, attached at 3/4- to 4/5-point of corolla tube. Thrum flowers 
with anthers 1.0-1.5 mm long, elliptic or oblong, whitish, sessile at corolla si- 
nuses, somewhat exserted, stigma included, extended toca. 1/2- to 2/3-point of 
corolla tube. Capsules 2-3 mm long and wide, 3/4-7/8 inferior, subglobose, 
broadly rounded or retuse at apex. Seeds 0.65-0.95 mm long, 0.35-0.6 mm wide, 
in outline elliptic, oval, or oblong, ventral face flat or slightly convex, testa shal- 
lowly reticulate. Chromosome number: 2n=26 (Lewis 1962). 

Phenology.—Flowering November to June. 

Distribution Beaches, dunes, salt flats, and rocky slopes and cliffs near 
beaches; Baja California between approx. 24° and 26° N; coastal areas along the 
Gulf of California and offshore islands from Coronados Island (ca. 26’) south to 
La Paz area (ca. 24°); also, Magdalena Bay area (S of 25°) on west side of B.C. pen- 
insula. This species is fairly frequent in suitable habitats, judging by the many 
specimens seen (27 collections). 


Selected representative specimens examined. MEXICO. Basa CatiFornia: Punta Prieta, Bahia de la 
Paz, Carter & Kellogg 3221 (K, MICH, US);Carmen Island, SE shore, Johnston 3836 (BM, F,GH, MO, NY, US); 
San Francisco Island, Johnston 3954 (GH, NY, US);Ceralbo Island, Gulf of California, Johnston 4035 (GH, 


910 BRIT.ORG/SIDA 19(4) 


MO, NY, US); Magdalena Bay, Mason 1947 (F, GH, K, NY, US); San Francisco Island, Moran 3726 (BM, 
MICH, TEX, US, WIS); Santa Margarita Island, Rose 16306 (US); 6.8 mi E of La Paz on road to Pichilingue 
Bay, Wiggins 14751 (ENCB, GH, K, TEX); Las Cruces, 22.5 mi SE of La Paz, Wiggins 15671 (BM, MICH, US). 


7. Stenotis peninsularis (Brandegee) Terrell, comb. nov. Houstonia peninsularis 
Brandegee, Zoe 5:160. 1903. Hedyotis peninsularis (Brandegee) WH. Lewis, Rhodora 63:222. 
1961. TyPE: MEXICO. BAJA CALIFORNIA SUR: Sierra de la Trinidad, Cape region, Nov 1902, T.S. 
Brandegee s.n. (LECTOTYPE, here designated: UC IsOLECTOTYPEs: GHI, USI, VT), 


Perennial woody herb or shrublet. Stems to 50 cm tall, woody toward base (to at 
least 1 cm thick), subterete, erect, very densely canescent or densely puberu- 
lent with grayish or brownish hairs to 0.2 mm long, epidermis brownish, shred- 
ding. Leaves to 40 mm long, 0.5-2 mm wide, sessile, sometimes fasciculate, lin- 
ear, very densely canescent, some hairs gland-tipped, obtuse, revolute. Stipules 
toca. 1 mm long and wide, truncate, with 1-few marginal hair-like teeth, often 
gland-tipped. Inflorescences terminal, cymose. Flowers sessile or with pedicels 
to5 mm long. Hypanthium densely canescent; calyx lobes 0.8-3 mm long, lan- 
ceolate or narrowly lanceolate, acute or obtuse. Corolla to 17 mm long, salver- 
form, purple, densely to sparsely pubescent externally, pubescent to glabrate 
within; tube 6-12 mm long, narrow, scarcely or only slightly wider at throat: 
lobes 2-6 mm long, 1-3.3 mm wide, ovate. Pin flowers with stigma lobes ca. | 
mm long, exserted ca. | mm beyond corolla throat, anthers included just below 
throat. Thrum flowers with anthers 1.1-1.7 mm long, linear-oblong, exserted on 
filaments toca. | mm long. Capsules 2-2.8 mm long and wide, 2/3-7/8 inferior, 
subglobose, dark brown varying to whitish with conspicuous nerves, hirtel- 
lous. Seeds (only seen from Purpus 427; one capsule), 0.6-0.75 mm long, 0.35- 
0.5 mm wide, in outline oblong, elliptic, or irregularly angular, dorsal face flat 
or slightly convex, ventral face broadly rounded or more angulate, testa shal- 
lowly reticulate. Chromosome number unknown. 

Phenology.—Flowering November to March. 

Distribution.—Mexico: Baja California Sur in the Sierra de la Trinidad, Cape 
region. Only the two collections known. Brandegee (1903) stated that “It grows 
abundantly ... generally on nearly perpendicular bluffs barren of other vegetation.” 


Additional specimens examined. MEXICO. Basa Catirornia Sur: Sierra de la Trinidad, Cape region, 
Jan-Mar 1901, CA. Purpus 427 (MO! US)), 

Brandegee (1903) stated that the species was based on collections by Purpus 
and himself, but he did not cite particular collections in the protologue for H. 
peninsularis. The collections by Brandegee and Purpus are the only two collec- 
tions known to me of this rare species. The better collection and the one labeled 
as a new species was the Brandegee collection. This little-known species ap- 
pears to be related to the other perennial Baja California species, H. brevipes 
and H. mucronata. Two Marcus E. Jones collections (24341: GH, and 27137:BM, F 


TERRELL, STENOTIS, 911 


MO, NY) were labeled H. peninsularis, but are actually H. australis and H. 
brevipes, respectively. 


ACKNOWLEDGMENTS 


I thank the curators of the herbaria cited here for access to their valuable col- 
lections. 1am especially grateful to John Wiersema and Piero Delprete for their 
meticulous reviews. Dan Nicolson and Harold Robinson contributed helpful 
assistance with the Latin description, and Gene Rosenberg provided the 
resumen. 


REFERENCES 


Branvecee, 1.S. 1903. Notes and new species of Lower California plants. Zoe 5:155-174. 

Bremer, B. and J.-F. Manen. 2000. Phylogeny and classification of the subfamily Rubioideae 
(Rubiaceae). Pl. Syst. Evol. 225:43-72. 

Cuurcu, S.A. Molecular phylogenetics of Hedyotis and Houstonia (Rubiaceae): descending 

\euploidy and cladogenesis in the North American lineage. Submitted for publication. 

oe ILM. 1924. Expedition of the California Academy of Sciences to the Gulf of Cali- 
fornia in 1921.Proc. Calif. Acad. Sci., ser. 4, 12:1173-1177. 

KiEHN, M. 1986. Karyosystematic studies on Rubiaceae: Chromosome counts from Sri Lanka. 
Pl. Syst. Evol. 154:213-223. 

Lewis, W.H. 1962. Phylogenetic study of Hedyotis (Rubiaceae) in North America. Amer. 
J. Bot. 49:855-865. 

Terrell, E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monog. 48: 

-118 

TerreLt, E.E. 2001. Taxonomy of Stenaria (Rubiaceae: Hedyotideae), a new genus including 
Hedyotis nigricans. Sida 19:591-614. 

Vasey, G. and J.N. Rose. 1890. List of plants collected by Dr. Edward Palmer in Lower 
California and western Mexico in 1890. Contr. U.S. Natl. Herb. 1:63-83. 

Wicains, |.L. 1964. Houstonia.|n:Shreve, F.and |.L.Wiggins. Vegetation and flora of the Sonoran 
Desert, 2 vols. Stanford University Press, Stanford, CA. Pp. 1398-1401. 

Wicains, |.L. 1980. Flora of Baja California. Stanford University Press, Stanford, CA. 


912 BRIT.ORG/SIDA 19(4) 


Book Notices 


ALAN E. Bessette, ARLEEN RAINIS BESSETTE, WILLIAM K. CHAPMAN, and VALERIE CONLEY 
CHAPMAN. 2000. Wildflowers of Maine, New Hampshire, and Vermont in 
color. (ISBN 0-8156-0586-2, pbk.). Syracuse University Press, 621 Skytop 
Road, Suite 110, Syracuse, NY 13244-5290, U.S.A. (Orders: http:// 
sumweb.syredu/su_press/,twalshO1@syredu, 315-443-5547, 315-443-5545 
fax). $24.95, pbk., 163 pp, 362 color photos, 61/2" x 91/4" 


The book includes ‘nearly 400 photographs’ of showy species, those most likely to be marveled at, 
each species with a brief, mid-technical description. The “primary reference source for nomencla- 
ture and terminology” is the “Revised Checklist of New York State Plants by R.S. Mitchell and G.C 
Tucker.” The photos are nicely done and most of them show aspects of the foliage as well as the flow- 
ers—the volume is worth having for the photos. Within each of the six color categories, the species 
are sorted by a series of key-like alternatives, using leatures of flowers, habit, and leas to help the 
user navigate toward matchings photo with plant in hand. A “Visual Glossary,’ “Glossary of Terms,” 
“Index of Common Names,” and “Index of Genera and Species” also provide help toward this goal. 


Lay it perhaps to the bias of a non-novice, but it’s hard for me to appreciate the value of this 
artificial color-arrangement of species, compared to the broader usefulness of grouping the photos 
by family. Users of this book who would want to check the identity of species by reference to the 
pithy descaphions ae semi- i-technical ACY s be have enough botanical back eae and savvy 

LRe 


vd Nesom. Botanica earch Insti- 


tute oe Texas, 509 Pecan Street, Bove Worth, 1X 76102- 4060, U.S 


Daniet ZonArY and Maria Horr. 2000. Domestication of Plants in the Old World, 
Third Edition. (SBN 0-19-850356-3, pbk.). Oxford University Press, 2001 
Evans Road, Cary, NC 27513, USA (Orders: www.oup-usa.org, 800-451-7556). 
$34.95, 316 pp, ens 9 a x 61/8". 


Asastudent of ethnobotany (fal } ical sid h 


as genu- 


Qu 


Alling I 
inely excited to see this book nim sticated Pla nisin the Ok Worl Id makes agieblew a wei salth of in- 
formation in a concise text that is both Leaman ae abe ne written. The book 


contains ten chz reo followed by site f rts. references, and an 


index. The book begins with a discussion of the sources of silence for the origin and spread of 
cultivated aos including archaeological evidence, evidence from living plants, and radiocarbon 
dating and dend eae The following seven chapters individually cover cereals, pulses, oils 
and fibre crops, fruit trees and nuts, vegetables and tubers, condiments, and dye ope : ea nine 
discusses plant remains in representative arc haeological sites — is f Mas ved aA a 
va f 


ing the ane of domestication and horticulture. 
int 


he previous editions 


> 


nat it includes the most recent findings from molecular ae about the genetic relations be- 


tween domesticated plants and their wild ancestors, added information on several new crop plants 
and incorporates extensive new archaeological data about - aes of agriculture, The chapter on 
“Fruit collected from the wild” that appeared in prey i s has been omitted from this edi- 


tion. | would highly recommend this book for eee and botany classes dealing with the 
origins of agriculture. It is a finely produced text ne shoul ane: on extended readership among 


lay readers interested in the subject.—Kevin D. Janni, Bot nstitute of Texas, Fort Worth 
TX 76102-4060, U.S.A, kjanni@britorg. 


SIDA 19(4): 912. 2001 


TAXONOMIC REVIEW OF HOUSTONIA ACEROSA AND 
H. PALMERI, WITH NOTES ON HEDYOTIS AND 
OLDENLANDIA (RUBIACEAE) 


Edward E. Terrell! 


Research Collaborator 
Department of Systematic Biology-Botany 
Nati oe Lee of Natural History 

n lactittition 


Washington, D.C. 20560-0166, U.S.A. 


ABSTRACT 


Generic relationships among Houstonia, Oldenlandia, and Hedyotis are discussed. Seed and other 
morphological characters, chromosome n aaa rs, and limited DNA data support the division of 


Hedyotis into smaller genera. The taxon a of Houstonia acerosa and H. palmeri 


are revised to accommodate two es eee varieties, H. acerosa var. tamaulipana and H. 
palmeri var. muzquizana. 


RESUMEN 


Se analizan las HelaCIOnes genericas entre Houstonia, Oldenlandia y Hedyotis. Datos de la semilla, 
’ datos limitados del ADN apoyan la division 


de Hedvotis : : Hos.'S ic | 


i AS a nomenclatura de Houstonia acerosa 


yvH. | i lar dos iedad i H. acerosad var. t imaulipan iy ag 


i if r 
palmeri var. muzquizana. 


GENERIC RELATIONSHIPS 


The first part of this paper discusses the Linnean genera Houstonia, Hedyotis, 
and Oldenlandia, whose relationships have been debated for several decades. 
The second part reviews and revises the taxonomy and nomenclature of 
Houstonia acerosa and H. palmeri from the southwestern United States and/or 
Mexico. These species were treated under Hedyotis by Turner (1995a, 1995b, 1997). 

The genus Houstonia, with 20 North American species, was the subject of 
a monograph (Terrell 1996a) that recognized two subgenera. Subgenus 
Houstonia has x=7 and 8 chromosomes and seeds with a ventral cavity lacking 
a hilar ridge (these species including the type, H. caerulea, are delicate herbs 
with salverform corollas). Subgenus Chamisme has seeds with an elongate hi- 
lar ridge in a ventral depression. Section Amphiotis (the H. purpurea group) 
has x=6 chromosomes and funnel-shaped corollas, and section Ericotis (the H. 
rubra group) has x=11 chromosomes, more variable corollas, and more com- 
plex seed morphology. The seeds of Houstonia sens. lat. are crateriform, refer- 
ring to the presence of a ventral cavity or depression. 


‘Address for correspondence: 14001 Wildwood Dr, Silver Spring, MD 20905, U.S.A. 
SIDA 19(4): 913 — 922. 2001 


914 BRIT.ORG/SIDA 19(4) 


The type species of Houstonia and the two related genera, Hedyotis and 
Oldenlandia, showed marked morphological differences (Terrell 1975). It is in- 
structive to review the taxonomic treatments of these three genera in the major 
floras of the eastern and mid-Atlantic United States. In the two latest editions 
of Gray’s Manual, Robinson and Fernald, 7th edition, (1908), recognized 
Houstonia with ten species and Oldenlandia with one species, and in the 8th 
edition Fernald (1950) listed eleven species of Houstonia and moved two spe- 
cies formerly in Oldenlandia to Hedyotis. Gleason (1952) in the widely-used 
Britton and Brown flora recognized Houstonia with nine species and 
Oldenlandia with two species. Gleason and Cronquist’s Manual (1991) recog- 
nized only the genus Hedyotis, with ten species. | believe that the three earlier 
floras provided more accurate taxonomic treatments of these genera. The 1991] 
Manual was apparently strongly influenced by the publications of ER. Fosberg, 
who consistently helda very broad concept of Hedyotis. Terrell 1996a) reviewed 
the history of all three genera in detail, and noted that Fosberg (1943) in his 
study of Polynesian Hedyotis recognized five subgenera while ing that 
“many botanists would regard these groups as genera.” 

Seed morphology (reviewed by Terrell 1996a) provides effective morpho- 
logical criteria when considered together with chromosome number. 
Oldenlandia and Hedyotis were found to have non-crateriform seeds (lacking 
ventral cavities or depressions with or without hilar ridges), in contrast to the 
crateriform seeds of Houstonia. Oldenlandia (Terrell 1990) generally hasa chro- 
mosome number of x=9 and seeds which are very small and trigonous with 
punctiform hilums on one of the three ridges; however, there are a few dissimi- 
lar species often included within Oldenlandia that do not have these charac- 
teristics. The genus has a worldwide distribution, but is most abundant in Af- 
rica, and several authors have provided taxonomic treatments of the African 
species, notably Bremekamp (1952) and Verdcourt (1976). 

Hedyotis sens. lat. is a worldwide, heterogeneous, “dust-bin” genus that in- 
cludes several distinct groups of species. Verdcourt (1976) and Halford (1992) 
commented that Hedyotis needs to be broken down into smaller, more homo- 
geneous units. The selection of a type species has been controversial. The spe- 
cies selected has been approved by at least one nomenclatural committee, as H. 
fruticosa L.,a native to Sri Lanka having a number of related species in Asia. 
Selection of the alternative species, H. auricularia L., would wreak havoc with 
the nomenclature and taxonomy, as the seeds and other characters suggest 
oldenlandioid relationships, and furthermore this species has been made the 
type of another genus, Exallage Brem. (Bremekamp 1952). Hedyotis fruticosa 
and its Asian relatives are not closely related to the approximately 20 North 
American, primarily Mexican, species of Hedyotis, nor to the varied Hawaiian 
species. Chromosomes of certain Mexican species have the numbers x=9, 13, 


TERRELL, TAXONOMIC REVIEW OF ACEROSA AND H. PALMERI 915 


and 17 (Lewis in Terrell et al. 1986). Preliminary data on chromosome numbers 
of Asian species suggest high numbers of chromosomes (Kiehn 1986). DNA data 
for Hedyotis and Oldenlandia indicated that both are paraphyletic (Bremer & 
Manen 2000). 


TAXONOMY OF HOUSTONIA ACEROSA AND H. PALMERI 


Houstonia acerosa (A. Gray) Bentham @ Hooker, a species of southwestern 
United States and Mexico south to San Luis Potosi, was among the 20 
monographed species of Houstonia (Terrell 1996a). Like the related species H. 
palmeri, H. acerosa was placed in the subgenus Chamisme Raf., section Ericotis 
(Terrell) Terrell. The species is quite variable; e.g., the leaves vary from whorled 
to opposite, fasciculate or scarcely fasciculate, the inflorescence is reduced and 
with sessile flowers and capsules or varies to pedicellate and cymose. It has gen- 
erally been treated as two varieties or subspecies under the epithets 
polypremoides or bigelovii (see nomenclatural summary below 

Earlier recognized two subspecies (Terrell 1979) or in later years sGanpublished) 
either two varieties or subspecies. In preparing my monograph, however, | de- 
parted from my previous views by stating that the variation seemed a continuum, 
and recognized only one variable species. My earlier course may have overem- 
phasized the intergradation, and here I recognize three varieties in H. acerosa. 

The species was discussed by Turner (1995a) under the genus name 
Hedyotis, and his publication in the August 1995 issue of Phytologia preceded 
the publication of my monograph by about two months. This resulted in my 
completing work on a Houstonia monograph before Turner’s paper appeared. 
He cited my overview of Hedyotis and related genera (Terrell 1991), which pro- 
vided the names and synonyms of North American species. He recognized the 
varieties acerosa and polypremoides and two new ones, var. potosina and var. 
tamaulipana. In 1996(b) I did not recognize Turner's varieties, and Turner (1997) 
provided what he called a vigorous rebuttal. I have reviewed Turner's varieties 
again, and present here a revision incorporating Turner's data and recognizing 
one of his two varieties. This has been an attempt to reconcile our differences of 
opinion, and I hope that we can henceforth leave the taxonomic matters as they 
are, and let future botanists study the species in the light of new knowledge. 

The following key outlines the new taxonomic treatment, and is followed 
by the nomenclature reorganized from Terrell 996a). 


= 


. Internodes usually 3- ” mm long; leaves OU ynio late fasciculate, numerous 
-| 


often crowded ile or on pedicels 
to 3mm long, sometimes d acre groupes var. acerosa 
1. Internodes usuall 25 some verticillate, not or some 


leaves fasciculate, not ere or Pouetoopine linear or acerose; flowers and cap- 
sules on pedicels 3-22 mm long, the inflorescence often open and diffuse. 


916 BRIT.ORG/SIDA 19(4) 


2. Corolla tubes 4-12 mm long; calyx lobes 2-7 mm long var. polypremoides 
2. Corolla tubes 3-4 mm long; calyx lobes 1.0-2.6 mm long var.tamaulipana 


A full description of Houstonia acerosa s.l. was presented in Terrell (1996a). This 
is here supplemented by the preceding key, a revised distribution map, and re- 
vised synonymy. 


Houstonia acerosa (A. Gray) Bentham & Hooker f. var. acerosa, 1, Gen. PL2 60: 
1873. Hedyotis acerosa A. Gray, Smiths. Contr. Knowl. 3:81. 1852. Olden! (A. Gray) 

A. Gray, ee Contr. Knowl. 5:67. 1853. Mallostoma acerosa (A. Gray) Hemsley, Biol. Centr.- 

Amer., Bot. 2:31. 1881. Ereicoctis Nee (eee Rev. Gen. PI. 1:281. 1891, (orth. var. of 
Eveicotis. TYPE: a S.A. TEXAS: Rissa Texas to El Paso, New Mexico ae -Oct 1849, C. Wright 


237 (HOLOTYPE: GH}; ISOT BN MO! US). As Turner (1995a) t, Wright's collec- 
id r 


tion was made in late June 1849 in present day Kinney or Val Verde County, southwest Texas. 


Habitat and distribution —Rocky or gravelly places, mountain slopes, mesas, washes, 
arroyos, desert grassland, desert scrub, often on limestone, also in clay, sandy, 
or gypseous soils; associated with Prosopis, Acacia, pinyon-juniper, or oak-juni- 
per; 400-2500 m, usually 900-2000 m (3000-6500 ft); United States: Central 
and southwestern Texas; Mexico: central Chihuahua to San Luis Potosi (Fig. 1). 


Houstonia acerosa (A. Gray) Bentham & Hooker f. var. piel Sata (A. Gray) 
Terr mb. nov. Houstonia polypremoides A. Gray, Proc. Amer. Acad. Arts 21:379. 1886. 
Hedyotis saat moides (A. Gray) Shinners, Field & Lab. 17:168. ie Hedyotis acerosa vat. 
polypremoides (A. Gray) WH. Lewis, Ann. Missouri Bot. Gard. 55:31. 1968, nom. supertfl. (see 


Hedyotis acerosa vat. esha Houstonia acerosa subsp. polypremoides (A. Gas) Terrell, 
Brittonia 31:168. 1979. Type: MEXICO. CHIHUAHUA: Santa Eulalia Mts., 26 Sep 1885, C.G. Prin- 
gle 356 (LECTOTYPE, designated by Lewis, 1968: GH! ISOLECTOTYPES: BM! CINC! F! K! MO! NA! 
NY! PH-2! US-2! VF-2)). SYNTYPE: MEXICO. C S Eulalia Mts., 29 May 1885, Prin- 
gle 16 (BM! CINC! F! GH! K! MO! NA! NY! PH-2! US-2! VT). 


Houstonia polypremoides var. bigelovii Greenman, Proc. Amer. Acad. Arts 32:29]. 1897. Hedyotis 
acerosd Vat. sl ae Lana Lewis, Ann. Missouri Bot. Gard. 55:397. 1969. TYPE: U.S.A. 
New Mexico: Mexican Boundary Survey, Florence Mts.,Jun 1852, Bigelow 437 (HOLOTYPE: GH). 
The rlbienee Mountains are thought to be the Florida Mountains, south of Deming, Luna 
Co., New Mexico (fide R. Spellenberg pers. comm.). 


The so-called autonym rule (Art. 22.3, Greuter et al. 2000) applies to Houstonia 
polypremoides var. bigelovii Greenman, as its publication in 1897 created a var. 
polypremoides, which then has precedence over var. bigelovii, when the latter is 
considered synonymous with Houstonia polypremoides A. Gray. 

Habitat and distribution.—Habitats similar to those of var. acerosa, New 
Mexico, western Texas, where it intergrades with var. acerosa; Mexico: Chihuahua, 
northern Coahuila (Fig. 1). 


Houstonia acerosa (A. Gray) Bentham & Hooker f. var. tamaulipana (B.L. Turner) 
Terrell, comb. nov. Hedyotis acerosa A. Gray var. tamaulipana B.L. Turner, Phytologia 
79:87-88. 1995. TyPE: MEXICO. TAMAULIPAS: Mpio. Villagran, | mi E of Ejido de San Lazaro ca. 
24° 35'N, 99° 13' W, 1500 ft, 11 Oct 1959, M.C. Johnston & J. Graham 4281k (HOLOTYPE: TEX). 
[Fig. 1] 


TERRELL, TAXONOMIC REVIEW OF ACEROSA AND H. PALMERI 


ia 


| 
E 


7 


@ 
: | e\e 
t 
‘ | e @ e e 
‘ Kl, e 
‘ 
he patel 
m/ em Ne 
: * 
‘ H f 
as 7= Con 
4 
\ ‘ a @ e® 
5 . Mw “ 
mm . A 8 
P 
Sen, . ® e t ) Sy ‘ 
. N Pe oe] e \ 
‘ i “teeter! e . 


Houstonia acerosa 
var. acerosa 
var. polypremoides 
var. tamaulipana 


| 
A 


ran ‘~ @: 
ve ws : Pa 
ve a om s 
. \ 4 ? 
‘ ae @ > 
. 103° 3 : @ ee 
‘ , “ e @: 0: 
‘ w 
eo, 
~ on 
« 
{ © 
400 km : @ 
wey 
. *. 


3 toe Ae Lal L 2 rf Lu 
Fic. 1. Distribution of the thr 


918 BRIT.ORG/SIDA 19(4) 


In my monograph I stated that | had studied three collections of H.acerosa from 
Tamaulipas at TEX that were possibly a distinct variety, and had finally con- 
cluded the differences were insufficient to merit their being distinct. Later, I 
had re-borrowed these three collections from TEX because of uncertainty 
whether they should be a new variety. I then decided again to not distinguish 
them as a new variety. As it turned out, these collections were recognized by 
Turner (1995a) as var. tamaulipana. In my 1996b paper I stuck by my previous 
conclusion and did not consider var. tamaulipana B.L. Turner asa distinct vari- 
ety. After further consideration I here recognize it as distinct and somewhat 
resembling var. polypremoides but differing in two overlapping floral charac- 
ters. It occurs in a separate area in Tamaulipas at the southeast end of the range 
of H. acerosa in somewhat different habitats, as noted by Turner, who provided 
a map of the distribution. 


Additional specimens examined. MEXICO. Tamautipas: Mpio. San Carlos, 6 mi S of San Carlos on the 
road to Padilla, 1600 ft, calcareous terraces of Arroyo de San Carlos, 13 Dec 1959, Johnston 5007A 
(TEX); Mpio. Casas, 5 mi E of Casas on Victoria-Soto la Marina hwy, 28 Sep 1960, Johnston 5784B 
(TEX). 


Hedyotis acerosa var. potosina B.L. Turner 
Turner (1995a) distinguished Hedyotis acerosa var. potosina, a new variety from 
Mexico, as follows: 


. Stems with internodes much-shortened, forming low pulvinate mat-like plants 
mostly 2-5 cm high; corolla tubes mostly 8-10 mm long; southernmost Coahuila 


and southwards to San Luis Potosi var. potosina 
1. Stems not as described in the above, mostly 5-15 cm high, forming well-defined 
rather naked stems; corolla tubes mostly 3-7 mm long (other varieties) 


I rejected this variety (Terrell 1996b), stating that I considered it as part of a 
cline, as the plants northward from Monterrey or Saltillo became larger and 
less pulvinate. 

To judge var. potosina more objectively | compared specimens from 22 col- 
lections (including Lundell 5048, HoLotype LL! isotype US!) at herbarium US from 
Mexico and southwest Texas, in the following characters: Height of plants, in- 
ternode length, corolla tube length, corolla lobe length, corolla length, and ca- 
lyx lobe length, the last three characters being possibly useful characters not 
mentioned by Turner. The collections were grouped in five categories as to ori- 
gin: San Luis Potosi; Coahuila south of Saltillo, Coahuila at or near Saltillo; 
Coahuila north of Saltillo; and from three southwest Texas counties. (Collec- 
tion data are listed in Appendix 1). 

Turner stated that var. potosina was a mat-forming plant “having a very 
different growth habit than found in var. acerosa.” My own finding is that it isa 
small, but erect plant in the southern part of its range with basal or near basal 
tufts of narrow needle-like leaves; however, I believe that northward the plants 
are gradually taller and appear less tufted, thus forming a cline. 


TERRELL, TAAQNOMIC REVIEW OF ACEROSA AND H. PALMERI 919 


Table 1 has only the raw data, as I believe that these are sufficient in them- 
selves to be easily interpreted. The data show that (1) specimens from near or 
south of Saltillo average slightly shorter; (2) Mexican specimens differed very 
little in internode length or corolla tube length, regardless of location, and there 
was much overlap; (3) the last three characters differed very little among the 
four geographic locations; (4) the southwest Texas collections generally were 
similar to Mexican collections, especially those from north of Saltillo. There is 
so much overlap in height and corolla tube length that there is no objective 
basis for recognizing var. potosina. The last three “additional characters” pro- 
vide information, but show no differences. 


Houstonia palmeri A. Gra 

Houstonia palmeri A. Gray, a species of northern Mexico, was, like H. acerosa, 
the subject of a short paper by Turner (1995b), which preceded the publication 
of my Houstonia monograph (1996a) by about two months. Turner’s paper rec- 
ognized a new variety, var. muzquizana. 

My monograph provided a description, synonyms, and distribution of var. 
palmeri, from which the data below are taken, with modifications from Turner’s 
data. Turner selected Palmer 395 (GH) as the type for the synonym, H. longipes 
S. Watson, whereas I selected Palmer 394 (see below). His selection has priority 
over mine. 


Houstonia palmeri A. Gray var. palmeri, Proc. Amer. Acad. Arts 17:202. 1882. 
lyotis palmeri (A. Gray) WH. Lewis, Rhodora 63:222. 1961. TYPE: MEXICO. COAHUILA: Lerios 
45 mi E of Saltillo, 10,000 ft, Jul 1880, Palmer 397 (LECTOTYPE, designated by Turner, 1995b, 
GH}; ISOLECTOTYPES: K! NA! NY! PH! USI). spi MES a 40 mi S of Saltillo, Si- 
erra Madre, 25 Jul- 7S 1880, Palmer 398 (F-2! Y! US-2! VT!). At GH 397 and 398 
are on the same shee 


Houstonia longipes S. Watson, Proc. Amer. Acad. Arts 18:97. 1883. Hedyotis longipes(S. Watson) WH. 
Lewis, Rhodora 63:222. 1961. TYPE: MEXICO. NUEVO LEON: Monterrey, 17-26 Feb 1880, Palmer 
395 (LECTOTYPE, designated - ae 1995b: GH! ; ISOLECTOTYPES: K! NA! NY! PH! US-2)). 


Habitat and distribution.—Gravelly, rocky, or sandy places, rock crevices, often 
over limestone, shale, or gypsum, associated with oak, pine, Agave, Acacia, Yucca, 
Larrea, Prosopis, usually 700-2300 m (2300-7500 ft; Mexico: Coahuila, Nuevo 
Leon, and San Luis Potosi. 


Houstonia palmeri var.muzquizana(B.L. Turner) Terrell, comb. nov. ae 
muzquizana B.L. Turner, Phytologia 79:91.1995. TypE: MEXICO. COAHUILA: M 
Muzquiz, ca. 130 road km NW of Muzquiz on Hwy 2A, Nesom & Mayfield 7380 aaa 
TEX! ISOTYPE: MEXU) 


In my monograph (Terrell 1996a) I commented that plants in the Muzquiz area 
have longer corollas, but later questioned (Terrell 1996b) whether Turner's vari- 
ety deserved recognition. I have now seen five additional collections in a further 
loan from Turner and accept var. muzquizana as consistently having longer 


920 BRIT.ORG/SIDA 19(4) 


Taste 1. Houstonia acerosa: Measurements for six characters from Mexico and Texas Collections. 


Collections Heightof Internode Corollatubes Corollalobes Corollalength Calyx lobes 
plants (cm) length length length totals length 
(mm) (mm) (mm) (mm) (mm) 
Mexico, el Luis Potosi 
Lundell 5 4-7 2-3 i 3 10 4-6 
Whiting “i 4-7 1-3 6-7 3 9-10 4-6 
Purpus ae 4-6 Q-2 7-8 3-4 10-12 3-4 
Nelson 4533 3-6 O-1 7 4-5 11-12 2.5-3 
Coahuila, south of Saltillo 
Palme 0-3 8-9.5 3--5 11-145 5-6.5 
Palmer 400 4-6 3-5 9-11 4-5 13-16 3-6 
Shreve 8540 5-7 2-5 8-11.5 3-4 11-15.5 4-6 
Coahuila, at /near Saltillo 
Palmer 12 3- Q-2 5-9.5 3-5 8-145 42-45 
Pennell 17264 3-4 2 6 4 10 
Pennell 17292 4-6 2-3 75 4 LS 4-45 
E.& B. Terrell 4431 4-6 O-2 5.5-10 3-5 8.5-15 45-65 
Stceay ane of Saltillo 
Palm 8 3-5 8 4 12 5-6 
Palm si 10-13 3-5 - 2 9 — 
Gian 7202 8 Q-2 6 2 8 3-4 
Chiang et al.7550Q 6-7 2 9 3 4-4.5 
Reveal et al. 2602 5-7 4-6 5-8.5 2.5-4 7.5-11.5 3.5-5 
Texas 
Pecos Co.: Lewis 5517 8-12 3-6 - - - 2-5 
TERRELL C 
BJ. Re 33530 8-11 4 4-6 3 7-9 3-4.5 
Orcutt 734 8-10 to 7 5 3 8 5-6 
Wooton 9/29/11 7-13 3-5 2-5 
VAL VERDE Co.: 
Rose 1/986 7-8 5 4 3 7 2-3 
E.J.Palmer 11057 6-10 4-7 7-8 2-3 9-11 4-6 


corollas. I note also that collections I have examined are mostly the pin form, 
with long styles and short stamens. 
All collections of var. muzquizana were shown in the detailed distribu- 
tion map provided by Turner (1995p). 
Turner (1995b) provided a key to the varieties as follows: 
Mature corolla tubes mostly (6-)8-10 mm long; pedicels mostly 20-30 mm long; n. 
Coahuila var. muzquizana 
Mature corolla tubes mostly 4—5(-8) mm long; pedicels mostly 5-20 mm long; s. 
Coahuila, Nuevo Leon var. palmeri 


TERRELL, TAXONOMIC REVIEW OF ACEROSA AND H. PALMERI 921 


APPENDIX 1 
Collections cited in Table 1. All collections in herbarium US. 


MEXICO. San Luis Potosi: Charcas, Lundell 5048; road between Doctor Arroyo, N.L.and Matehuala, 
S.L.P, Nelson 4533; Minas de San Rafael, Purpus 5013; Charcas, Whiting 911. Coahuila: 22 km.ESE of La 
Cuesta del Plomo on Muzquiz-Boquilla Hwy., Chiang et al. 7550Q ; 25 mi S of Monclova, Johnston 
7202; Saltillo, Palmer 126; 27 mi S of Saltillo, EAE 353; 40 mi S of Saltillo, Palmer 400; 25 mi S of 
Monclova, Palmer 401; 100 mi N of Monclova, Palmer 402; Saltillo, Pennell 17264; Saltillo, Pennell 17292; 
along rt.57, 14 mi S of Castanos, 8.5 mi S of ‘a ict of 57 and 53, Reveal et al. 2602; 11 mi S of Saltillo, 
Shreve 8540: 10.5 miE of Saltillo, E.& B. Terrell 4431.U.S.A. Texas. Pecos Co.: 11 mi S of Fort Stockton, 
Lewis 5517. Terrell Co.: Sanderson, Orcutt 734; near Feodora, £. J. Palmer 33530; Sanderson, Wooton 
s.n., 9/29/11.Val Verde Co.: Comstock, E. J. Palmer 11057; vic. Del Rio, Rose 17986. 


ACKNOWLEDGMENTS 
I a BLL. Turner for loan of TEX and LL specimens, and the curators of US for 
d research facilities. lam grateful to Piero Delprete, Steven 
Hill, and Joseph Kirkbride for very helpful reviews. Christina Kirkbride and 
Paul Peterson provided the Resumen. 


REFERENCES 


Bremekamp, C.E.B. 1952. The African species of Oldenlandia L.sensu Hiern and K.Schumann. 
Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. 48:1—297. 

Bremer, B. and J.-F. MANeN. 2000. Phylogeny and classification of the subfamily Rubioideae 
(Rubiaceae). PI. Syst. Evol. 225:43-72. 

Fernato, M.L. 1950. Gray's Manual. 8th Ed. American Book Co, New York. 

FoseerG, F.R. 1943. The Polynesian species of Hedyotis (Rubiaceae). Bernice B. Bishop Mus. 
Bull. 174:1-102. 

Gteason, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United 
States and adjacent Canada. 3 vols. New York Botanical Garden, New York. 

Gteason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada. 2" ed. New York Botanical Garden, New York 

Greuter, W., J. MCNEILL, E.R. Barrie, HM. Buroer, V, DemMouLIN, T.S. Ficcueiras, D.H. Nicotson, P.C. Sitva, 
J.E. Skos, P. TrReHane, N.J. TurLano and D.L. HawskwortH (eds. and compilers). 2000. Interna- 
tional Code of Botanical Nomenclature (Saint Louis Code). Regnum Veg. 138. Koeltz 
Scientific Books, KOnigstein 

Harford, D.A. 1992. Review of the genus Olden/andia L. (Rubiaceae) and related genera in 
Australia. Austrobaileya 3:683-722. 

KicHNn, M. 1986. Karyosystematic studies on Rubiaceae: Chromosome counts from Sri Lanka. 
Pl. Syst. Evol. 154:213-223. 

Lewis, W.L. 1968. Notes on Hedyotis (Rubiaceae) in North America. Ann. Missouri Bot. Gard. 
55:31-33. 

Rosinson, B.L.and M.L. FerNatp. 1908. Gray’s New Manual of Botany. Seventh ed. American 
Book Co, New York. 


922 BRIT.ORG/SIDA 19(4) 


Terrett, E.E. 1975. Relationships of Hedyotis fruticosa L. to Houstonia L. and Oldenlandia L. 
Phytologia 31:418-424. 

Terrett, E.E. 1979. New species and combinations in Houstonia (Rubiaceae). Brittonia 
31:164-169. 

Terrewt, E.E.,W.H. Lewis, H. Rosinson, and J.W. Nowicke. 1986. Phylogenetic implications of di- 
verse seed types, chromosome numbers, and pollen morphology in Houstonia 
(Rubiaceae). Amer. J. Bot. 73:103-115. 

Terreit, EE. 1990. Synopsis of Oldenlandia (Rubiaceae) in the United States. Phytologia 
68:1 25-133. 

Terreit, EE. 1991. Overview and annotated list of North American species of Hedyotis, 
Houstonia, Oldenlandia (Rubiaceae) and related genera. Phytologia 71:212-243. 

Terreit, EE. 1996a. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monograph 
48:1-118. 

Terrett, E.E. 1996b. Taxonomic notes on Texan and Mexican species of Hedyotis and 
Houstonia (Rubiaceae). Phytologia 81:108-114. 

Turner, B.L.1995a. Taxonomy of the Hedyotis acerosa complex. Phytologia 79:83-88. 

Turner, B.L.1995b. Taxonomic study of Hedyotis palmeri (Rubiaceae). Phytologia 79:89-92. 

Turner, B.L. 1997, Rebuttal to Terrell’s taxonomic notes of Turner's treatment of Texan and 
Mexican Hedyotis. Phytologia 82:82-85. 

Verocourt, B. 1976. Oldenlandia, Rubiaceae. In: Flora of Tropical East Africa, R.M. Polhill, ed. 
Crown Agents, London. 1:268-315. 


NOMENCLATURAL CHANGE IN THE DIGITARIA COGNAIA 
COMPLEX (POACEAE: PANICEAE) 


Joseph K.Wipft 


Herbarium 
Pure Seed nonin Inc. 
PO.B 


Hubbard, a ae USA. 


ABSTRACT 


Digitaria pubiflora is raised to specific rank. 


RESUMEN 
Digitaria pubiflora se eleva al rango especifico. 
During the preparation of Digitaria A. Haller for the Manual of Grasses for North 
America and after a reevaluation of the subspecies of D. cognata (Schult.) Pilger, 
it was concluded that the elevation of Digitaria cognata subsp. pubiflora (Vasey) 
Wipff to specific rank was warranted. 

Wipff and Hatch (1994) recognized two subspecies of D. cognata, but a re- 
evaluation of these taxa using principal components analysis showed the two 
taxa to be distinct and easily distinguished from one another. The key morpho- 
logical characters separating the two taxa are: 1) the number of veins; 2) vena- 
tion pattern (equidistant vs. non-equidistant veins); 3) the vestiture pattern of 
the lower lemmas; and 4) spikelet shape. Also, in the area of sympatry the two 
taxa maintain their morphological distinctness and no putative hybrids be- 
tween the two taxa were found in the populations studied or from the 1,839 
herbarium specimens examined. A future paper will discuss the data warrant- 
ing the following new combination, which is made now for its use in the forth- 
coming Manualof Grasses for North America. 


ee pubiflora (Vasey) Wipff, comb. et stat. nov. BAsIONYM: Panicum autumnale 
ea var. pubiflorum Nae, De Cat. Grasses US: 9.1885. nom. nud_] Dept. Agric. 
Bot. Div. Bull. 8:35. 1889. Digit It.) Pilger subsp. pubifl Vi Wipff, Sida 
13120. 1988. TYPE: UNITED STATES. TEXAS. El Paso Co.: “Mts. near Paso del Nonte® May 1881, 
G.R. Vasey s.n. (LECTOTYPE: Wipff & Hatch 1994: ane ISOLECTOTYPE: US)). 


KEY TO TAXA 


‘Lbalatr lakh 


Lower lemmas 7-veined, veins non with a narrow strip 

of pubescence between the veins; 5; spikelets obovcl to broadly elliptic 2.D.cognata 
Lower lemmas 5-veined, veins equidistant; spikelets densely pubescent between the 

veins; spikelets narrowly elliptic .D. pubiflora 


SIDA 19(4): 923 - 924, 2001 


924 BRIT.ORG/SIDA 19(4) 


ACKNOWLEDGMENT 


I would like to thank Kanchi Gandhi (GH) for his review of the nomenclature. 


REFERENCES 


Wire, J.K. and S.L. Hatcu. 1994. A systematic study of Digitaria sect. Pennatae (Poaceae: 
Paniceae) in the New World. Syst. Bot. 19:613-627. 


NEOTYPIFICATION OF ENSLENIA ALBIDA AND 
A NEW COMBINATION IN AMPELAMUS FOR CYNANCHUM 
LAEVE (APOCYNACEAE: ASCLEPIADOIDEAE) 


Alexander Krings 


Herbarium, Department of Botany 
North Carolina State University 
Raleigh, NC 27695-7612, U.S.A. 

Alexander_Krings@ncsu.edu 


ABSTRACT 


D . ] 


f recognition of various climbing milkweed genera pre- 


viously submerged in Cynanchum L. Although caution regarding resurrection of unispecific 
Ampelamus Raf. has been suggested, recent acceptance has been implicit through a listing of the 
genus in a synopsis of the subtribes and genera of the tribe Asclepiadeae. As the name is currently in 
“standardized” use among weed scientist and has additionally appeared in recent drafts, a note re- 

+4 ; pitas 7 


garding the correct nomenclature is warranted. This pap | 5 

of the entity variously known as Gonolobus laevis Michx., Cynanchum laeve (Michx.) Pers., 
Vincetoxicum gonocarpus Walt. var. laevis (Michx.) Britton, Enslenia albida Nutt., and Ampelamus 
albidus (Nutt.) Britton. 


RESUMEN 


Los avances taxonomicos recientes apoyan el reconocimiento de varios géneros de algodoncillos 

trepadores incluidos previamente en Cynanchum L. Aunque se ha pedido precaucion con respecto a 

la resurreccién del género monotipico Ampelamus Raf, ha sido implicita la aceptacion reciente 

mediante un listado del género en una sinopsis de las subtribus y géneros de la tribu Asclepiadeae. 

Como el nombre esta actualmente en uso “estandarizado” entre especialistas en malas hierbas y 
t4i if; d t vetr 


ademas ha aparecido en recientes borradores j | ate de la nomenclatura 


correcta. Este articulo trata de la tipificacion y sinonimia de la entidad conocida como Gonolobus 
laevis Michx., Cynanchum laeve (Michx.) Pers., Vincetoxicum gonocarpus Walt. var. laevis (Michx.) 
Britton, Enslenia albida Nutt. y Ampelamus albidus (Nutt.) Britton. 


INTRODUCTION 


Cynanchum laeve (Michx.) Pers. was originally described as Gonolobus laevis 
Michx. in 1803. However, due to mixed elements on the type sheet, namely flow- 
ers and angled follicles of G. suberosus (L.) R. Br. (Gonolobus gonocarpus (Walter) 
Perry) and leaves of Gonolobus laevis Michx., confusion has surrounded the cor- 
rect nomenclature of this entity (see Vail 1899, Woodson 1941, Drapalik 1969 
for detailed di ions). R iling the material with Michaux’s descrip- 
tion of Gonolobus laevis, Vail (1899) accepted the corresponding elements on 
the sheet as belonging to the taxon in her time known primarily as Enslenia 
albida Nutt. or Ampelamus albidus (Nutt.) Britton and designated the appro- 


SIDA 19(4): 925 — 929. 2001 


926 BRIT.ORG/ SIDA 19(4 


priate elements on the sheet as the type for Gonolobus laevis Michx. An image 
of the specimen is available in the IDC microfiche (#6211) set of the Michaux 
herbarium on fiche 29 of volume I. 

A type specimen was not designated by Nuttall (1818) in his original descrip- 
tion of Enslenia albida on the other hand, although he clearly described (p. 165) 
the 5-parted corolla of linear-oblong petals and the entirely free, deeply bifid 
coronal segments of the species. Based on his description in the “Genera of North 
American Plants” (Nuttall 1818), it is likely that he saw at least three specimens 
in the course of the work—presumably collected from the following localities: 


“Near Shepherdstown, on the gravelly banks of the Potomac, Virginia” (U.S.A), "on the high sandy 
banks of the river Scioto” (Ohio, U.S.A), and “near Cincinnati (Ohio)’ (U.S.A). 

Unfortunately, no specimens bearing these localities, or the distinctive asterisk 
notation used by Nuttall on herbarium sheets when describing new genera or 
species, could be found at PH—the repository for the majority of types from his 
“Genera” (Pennell 1936; Stuckey 1966). In addition, no such specimens could be 
found at BM (the eventual repository for Nuttall’s private herbarium and most 
post-“Genera” collections), E, F GH, K, LIV. MANCH, MO, and NY—all potential 
collections to host Nuttall material (Pennell 1936: Stuckey 1966; Stafleu& Cowan 
1981). The herbaria of DWC, FI, and P were also contacted, but no responses 
were received. Specimens of Enslenia collected by Nuttall in Arkansas (ie. post- 
“Genera”) were found at BM, G-DC, NY, and PH, however In addition, two speci- 
mens from Ohio were found respectively at G-DC and OXE Of the two speci- 
mens residing at G-DC, the one from Arkansas was received by DeCandolle in 
1825 and the one from Ohio in 1824. Unfortunately, labels on both specimens 
are in DeCandolle’s hand and not in Nuttall’s. Thus, aside from DeCandolle’s 
word, there is no other evidence that Nuttall actually saw the specimens. On 
the other hand, there is no reason to believe that DeCandolle would forge the 
specimens’ history. Nuttall apparently passed through Ohio only three times 
in his life: Jun-Jul 1816, Oct-Nov 1818, and Mar 1834 (Pennell 1936). Interest- 
ingly, the Ohio specimen at G-DC bears inflorescences in full bloom. In the 
northern United States, the species is known to flower primarily through Au- 
gust (Gleason 1952). It is unlikely that flowering would occur in mid-Oct 
through November or in early March. Thus, it can be deduced that the material 
is likely part of Nuttall’s original collection of the species in the summer of 
1816, prior to his later description of the novelty in his “Genera.” In the absence 
of any other material with labels in Nuttall’s own hand, the Ohio specimen at 
G-DC is here chosen as the neotype of Enslenia albida Nutt. It should be noted 
however, that the Ohio collection at OXF may also belong to Nuttall’s original 
1816 collection. Aside from the species name, the OXF specimen label bears the 
locality (“Ohio”), “com. Nuttall,” and a year which I could not completely make 
out (“1820”). The specimen also bears an inflorescence and flowers, leading to 


KRINGS, A NEW COMBINATION OF AMPELAMUS 927 


the same deduction concerning its potential year of collection as the Ohio speci- 
men at G-DC. 

Unfortunately, due to an earlier homonym (ie., Enslenia Raf., Fl. Ludovic. 
1817), the generic name Enslenia was unavailable for the climbing milkweed 
taxon named by Nuttall. Rafinesque (1819) instead suggested that Ampelamus 
(Ampelos [opnedo€] = vine, Psammos [yooppol] = sand) be used. Britton (1894) 
first validly published the requisite combination for the species name. Unfor- 
tunately, Britton (1894) spelled the genus “Ampelanus.” This spelling was fol- 
lowed by Vail (1899) and appears again in the key to the Asclepiadaceae in 
Gleason (1952), although not in the description of the genus and species. How- 
ever, as Britton (1894) explicitly noted that he made the combination “to main- 
tain Rafinesque’s very suggestive name,” most recent authors (e.g, Woodson 1941; 
Shinners 1964; Liede 1997a, 1997b) have recognized “Ampelanus” as an ortho- 
graphic error to be corrected to Ampelamus. 

Numerous authors have treated the respective names in Gonolobus, 
Enslenia,and Ampelamus in synonymy within Persoon’s 1805 combination of 
Cynanchum laeve (Michx.) Pers., based on Michaux’s Gonolobus laevis (see 
Woodson 1941; Shinners 1964; Radford et al. 1968; Correll & Johnston 1970). 
However, recent taxonomic advances are supportive of recognition of various 
climbing milkweed genera previously submerged in Cynanchum L. (Liede 
1997a, 1997b; Liede & Meve 1997). Although Liede (1997a) suggested caution 
regarding resurrection of Ampelamus Raf., she implied acceptance through list- 
ing of the genus in a synopsis of the subtribes and genera of the tribe Asclepiadeae 
(Liede 1997b). As Ampelamus albidus is currently in use as the “standardized” 
name accepted by the Weed Science Society of America (Patterson et al. 1989) 
and has additionally appeared in recent drafts (e.g., Weakley, in prep.), a note 
establishing the correct name of the entity is warranted. Resurrection of this 
unispecific genus requires a new combination based on Gonolobus laevis Michx. 


SYSTEMATIC TREATMENT 


Ampelamus laevis (Michx.) Krings, comb. nov. Gonolobus laevis Michx., Fl. Bor-Amer. 
L119. 1803. C hx.) Pers., Syn. pl.:274.1805. Vincetoxicum gonocarpus Walt. 
var. laevis (Michx.) Britton, Mem. Torrey Bot. Club 5:26, 1894. TYPE: U.S.A. ILLINOIS: Michaux 
(HOLOTYPE: P, IDC-#6211, 1:29). 


Ensleniaalbida Nutt. Gen. N. Amer. pl. 1:164.1818. Ampelamus albidus (Nutt.) Britton “Ampelanus”), 
ull. Torrey Bot. Club 21:314. 1894. Type: U.S.A. OHIO: [1816], Nuttall (NEOTYPE: here desig- 
nated, G-DC!) 


ACKNOWLEDGMENTS 


For assistance, support, or insight, | thank Paul Fantz, the Interlibrary Loan 
Department of the North Carolina State University Libraries, Sigrid Liede, Jon 
Stucky, and Wendy Worley. In addition, for kind assistance in searching for or 


928 BRIT.ORG/SIDA 19(4) 


arranging loans or digital images of specimens in their collections, I thank David 
Chamberlain (EF), John Edmondson (LIV), Sean Edwards (MANCH), David 
Goyder (K), Stephen Harris (OXF), Peta Hayes (MANCH), Helen Hoy (E), Rob 
Huxley (BM), Fernand Jacquemoud (G, G-DC), Walter Kittredge (GH), James 
Lendemer (PH), Lindsey Loughtman (MANCH), James Macklin (PH), Serena 
Marner (OXF), Greg Mueller (F), Christine Niezgoda (F), Simon Owens (K), Dou- 
glas Stevens (MO), Roy Vickery (BM), and Emily Wood (GH). Fred Barrie and 
Eric Sundell provided thoughtful reviews of the manuscript. 


REFERENCES 


Britton, N.L. 1894. List of pteridophyta and spermatophyta growing without cultivation in 
northeastern North America: Asclepiadaceae. Mem. Torrey Bot. Club 5:263-266. 

Britton, N.L. 1894. Note on the genus Ens/enia Nutt. Bull. Torrey Bot. Club 21:314. 

Corrett,D.J.and M.C. JoHNsToN. 1970. Manual of the vascular plants of Texas. Texas Research 
Foundation, Renner 

Draratik, D.J.1969.A biosystematic study of the genus Matelea in the southeastern United 
States. Ph.D. Dissertation, University of North Carolina, Chapel Hill. 

Ewan, J. 1974. Classica botanica americana, Vol.3: Andre Michaux, Flora Boreali-Americana. 
Reprint, fascimile of the 1803 edition. Hafner Press, New York. 

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

HARRIMAN, N.A. 1981. A generic index to the IDC microfiche (#6211) of the Michaux Her- 
barium arranged according to the alphabetical generic index of Michaux's Flora Boreali- 
Americana, facsimile of the 1803 edition, with many added generic names. Published 
by the author, Oshkosh. 

Lieve, S.1997a. American Cynanchum (Asclepiadaceae)—A preliminary infrageneric classi- 
fication. Novon 7:172-181. 

Liebe, S. 1997b. Subtribes and genera of the tribe Asclepiadeae (Apocynaceae, 
Asclepiadoideae)—A synopsis. Taxon 46:233-247. 

Liebe, S. and U. Meve. 1997. Some clarifications, new species, and new combinations in 
American Cynanchinae (Asclepiadaceae). Novon 7:38-45. 

PeNneLt, F.W. 1936. Travels and the scientific collections of Thomas Nuttall. Bartonia 18:1—51. 

Nurtatt, 1.1818. The genera of North American plants, and a catalogue of the species, to 
the year 1817. Printed for the author by D. Heart, Philadelphia. 

Persoon, C.H. 1805. Synopsis plantarum. Parisiis Lutetiorum [France]: apud C.F. Cramerum. 

Patterson, D.T., J.F. Avex, G.A. Bozarth, C.T. Bryson, J.W. Everest, E.P. Fun, F. Forcetta, D.W. HALL, 
H.F. Harrison, Jr., L.W. Henorick, L.G. Hou, D.E. Seaman, V, Sorensen, H.V. Strek, and R.H. WALKER. 
1989. Composite list of weeds, revised 1989. Weed Science Society of America, 
Champaign. 

Ravroro, A.E., H.E. AHies, and CR. Bett. 1968. Manual of the vascular flora of the Carolinas. 
University of Chapel Hill Press, Chapel Hill. 


KRINGS, A NEW COMBINATION OF AMPELAMUS 929 


Rarinesque, C. 1819. Nuttall’s North American plants. Amer. Monthly Mag. & Crit. Rev. 4:192. 

SHinNeRS, LH. 1964. Texas Asclepiadaceae other than Asclepias. Sida 1:358-36/7. 

Stateu, LA.and R.S. Cowan. 1981.Taxonomic literature: A selective guide to botanical pub- 
lications and collections with dates, commentaries and types, Vol. Ill: Lh-O. Bohn, 
Scheltema, and Holkema, Utrecht. 

Stuckey, R.L. 1966. Thomas Nuttall’s 1816 Ohio Valley plant collections described in his 
“Genera” of 1818. Castanea 31:187-198. 

Vai, A.M. 1899. Studies in the Asclepiadaceae, IV. Bull. Torrey Bot. Club 26:423-431. 

Weaktey, A.In prep. Manual of the flora of the Carolinas and Virginia. Unpublished, draft-in- 
progress, courtesy of the author, Chapel Hill. 

Woooson, R.E. 1941. The North American Asclepiadaceae. Ann. Missouri Bot. Gard. 28: 
193-244. 


BRIT.ORG/SIDA 19(4 


Book NOTICE 


WarreN L. WaGNeRr, Derr at R. Hersst, and S.H. SoHMER. 1999, Manual of the Flowering 
of Hawai'i, Revised Edition, (2 volumes). (ISBN 0-8248-2166-1, hbk.). Published 
in association with Bishop Museum Press, University of Hawai'i, Press, 2840 
Kolowalu Street, Honolulu, HI 96822-1888, uhpbooks@hawaii.edu, 808-956- 
8697). $95.00 (2 vols.), 1919 pp, illustrated. 


The 1999 volumes update the first edition (1990). “Over 180 additional naturalized s pecies have been 
documented. ... “Thirty-three new taxa of native plants have been discovered. ... Field studies have 
resulted in the rediscovery of 27 species presumed extinct, the resurrection of 25 taxa relegated to 
synonymy in the Manual, and more precise information on the condition of much of the endemic 
and indigenous flora.” “Species added in the past eae bring the totals to 152 families [vs. 146 it 
edition 7 737 genera [vs. 649], and 2089 species [vs. 1817].” 

f the ‘revised-ness’ of the 1999 edition is in the extensive Supplement, which includes 
‘eaueen on newly described species, nomenclatural changes, and new island records, a bibliog- 
raphy of important taxonomic references published since the first edition, and a new preface. The 
text has been modified where corrections would not alter pagination of the first edition—primarily 
an updating of the status of 271 species newly listed as endangered or threatened. The types of infor- 
mation included in the Supplement are: “nomenclatural (7) and taxonomic name changes (120); 
urrections of taxa from synonymy (25); new native taxa (33); records of naturalized species previ- 
ously nanan (108); records for naturalized species previously known only in cultivation or 
as adventives (75); new island (233) or significant intraisland range extensions of both naturalized 
and native species (13): the rediscovery of species presumed extinct (27); correct names for plants 
previously misidentified (25), particularly in the Manual; changes in distributions status (5): and a 


few miscellaneous notes.” 
The Supplement for the revised es is leuaies asa PDF Bile at the Smithsonian Systematic 


Biology /Botany web page, http://w iny.htm] under Research.— 
Guy L. Nesom, Botanical Research ane of texas, 509 Pecan Street, Foi Worth, TX 76102-4060, U.S.A. 


SIDA 19(4): 930. 2001 


PRELIMINARY RESULTS TOWARD A 
REVISION OF THE AMARANTHUS HYBRIDUS 
SPECIES COMPLEX (AMARANTHACEAE) 


Mihai Costea Andrew Sanders 


University of Agronomical Sciences Herbarium 
a Dept. as and Plant Science 


Bd. Marasti, 71331, Sector 1 mie eee ria 
Bucuresti, ROMANIA Pra CA, 92521-0102, U.S.A. 
coste_amihai@hotmail.com a ee eae 
Giles Waines 
Dept.o fBo tar and P. lant Science 


University of California 
Riverside, CA, 92521-0102, U.S.A. 
giles.waines@ucr.edu 


ABSTRACT 


The present study is a review of the Amaranthus hybridus complex based on new morphological 
and anatomical characters. Two new combinations are proposed—A. hybridus subsp. quitensis and 
A. powellii subsp. bouchonii. Based on characters such as phyllotaxy, leaf traces and other morpho- 


logical characters the division of the genus into subgenera is also discussed. 


RESUMEN 


El presente estudio es una revision del complejo Amaranthus hybridus basado en caracteres 


morfologicos y anatomicos. Se proponen dos nuevas combinaciones—A. hybridus subsp. quitensis y 
A. powellii subsp. bouchonii. También se analiza la division del género en subgéneros en base a la 
filotaxis, los rastros foliares y otros caracteres morfolégicos. 


INTRODUCTION 


The genus Amaranthus L. consists of about 70 species, of which about 40 are 
native to the Americas and the rest to Australia, Africa, Asia and Europe. The 
most frequently used infrageneric classification involves 2 subgenera and was 
suggested by Sauer (1955)—Acnida (L.) Aellen ex K.R. Robertson which com- 
prises dioecious species and Amaranthus that includes monoecious species. 
Traditionally subgenus Amaranthus is divided in two sections, Amaranthus and 
Blitopsis Dumort. Recently, Mosyakian and Robertson (1996) proposed elevat- 
ing the two sections of subgenus Amaranthus to subgeneric rank, subgenus 

marantl d subgenus Albersia (Kunth) Gren & Godr. (= section Blitopsis). 
Yet, they did not provide new characters to justify their revised treatment and 
the question of the most appropriate subgeneric classification apparently has 
remained open. 


SIDA 19(4): 931 — 974. 2001 


932 BRIT.ORG/SIDA 19(4) 


In spite of the fact that it has been the object of many studies, the genus 
Amaranthus is still poorly understood, being widely considered a “difficult” 
genus. Currently, the taxonomic problems are far from being clarified especially 
because of the widespread nomenclatural disorder caused chiefly by repeated 
misapplication of names. Most of the problems are concentrated in the most 
studied group of species, the A. hybridus aggregate. Sauer’s monograph of grain 
Amaranths (1950, 1967) has solved most of the nomenclature problems and 
provided an extensive taxonomic treatment for the species involved. Yet nomen- 
clature and taxonomic confusion among these closely related taxa has persisted 
in the literature and herbarium collections, especially in Europe. Because of 
such “serious nomenclature discrepancies” Jalas and Suominen (1980) were 
forced to present a collective map for the occurrence of “A. hybridus, A. 
paniculatus, A. patulus and A. powellii” in Europe. This quotation is a good ex- 
ample of the present confused situation. One wonders what the authors under- 
stood by “A. patulus” since it is known that this name isa synonym of A. hybridus 
or is sometimes comprehended as A. cruentus (Aellen 1964). The actual taxo- 
nomic concepts in this group of species—assuming that the nomenclature prob- 
lems are solved—ranges between two different extremes. At one extreme is 
Sauer’s treatment (1950, 1967) that recognizes as species the cultivated taxa 
(Amaranthus caudatus, A.cruentusand A. hypochondriacus) and at the other is 
Greuter’s (1981, 1984) who lumps the cultivated species with their putative wild 
oe (A. ys A. hybridus and A. powellii respectively). All possible 
intermediate ions between these two opposed treatements—many of 
them abliene since the beginning of the century by Thellung (1907, 1914, 
1919) —were also used (Aellen 1959, 1964, 1972: Dostal 1950; Morariu 1952: Brenan 
1961, 1981; Gusev 1972; Ehrendorfer 1973; Townsend 1974, 1985, 1988: Carretero 
1979, 1985, 1990; Stace 1991, 1997; Lambinon 1992; Cherepanov 1995; etc.). 

The evolutionary origins of grain amaranths are still unclear. Sauer (1967, 
1976, 1993) suggested two possible hypotheses. The monophyletic hypothesis 
states that the three cultivated species originated from a single wild progeni- 
tor—A. hybridus, followed by subsequent introgressive hybridization with two 
other wild species in different regions. According to this hypothesis, the first 
domesticated species was A. cruentus, derived from A. hybridus in Central 
America, followed by the domestication of A. hypocondriacus by repeated cross- 
ing of A. cruentus with A. powellii in Mexico and of A. caudatus by crossing 
with A. quitensis. The polyphyletic hypothesis suggests that each of the three- 
grain species was domesticated separately from a different wild species. In con- 
cordance with this hypothesis, A. hypochondriacus was domesticated in Mexico 
from A. powellii, A. cruentus from A. hybridus in Central America and A. 
caudatus from A. quitensis in South America. Detailed studies of the relation- 
ship among amaranth species using cytological or molecular methods are con- 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 933 


tradictory, supporting separately both hypotheses (Pal & Khoshoo 1972, 1973; 
Hauptli & Jain 1984; Greizerstein & Poggio 1992; Greizerstein et al. 1997; Transue 
et al. 1994: Lanoue et al. 1996; Chan & Sun 1997). The taxonomic achievement 
of all these studies was that they proved the individuality of the taxa involved, 
against the lumping tendencies. 

Other remaining taxonomic questions concern the status of A. bouchonii 
Thell. and A. quitensis Kunth. The first taxon is regarded as conspecific with A. 
powellii S. Watson by Sauer (1967), Carretero (1990), Akerroyd (1993), but is 
maintained at the species level by Htigin (1987), Stace (1991, 1997), Kerguélen 
(1993), Wilkin (1992). The second taxon, Amaranthus quitensis, was synonymised 
with A. hybridus by Coons (1975, 1978) but kept separate by almost all other 
authors. 

This study is only the beginning of a revision of this difficult complex. Set 
aside for the moment are the following topics: the relationship between the 
closely related taxa, A. dubius Mart. and A. spinosus L.and their affinity with A. 
hybridus; the appropiate position of some other taxa (A. brandegei Standley, A. 
bigelovii Uline & Bray, A. viscidulus Greene, A. scariosus Benth., A. lepturus 
S. F Blake, A. celosioides H.B.K. and A. pallidiflorus F Muell) and the role of hy- 
bridization and its consequences within the A. hybridus aggregate and among 
the dioecious amaranths. 

Amaranthus caudatus, together with A. hypochondriacus and A. cruentus, 
have created a great interest during recent years as agricultural crops in many 
regions of the world, due to the exceptionally high nutritional value of their 
seeds and leaves. Hundreds of articles document the nutritional value, the 
agronomical potential, genetic resources and breeding of amaranths. For a good 
review on the nutritional value and production methods see Kauffman and 
Wagoner (1984), Kauffman and Weber (1988), Kauffman (1992). For a compre- 
hensive review on genetic resources and breeding see Brennan et al. (2000). 


MATERIAL AND METHODS 


To assess anatomical variation the taxa were sampled between 199] and 1996. 
Seeds collected from the wild flora or received from various Botanical Gardens 
were cultivated in the Botanical Garden of the University of Agronomy 
Bucharest, Romania. Generally 4 to 6 populations or accessions were used for 
each taxon involved in the study. Seedlings, young shoots, stems of different 
ages, mature leaves, fruits and seeds were liquid preserved in a 50-70% mixture 
of formalin, acetic acid and alcohol (FAA) and embedded in Paraplast or paraf- 
fin. For the study of the vascular system of the stem, plants were chosen that 
lacked sylleptic branching, to avoid examination of branch trace bundles. Typi- 
cally stems from 25 plants, for each population were serially sectioned at 10 
mm, stained with safranin and fast green, and studied with transmission and 


934 BRIT.ORG/SIDA 19(4) 


polarized light. Furthermore, 100 seedlings for each population were grown in 
order to assess the cotyledon shape and the stem anatomy in juvenile stage. 

Regarding the pericarp structure, fruits ranging from very young to ma- 
ture were examined, but characters refer only to the latter ones. Fruits were 
embedded in white paraffin and transverse and longitudinal sections of 5-7 
um thickness were stained with Toluidine Blue. 

The SEM study of fruits, seeds and pollen was carried on with a Hitachi S- 
4100 SEM at I5KV, using a Bio-Rad Sputter-Coatter SC-500. Fifty fruits and seeds 
were collected from each of the 50 individuals per population examined. The 
same number of pollen grains per taxon was used in order to assess the mor- 
phology of the pollen grains. 

Morphology (without SEM) was studied using plants collected in the field 
in different parts of Europe and United States. Loans or other collections were 
examined from the following herbaria: ARIZ, B, BH, BP. BPI, BRIT. BUAG, BUC, 
BUCA, BUCF BUCM, CAS, CL, CLA, CM, CRA, DAV, DS, DUKE, FLAS, I, IA, IAGB, 
IASI, IBE, ISC, LA, LIL, MICH, MIL, MIN, MO, MSC, NA, OKL, PRH, RB, RSA, 
SMU, SOM, TEX, UCR, UMO, US, UTEP. VAL, VALA, VAB, VDB, VE WIS, Her- 
barium University of Agronomical Sciences Timisoara, Romania: Herbarium 
Faculty of Silviculture Brasov, Romania; Herbarium Natural Sciences Museum 
Ploiesti, Romania (not listed in Index Herbariorum). 

Taking into account the confusion that prevails in many herbaria, we con- 
sidered it useful to cite the typical accessions of grain amaranth species culti- 
vated at Ames, lowa during 1984-1989, deposited now at NA (Appendix 1). Al- 
most all these plants bear Lehmann’s name and are part of the U.S. National 
Plant Germplast System. We often cited Sauer’s specimens which were culti- 
vated at Berkley, Davis, Madison, St. Louis or Winsconsin. In addition, we cited 
Grant's specimens which were cultivated at MacDonald College of Mc Gill 
University. In specimens citation we abbreviated: s.l. = same location and sf. = 
seeds from (for the cultivated specimens). 

Seed samples for most amaranth species are available to future researchers 
from a large and diverse germplasm collection of the U.S. National Plant 
Germplasm System http://wwwars-grin.gov. Also, for European material we 
recommend Gatersleben (Germany) germplasm collection. 


Characters 

Many species of the genus are greatly affected by environmental factors (nutri- 
tional elements, water availability, light conditions, injurious factors, etc.) ex- 
hibiting a great morphological variability with little taxonomic significance. 
An enormous number of such inconsistent states (mostly based on extremely 
variable characters like: plant height, branching, color of the whole plant or 
only of stems or petioles, abnormalities in the shape of inflorescence etc.) have 
received in the past various taxonomic ranks, increasing artificially the com- 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 935 


plexity of the genus. For example, in Amaranthus retroflexus more than 60 va- 
rieties, forms and subforms (Thellung 1914; Priszter 1953; Morariu 1952) have 
been described. In the present study, besides the recognized floral characteris- 
tics, emphasis is placed on less well known characters, but ones which appear 
to be more stable. 

Our understanding of variation in Amaranthus would improve if more care 
were taken in the preparation of specimens. Plants should preferably be col- 
lected when well-developed fruits and seeds are present. In addition, to improve 
chances of correct identification, a range of individuals from the same popula- 
tion anda range of flowers, fruits and seeds belonging to the same plant should 
be collected and examined. 


Leafand Stem.—The phyllotaxis and the courses of vascular bundles were previ- 
ously studied only for A.caudatus (Gravis & Constantinesco 1907), A.graecizans 
and A. hybridus (Wilson 1924). We extended these studies to the other com- 
mon species of the A. hybridus group and to eight more species from the subge- 
nus Albersia (Costea & DeMason 2001). The phyllotaxis varies within the same 
plant: the basal leaves (from the first 4 nodes) are arranged according toa 1/2 
divergence. However, the phyllotaxis of the leaves from nodes 5-21(-23) may be 
regarded as constant for each species. Phyllotaxis varies within the genus, be- 
ing predominantly 2/5 for the species of subgenus Amaranthus and 1/3 for the 
species of the subgenus Albersia. Towards the apex of the stem it may be 3/8 in 
the species of the subgenus Amaranthus and 2/5 in the species of the subgenus 
Albersia. The vascular system in Amaranthus is closed. The leaf traces consist 
of one large median bundle (M), two lateral (L) bundles, and, depending on the 
level in the stem, 2 intermediary (i) and 2-6 (1-3 orders) of marginal small 
bundles, m, m’, m”. The bundles within a leaf trace have a symmetrical, and 
characteristic, zigzag arrangement. The configuration of leaf traces can be ex- 
pressed by means of a formula as follows: m” m’m LiMiLmm’m”. Although 
the number of bundles varies along the stem, there is always a maximum num- 
ber of bundles within a leaf trace that can be found ata certain level in the stem, 
this number being constant for each species. It varies from 11 bundles in 
Amaranthus caudatus, A. cruentus and A. hypochondriacus to 9 bundles in A. 
hybridus, A. powellii and A. retroflexus, 7 bundles in A. albus, 5 bundles in A. 
blitum, A. viridis, A. deflexus, A. blitoides, A. graecizans and 3 bundles in A. 
crispus. For more detailed information about phyllotaxis, the vascular system, 
trichomes and the anatomy of the stem in amaranths see Costea and DeMason 
(2001). 

The general shape, the apex and the base of lamina and cotyledons were 
described following Hickey (1978). The leaf anatomical pattern is constant 
within the genus. Stomata in both upper and lower epidermis are anomocytic 
for all species examined—observations that disagree with Timonin (1986a,b) 


936 BRIT.ORG/SIDA 19(4) 


who found even on the same leaf anomo-, haplo-, dia-, para-, latero-, ecyclo- 
and helicocytic stomata. The indument consists of uniseriate trichomes or mixed 
multiseriate and uniseriate trichomes. Lamina is dorsiventral, exhibiting 
“Kranz” structure, without taxonomic significance within the genus, due to lack 
of variability. 

The Inflorescence.—The flowers are arranged in small and very contracted 
cymes, the first flower within each cyme, in monoecious species, being male. 
For this reason, young plants of dioecious species are often misidentified as 
monoecious species (commonly as A. hybridus or A. powellii). Exceptions to 
this rule are A. dubius and A. spinosus which have the male and female flowers 
in separate cymes (Murray 1940; Eliasson 1988). The cymes are agglomerated, 
axillary and additionally arranged in racemose or spiciform terminal, large and 
complex synflorescences. Although extremely variable, there is usually a ten- 
dency towards a morphological “type.” Therefore the inflorescence can be char- 
acterized by the number, thickness, orientation and density of branchesand thus 
can be useful for a preliminary identification of Amaranthus species. 


The Flowers.—The length of bracteoles in some species (A. hybridus and A. 
retroflexus) is variable and continuous, therefore without taxonomic signifi- 
cance. The sterile flowers of hybrids tend to be associated with longer (than 
normally) bracteoles (Brenner unpublished). Asa result of selection, grain ama- 
ranths have usually short bracteoles, but sometimes, A. hypochondriacus may 
show bracteoles twice as long as the tepals, as in the wild species. The morphol- 
ogy of the tepals and the ratio between the tepals and the fruit provide the best 
technical and quick identification characteristics. The color of the mid-vein of 
the tepals may be green (A. hybridus) or yellowish and inconspicuous (A. 
powellii) and it can be examined on dried material (on old specimens the color 
fades, but generally remains visible). 


The Pollen. —Erdtman (1966) defined the “Amaranthus” pollen grain type also 
present in other Amaranthaceae genera and even in several other 
centrospermous families (Nowicke 1993). The pollen grain is pantoporate, apo- 
lar, small (with D = 18-28um). Generally it has more than 18 sunken pores, uni- 
formly distributed and having the apertural membrane granulated. Tectum 
with granules or spinules (see also Eliasson 1988; Costea 1998a,b). Characters: 
the diameter of the pollen grain; number and diameter of pores; density of gran- 
ules or spinules on the surface of the pollen grain according to the following 
scale: high = 20-30 granules or more/ljum?; medium = 10-19 granules/um2and 
low = 2-9 granules /lum-. 


The Fruits.—a. General characters as seen under 50 x magnification or more 
(see also Klopfer & Robel 1989a; Costea 1997b). 


Fruit transverse circumscissile, indehiscent or irregularly dehiscent.—Usually 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 937 


transverse dehiscence versus indehiscence is a constant characteristic, useful 


for tion. In ies with normally circumscissil hiscence 
(A. hybridus and toa less extent A. powellii), a transition between indehiscent, 
irregularly dehiscent and circumscissile fruit may be observed in the same 
population or even on the same plant. Other characters are: the general form of 
the fruit: spherical, ovoid, ellipsoidal etc.; the form of the fruit toward the stigma 
region: abruptly narrowed toward the stigma region, a short beak being con- 
spicuous (A. cruentus) or fruit gradually narrowed toward the stigma region 
(A. hypochondriacus, A. powellii), with the apex truncate to rounded, or acute; 
the ratio between the length and the width of the fruit, position of the dehiscence 
line: in the upper half, at the middle or in the lower half of the fruit; general 
pattern of pericarp wrinkling, size of stigma branches: width at the base and 
length; position of stigma branches: erect or recurved. 

b. SEM characters of fruits.—Surface of pericarp observed with the SEM at 
300 magnification or more. To describe the patterns of the pericarp surface, 
we slightly modified the classification of Klopfer and Robel (1989a) as follows: 
Type A (“paniculatus” type, Klopfer & Robel 1989a). -Cells very irregularly 
shaped and ramified. Principal axis of the cells not more than three times longer 
than the longest secondary axis. Anticlinal walls irregularly waved (Fig. 1, A, 
B). Type B (‘bouchonii” type, Klopfer & Robel 1989a). -Cells ramified, 4-6 times 
longer than wide. The anticlinal walls Q or S-waved (Fig. 1, C, D, E). Type C 
(“patulus” and “deflexus” types, Klopfer & Robel 1989). -Cells less ramified or 
not ramified, 8-12 times longer than wide. Anticlinal walls mostly S-waved (Fig. 
1, F). Type D (‘albus” type, Klopfer & Robel 1989a). -Cells less ramified or not 
ramified 1.5-2 times longer than wide. Anticlinal walls weekly S-waved. Type E 
(‘lividus” type, Klopfer & Robel 1989a). -Cells not ramified, 3-4 times longer 
than wide with anticlinal walls straight to S-waved. The delimitation of these 5 
types is somehow arbitrary because with respect to pericarp surface features, 
the species of subgenus Amaranthus are more variable than was suggested be- 
fore (Klopfer & Robel 1989a). Almost all species examined in the hybridus com- 
plex present, even ona single fruit, a continuous transition between two types. 
Therefore, the character of the surface patterns should be regarded more as a 
tendency toward a type. Even so, in some cases it is useful to separate, for ex- 
ample, the subspecies of A. powellii (subsp. powellii presents the type A surface 
character, while subsp. bouchonii shows variation between B and C types). In 
the subgenus Albersia nearly all species are characterised by a unique type of 
surface organisation, D or E (Klopfer & Robel 1989a; Costea 1998a). 

c. Structure of pericarp.-The mature pericarp has a very simple structure, 
being 2-4 layered. The epicarp and endocarp are always l-layered, while the 
mesocarp, according to the species, may be I-layered (A. powellii, A. hybridus, 
A. retroflexus) or 2-layered (A. hypochondriacus, A. caudatus and A. cruentus). 


938 BRIT.ORG/SIDA 19(4) 


I i powellii: B. Amaranthus 


Fig. 1. (Plate 1). Surf y 99 | 
retroflexus; C.A th tus; D.A hus hybridus; €.-F, Amaranthus powellii subsp. bouchonii. 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 939 


In some cases the mesocarp originally consists of a single cell layer but finally 
is crushed and becomes usually no longer distinguishable (A. hybridus). As a 
consequence, the pericarp in these taxa is apparently 2-layered. The taxonomic 
importance of pericarp structure, mechanism of dehiscence and analysis of the 
dehiscence-indehiscence character are the object of different study (Costea, 
Waines & Sanders, in ed.). 


The Seeds.—The diagnostic features of the seeds were largely neglected due to 
their small size. Descriptions of seeds were also produced by Kowal (1954), 
Klopfer and Robel (1989b), Esparza-Sandoval et al. (1996) and Costea (1997). 
The seeds of grain species are more variable, dark or light colored, the character 
being controlled by multiple alleles and additional loci (Kulakow et al. 1985; 
Kulakow 1987; Kulakow & Jain 1990). The following features (including the 
sculpture) should be observed under 50x magnification or more (Fig. 2, A): The 
shape of the seeds as seen from above, is variable between species but more or 
less constant within a species: circular, elliptic, obovate, etc. As seen laterally, 
the seeds usually exhibit a lenticular shape. The seeds are usually differenti- 
ated into a central, convex zone and a marginal, plane zone, exceptions to this 
rule being rare and diagnostic. This shape allows the seeds to float and disperse 
by water. The size—defined by the length of the two diameters—is constant when 
the plants grow in the same environmental conditions. Otherwise, variations 
of 0.1-0.2 mm from the average are possible. Appearance of the seed base, where 
the hilum and radicle are located: the tip of the hilum surpasses the tip of the 
radicle (subgenus Amaranthus, exception—A. hybridus) or the tip of the hilum 
is below the tip of the radicle (subgenus Albersia) (Kowal 1954; Costea 1997c). 
The presence between the hilum and the radicle, toward the center of the seed, 
of a furrow; a concavity or a concavity continued by a furrow can be also an 
useful additional character. The shape of the seed margin observed from profile 
may be acute (usually), rounded or truncated (rare). The sculpture of the seeds is 
the result of the concentrically distribution of the cells from the exotesta. The 
following variations are possible: the sculpture is evident only on the marginal 
zone, in the central zone being inconspicuous (usually); both the central and 
the marginal zone are sculptured; both the central and the marginal zone are 
almost smooth, and the central zone is sculptured while the marginal one is 
smooth or distinctly wrinkled. The color of the seeds is commonly dark-brown 
to blackish, or whitish-yellowish, sometimes with reddish nuances at the spe- 
cies cultivated as cereals. Many cultivars of A. caudatus have pink cotyledons 
visible through the seed coat. The color may be uniform or not in the last case 
usually with the marginal zone paler. 

SEM characters of seeds refer to the ornamentation of the exotesta in the 
marginal zone of the seeds, as observed under 300 x, or more magnification. 
We used the terminology proposed by Barthlott and Ehler (1977). The aspect of 


d. General view 
oe far 


th 


BRIT.ORG/SIDA 19(4) 


WeC\ A 


I hybridus: F.A 


powellii; B.S 


n 


pow 
f 
J 


ellii: D.A 


quitensis 


Fic. 2. (Plate 2).A.See 


of the see 


subsp. bouchonii; E. Pollen 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 941 


the anticlinal (prominent or inconspicuous) and periclinal (flat, concave or 
convex with the sculpture of the epicuticular waxes smooth or punctiform) 
walls were noted. 

All the above mentioned characteristics are useful for the taxonomy of the 
genus but difficult to use for the current identification of taxa. For this purpose 
we recommend the same identification keys surveyed by Brenner et al. (2000) 
which are mostly based on the traditional characters. 


Hybrids 

Hybridization is the main source of the taxonomic problems within the A. 
hybridus aggregate. Yet the frequency of hybridization within populations and 
sometimes the fertility of hybrids has been over estimated (Priszter 1949, 1958). 
In fact, experimental hybridizations undertaken (Murray 1940; Greizerstein & 
Poggio 1992) showed that such events occur infrequently and that the Fl plants 
have a reduced fertility (usually 80-98%, rarely 60%). The mean outcrossing 
rate for the cultivated species is 3.5 to 34 % (Jain et al. 1982; Hauptli & Jain 1985, 
Agong & Ayiecho 1991; Espitia-Rangel 1994). Not surprising|y, considering that 
often amaranths grow in large, mixed species populations, introgression does 
occur, the result being formation of hybrid swarms. Hybrid plants are often 
difficult to ascribe to one of the species, or even to associate with the correct 
parental species. In the temperate regions the phenomenon commonly involves 
the three weedy species—A. retroflexus, A. hybridus and A. powellii. Gene flow 
may also occur between the domesticated species (A. hypochondriacus, A. 
cruentus and A. caudatus) or between the domesticated species and their wild 
relatives. The last situation occurs frequently in the regions where grain ama- 
ranths are cultivated and the related weed species grow in the vicinity (for ex- 
ample in Mexico, Ecuador, Guatemala, Peru, and Africa). In such regions the 
species limits blur resulting in an extremely difficult complex. Often, the pat- 
tern of variation is impossible to explain if we take into consideration only the 
“classical” species, suggesting that some other local (unknown ?) taxa are in- 
volved too. A possible explanation of this variability in some areas could be 
that outcrossing rates are higher in certain environmental conditions, depend- 
ing for example on the pollinators (Hauptli & Jain 1985) and probably other 
factors too. It is hazardous to evaluate such plants only morphologically; de- 
tailed studies using molecular markers are necessary in the future if the conse- 
quences of hybridization are to be properly understood. However, to date mo- 
lecular methods have not contributed much to the understanding of this 
complex of species because of their contradictory results. The attempt to evalu- 
ate the relationships between cultivated and wild amaranths, using plants pos- 
sessing a degree of introgression (or even misidentified), can lead to unrealistic 
results and the future studies must involve careful use of both molecular and 
morphological methods. 


942 BRIT.ORG/SIDA 19(4) 


The species of the subgenus Amaranthus hybridize with the species of the 
subgenus Acnida (Murray 1940; Sauer 1955) complicating even more the inter- 
pretation of the variability in A. hybridus agg. The gene pool of the grain ama- 
ranths also includes the 10 dioecious species (Brenner 1990; Brenner et al. 2000), 
the potential of variation being enormous. The consequence of hybridization 
between species of the subgenus Amaranthus and subgenus Acnida, in North 
America, isa subject that deserves future attention. The Fl hybrids within sub- 
genus Amaranthus and those between species of the subgenus Amaranthus and 
subgenus Acnida, are often not strictly intermediate morphologically between 
their two parents, having abnormal inflorescences with very dense, crowded 
branches. They can easily be recognized by the great number of densely packed 
bracteoles that subtend the often-sterile flowers. For their accurate identifica- 
tion, one has to take into careful consideration what potential parent species 
are present in the field where the hybrids were collected. Fortunately, the spe- 
cies of the subgenus Amaranthus (and Acnida ?) do not hybridize with the spe- 
cies of the subgenus Albersia (Priszter 1949, 1958). 


TAXONOMIC TREATMENT 


The division of the genus into three subgenera—Acnida, Amaranthus and 
Albersia—suggested by Mosyakin and Robertson (1996), appears to be most 
appropriate because it would permit revision at the section level of the hetero- 
geneous subgenus Alhbersia (= sect. Blitopsis Dumort.) already started by the 
two authors. We support this classification with new morpho-anatomical char- 
acters. Also the fact that hybrids between the Subgenus Amaranthus and the 


Subgenus Albersia are unknown confirms the separation of Albersia as a sub- 
aoe 
. Dioecious ae Subgenus Acnida 


. Monoecious p 
Zz alee ea) 2/5; largest leaf traces in stems with 9-11bundles; 
seeds with hilum at or above the radicle leve panied lander 


ranthus) 


2. Phyllotaxis predominantly 1/3; largest leaf traces in stems with 3,5, or 7 ae 
seeds with hilum beneath tip of the radicle Subgenus Albersia 
= Subgenus Amaranthus section Blitopsis Dumort.) 


Within the subgenus Amaranthus, we support the treatment of Sauer (1950, 
1967) in maintaining the cultivated taxa distinct as species from their supposed 
wild progenitors, supporting it with new morphological and anatomical evi- 
dence. We do not claim that combination may not be necessary in the final re- 
vision of this difficult group. However for the moment, lumping appears to be a 
too simple and arbitrary solution. Until the day that the variability and the re- 
lationships between all the taxa involved will be understood we consider it more 
appropriate to maintain them as separate species. 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 943 


1. Amaranthus caudatus L., Sp. Pl. 990. 1753. Type: “Habitat in Peru, Persia, Zeylonia’: 
LINN 1117/26 
Amaranthus mantegazzianus Passerini, Ind. Sem. Hort. Bot. Parma 4. 1865. 
Amaranthus edulis Spegazzini, Physis (Buenos Aires) 3:163. 1917. 
Stout erect to arching annual up to 1-1.5(-2) m high. Cotyledons narrow-elliptic, 
3 x 4-5 mm, with acute apex and base; petiole about 10 mm long. Leaves rhom- 
bic-ovate to elliptic. Trichomes multicellular with uniseriate cells. Inflorescence 
large and showy, with the terminal part 10-30(-40) cm long, thick, pendent to 
erect, usually red or purplish, rarely white or yellow. Sometimes the inflores- 
cence is similar to A. hybridus, consisting of many lateral, more or less perpen- 
dicular branches. Bracteoles 3-4 mm long, about 1.2-L.5 times longer than the 
tepals, usually not exceeding stigma branches. Tepals 5, equal, outwardly curved 
or erect, 1.9-2.5(-3) mm long, obovate to broadly-spathulate, mucronate, over- 
lapping each other, with the mid-veins uniform, yellowish-brown. Fruit circum- 
scissile dehiscent, longer than the tepals, 1.5-2.5 mm long, a little longer than 
wide, ovoid, weakly longitudinally wrinkled, with the dehiscence line in the 
lower half, gradually or abruptly narrowed toward stigma region. Pericarp sur- 
face is type A. Seeds 1.3-L5 x 125-135 mm, round to asymmetrically-round. 
Between the hilum and the radicle a furrow extends almost to the middle of the 
seed. The seeds are differentiated into central and marginal zones. Two distinct 
types of seeds exist: a) whitish-yellowish in color, ovoid in lateral view, with 
marginal zone smooth or irregularly wrinkled, often with the pink embryo vis- 
ible through the seed coat, and the central zone conspicuously sculptured; the 
border of the seed rounded or truncated; cells of the exotesta in the marginal 
zone elongated (30-50 pm length) with prominent anticlinal walls and 
periclinal walls plane and verrucose (Fig. 2, B). b) dark-brown seeds, lenticular, 
with marginal zone sculptured and central zone smooth; uniformly colored; 
cells of the exotesta as in the whitish-yellowish seeds but the periclinal walls 
smooth; border of the seeds is acute. Pollen grains 22-24 um with 30-40 pores 
of 1.5-1.8 um diameter. Density of granules medium. 2n =32. 
The relationships within A. caudatus were studied by Coons (1975, 1982) 
and we share that author's view of infraspecific classification 
Distribution and ecology.—In Europe, A. caudatus is grown primarily asan 
ornamental, and rarely escapes from cultivation in waste places. A.caudatusas 
a grain crop, originated at high elevations in South America, in northwestern 
Argentina, Ecuador, Peru and Bolivia, the closest wild taxon being A. hybridus 
subsp. quitensis. In the native areas you can often find transitional forms be- 
tween the two taxa. The question remains whether A. cruentus also participated 
in the development of A. caudatus. Using the data from nuclear DNA and re- 
striction-site chloroplast variation, Lanoue et al. (1996) found that A. caudatus 
and A. cruentus are more closely related to each other than to their respective 


944 BRIT.ORG/SIDA 19(4) 


putative progenitors. Chan and Sun (1997) stated also that A. caudatus is more 
closely related to A. cruentus, and both of them nearer to A. hybridus than to A. 
quitensis. These data are not confirmed by cytological studies (Pal @ Khoshoo 
1972, 1973; Hauptli & Jain 1984; Greizerstein & Poggio 1992; Greizerstein et al. 
1997). Also, other mollecular studies, using RAPD markers indicate that A, 
caudatus is more closely related to A. hypochondriacus than to A. cruentus 
(Transue et al. 1994). Sauer (1967) stated that when growing together with the 
other two cultivated grain species, A. caudatus does not usually form hybrids 
and crossing barriers in A. caudatus have also been observed by Coons (1975, 
1982). However, among the accessions cultivated at Ames, lowa between 1984- 
1989 and deposited at NA, many of the plants impossible to ascribe toa definite 
species were obviously related morphologically to A.caudatus. In all cases such 
plants have obovate to spathulate tepals, only partially overlapping each other, 
and narrower than in typical A. caudatus but wider than in any other species, 
suggesting a degree of introgression. Two morphologic types are commonly 
encountered: 

1) The plants from Morelos, Mexico identified as “A cruentus” by Lehmann 
or “A. hybridus subsp paniculatus” by Spjut, apparently represent a definite land 
race probably involving A. cruentus and A. caudatus [(Spjut 8808, 8810, 8821; 
Lehmann: AMES 5501 Clype Land 2), 5502, 5179, 5182, 5183, 5188, 5193, 5195, 5196, 
5197 (Type land 2) 5198 (Type 1 and 2), 5200, 5201, 5202, (NA)I. Besides this rec- 
ognizable type, many other accessions bear morphological “traces” of A. 
caudatus but there are more heterogenous suggesting either the participation 
of A. cruentus or A. hypochondriacus, or both, in their origins. 

2) The plants from Ecuador identified as “A. hybridus” by Lehmann or “A. 
hybridus subsp. quitensis” by Spjut, probably involve A. caudatus and A. 
hybridus (Ecuador, Lehmann, PI 490663, PI 490664, PI 490666, PI 490667, PI 
490670, PI490716, PI 490718, PI1490719, PI 490722, P1 490672, PI 490715, PI 490723, 
Pag 24, P1907 5.214907 20, Pl 4907 28, Pl 90730, PLA90 731, P1407 32, Pl 
HT: FP e9U OT, FLAT Z, F 1490/40, PL 490747, Fl4 90/46, Fl 490752. These 
specimens are different from the red Ecuadorian “sangorache,” another example 
of taxonomic incertitude in grain amaranths. Sauer included it within A, 
quitensis and Brenner et al. (2000) suggest the same. Coons accepted it as a va- 
riety of A. hybridus (s.l, including. A. quitensis) (1975, 1978). As noted by Coons, 
even if closely related to A. caudatus and A. hybridus Gincl. A. quitensis) and toa 
less extent with A. cruentus, “sangorache” presents some unique features such 
as the tepals position and the form of the fruits. We also think that it deserves 
separate recognition—at least as a variety (if not as a subspecies)—but its posi- 
tion in relation within the species complex requires further investigation. 


Representative specimens examined. AFRICA. ETHIOPIA. Harege Prov.: Harar, escaped from culti- 
vation, 9°5'N, 42°17°E, 7 Sep 1963, Burger 3237 (US). 20 km from Harar Leprosarium “Besadimo,”" 1425 
m, 3 Aug 1967, E. Westphal & Westphal-Stevels 994 (MO). IIlubabor Prov.: 8 km SW Teppi, 7°9'W, 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 945 


35°18'E,1300 m,9 Dec 1964, Meyer 8995 (NA). Kefa Prov.: E of Jima, Oct 57,Anderson s.n. (WIS).Shoa 
ieee : Addis Abeba, Hugh Rouk, 12 Jan 1965, Meyer 9093 (NA). KENYA. Eastern Prov.: Mount Kenya, 
m, 21-27 Sep 1909, Mearns 1377 (A). SOUTH AFRICA: Pretoria-Riviera, cultivated, 1 Jun 1964, 
eae ae 9983 (A). 
ASIA. ISRAEL: Sharon Plain, Herzliyah, near Yarkon bridge, 21 Apr. be Zohary & Grizi 730 
(A, BH, FLAS, MO, NA). NEPAL: sf Marku Valley, Sirsagarhi,“grain crop,” 30 Jan 1951, Sauer 1 ae IS). 
s.f. the SW Slopes of Annapurna, ‘cultivated as grain,” 28°20'N, 83°45’, na m, 1 Nov 4, Sauer 
1776 (WIS). PAKISTAN. Ladak Prov.: Indus Valley, Leh to Kaltse, 12-14 Jul 1856, eae 1486 
(GH). INDIA: s.f. unknown locality, collected in Aug 1910 and cultivated at New York, Arnold s.n. (BH). 
Tamil Nadu: s.f. Madras, 23 Sep 1964, Sauer ae (NA, WIS); 5.1, 30 Aug 1964, Sauer 3962 (MO, NA, 
WIS); s.L., 16 oe 1964, Sauer 3964 a (WIS). CHINA: s.f. Yunnan, 3 Sep 1940, Cowgill 2087 (BH). s.f. 
Mowhsien, Szec n, 2000 m, “cultivated crop,” 9 Jan 1950, Sauer 1503 A (WIS). JAPAN. Hondo: 
ase cote "26. es oe manne het (C 
cao B ical Garden, Grant 719 (WIS). ENGLAND: 
sf. Pane cl Aneee at New a Ithaca, 7 Aug 1960, Bailey 8417 (BH). FRANCE. Doubs: 
s.f.Besan | Garden of the University of Besancon, Grant 718 (WIS). Loire-Atlantique: s.f. 
Nantes, cone Garden of Nantes, Grant, 216, 217 (WIS). ROMANIA. Constanta Co.: Medgidia, 
“cultivated as ornamental and ruderal,"15 Sep 1995, Costea s.n. (B 
NORTH AMERICA. UNITED STATES: CALIFORNIA. Santa Barbara Co.: Santa Barbara, 22 Jan 
1958, Pollard s.n. (CAS). San Bernardino Co.: Rialto, 1 Oct 1933, Wheeler 2141 (DS). Ventura Co.: 
Casitas Pass Rd., Foster Park, 23 Oct 1965, Pollard s.n. (CAS). Santa Clara Co.: Palo Alto, 22 Jan 1958 
Pollard s.n. (CAS).18 Sep 1968, McClintock s.n. (NA). FLORIDA. Alachua Co.: Gainesville, 14 Sep 1953, 
West s.n. (FLAS). ILLINOIS. Chicago (cultivated), 13 Jul 1958, Pollard s.n. (DUKE, FLAS); 5.1, 13 Jul 1987, 
Plowman 14507 (US). MINNESOTA. Stearns Co.: St Cloud, 807 S,6"™ Avenue, 26 Sep 1991, Lindstrom 
1997 (KSC). NEW YORK. Tompkins Co.: Ithaca, 28 Sep 1936, Allen 6397 (BH);s..,11 Sep 1914; 5.1, 29 
Jul 1920;s.1.,,20 Oct 1920;5.1.,17 Sep 1924: 5.|.,23 Jul 1928, Bailey s.n. (BH); s.1.,3 Aug 1948, Dress 1405 
(BH, NA);s.1,31 Aug 1948, Dress 1232 (BH, NA). VIRGINIA. Buchanan Co.: Upper Prater, 11 Aug 1988, 
Churchill 88234 (MSC). WISCONSIN. Dane Co.: Madison, Oct 1924, Davis s.n. (WIS). 
SOUTH AMERICA. PERU. Dep. Ayacucho: Prov. Cercado, Ayacucho, 2800 m, 10 Mar 1964, Gade 
s.n. (WIS). Dep. Cuzco: Urubamba, 21 Apr 1915, Cook & Gilbert 256 (NA). Paruro, 10 May 1964, Gade s.n. 
(WIS). Dep. fara ccpees Mejorada, 31 Oct 1948, Sauer 1283D (WIS). BRASIL. Mun. Curtiba: Parana, 
“ornamental,” 26 Jul 1980, Kammrow 1368 (WIS). BOLIVIA. Prov. Cochabamba: Cochabamba, 31 
Oct vee a ee 1303 (DAV, DS,GH, NA, ne} 5.1, 12 Dec 1948, Sauer 1179 (MO, DAV, DS, GH, MO, 
NA, WIS).s.f. Cochabamba, grown at Botanical Garden Illinois, March-May 1948, Fuller s.n. (NA). Prov. 
La Paz: s.f.Sacaba, 19 Nov 1959, seen a WIS).s.f. Tajma, 17 Dec 1959, Sauer 2542 (NA,WIS).s.f. 
Chulumani, Nov 1959, Sauer 2539 ARGENTINA: cultivated, 20 Apr 1942, Hunzinker 2083, 2083 
Bis (BH). Prov. Salta: Dep La Vina, eee de Dias, cultivated, 1200 m, 1941, Hunzinker 1321 (A). 


— 


— 


2. Amaranthus cruentus L., Syst. Nat. ed. 10, 1269. 1759. (Fig. 3). Type: “CHINA”, LINN 
117/19. A. hybridus L. subsp. cruentus (L.) Thell. var. paniculatus (L.) Thell., Fl. Adv. Montpell. 
205.1912. A. hybridus L.su ade Thell proles paniculatus(L) Thell., Ascherson & 
Graebner, Syn. Mitteleur. Fl. 5:247. 1914. A. hybridus L. subsp. paniculatus (L.) Hejny, Dostal, 
Kvetevna CSSR. 444. 1950. A. a ee iar Ti ex Gren. & Godr.) Brenan 

var. cruentus Mansf., Die Kulturpflanze 2 
Amaranthus Sanniog Sp. Pl. ed. 2, oe 1763. 
Amaranthus sanguineus L. p.p., Sp. Pl. ed. 2, 2:1407. 1763. 
? Amaranthus chlorostachys Willd, Hist cae 34.1790. 


o 


Erect annual 0.5-1.5 m, usually reddish throughout. Cotyledons lanceolate to 
narrow-ovate, 16 x 5-6 mm, with rounded apex and cuneate base; petiole about 


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Amaranthus cruentes L 


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CULTIVATED PLANTS 


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Fic. 3 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 947 


7 mm long. Leaves rhombic-ovate to broadly-lanceolate, 4-15 x 2-18 cm. Tr 
chomes multicellular with cells uniseriate. Inflorescence usually with many 
lateral, perpendicular, thin branches. Bracteoles 2-3 mm long, equalling or 
slightly longer than the tepals. Tepals 5, almost equal (one is approximately 0.5 
mm longer than the other 4), oblong, acute, with the mid-veins uniform, yel- 
lowish-brown. Fruit circumscissile dehiscent, 2-2.5mm long abruptly narrowed 
toward the stigma region, a short, thin rostrum being evident. Fruit about 1.5 
times longer than wide, obovate to rhombic, 2-2.5 mm long, with the dehis- 

cence line at the middle or in the upper half. The pericarp is almost smooth or 
weakly wrinkled above the dehiscence zone. Type of pericarp surface variable: 
A to B. (Fig. 1, C). Seeds obovate to elliptic-asymmetrical 1.25-L6 x 11-12 mm, 
whitish or yellowish, rarely dark-brown. The pale seeds are very much like the 
ones described for A. caudatus. The differences are that the furrow between the 
hilum and the radicle is prolonged only about 1/3 of the way through the di- 
ameter of the seeds and that the pink tint is usually absent. The dark seeds ex- 
hibit a concavity above the hilum and have both the central and the marginal 
zone (especially) sculptured. The cells of the exotesta as in A. hybridus. The 
border of the seed is acute. Pollen grains 20-25 um, with 33-45 pores of 1.9-2.1 
um; density of spinules medium. 2n = 32, 

Distribution and ecology.—Apparently originated in Central America from 
the wild species A. hybridus, asa grain crop. Sauer (1950) and Sanchez-Del Pino, 
Flores Olvera and Valdes (1999) noted transitional forms between the two spe- 
cies in Mexico but such examples also occur in Central America and Africa. 
Together with the other two grain amaranth species, it was introduced into the 
Old World by the Spaniards and then into Asia and Africa. Today it is culti- 
vated worldwide, mostly as a garden ornamental. In United States, South 
America, Zaire, Sierra Leone, Ethiopia, India and Middle East it is cultivated as 
a cereal and in many other countries it is being evaluated experimentally for 
this purpose. The commercial grain amaranth crop involves both A. cruentus 
and A. hypochondriacusas wellas hybrids between the two. In Africa A.cruentus 
is also cultivated as a potherb, bearing the name “African spinach” or “Sudan 
spinach,” or is used as animal focleioe The leaves have a high content of protein 
and vitamins. Thi pes from cultivation in ruderal ae 


db 


Representative specimens examined: AFRICA. EGYPT: Bashtil, 21 Oct 1974, Mosein & Riad Higaz) 
(MO). ETHIOPIA: Road from Arba Minch to Soddo, 108 km from Arba Minch; 1180 m, 7 Feb 1 068, : 
Westphal & J.M.C. Westphal 323b (US). NIGERIA. State North East: Distr.Gembu, Mobilla Plateau, 10 
May 1972, Gbile et al. 1337 (MO). ae Prov. Sud-Ouest: Dep. Fako, Bakingili, 4°04'N, 9°02’E, 
0-30 m, 12 Jun 1984, Thomp: 3 (MO, CM). Nkoubisson, Collection Garden of Depart- 
ment of Agriculture ENSA, 17 Dec 1976, ie C. Westphal 7375 (NA); 5.1, 18 Dec 1976, .M.C. Westphal 
9010 (NA). UGANDA. Distr. Kampala City: Botanical Garden of Makerere University, 0°17'N, 32°34'E, 
1200 m, 27 Feb 1972, Katende 1735 (MO). KENYA: s.f. unknown locality cultivated at Miami, Florida, 7 
ep 1920, Shantz 51847 (BH). ZAIRE (CONGO). Distr. Shaba: 15 sia of Lubumbashi, 22 Dec 1961, 
Schmitz 7522 (MO). BURUNDI. Distr. Bujumbura: Bujumbura, 780 m, 15 Sep 1974, Auquier 4090 


948 BRIT.ORG/SIDA 19(4) 


(MO);s.1.,800 m, May 1978, Lambinon 78/388 (MO).Rushubi, 3°22'S, 29°28'E, 1650 m, Mar 1981, Reekman 
9612 (MO). SOUTH AFRICA. Eastern Cape: Grahamstown, cultivated, 2400 m, 30 Jul 1972, Bayliss 
1377 (BH). 

AUSTRALIA AND OCEANIA. PAPUA NEW GUINEA: East Highlands Distr.: Norey Kora 
Swamp, 6°30S, 145°75'E, 1400 m, cultivated, Oct 1966, Wheeler & Anu 5857 (A). 

ASIA. INDIA: s.f unknown locality cultivated at New York, Ithaca, 15 Aug 1933, Gilmore 13600, 
15595 (BH).Pembra, Malabar Wynsod, Millet’s Garden, 800 m, 16 Apr 1963, Noble # O (NA, WIS). CHINA. 
Kwangsi: near San-t’ai-ling, 28 Aug 1937, Taam Ying-Wah s.n. (BH). VIETNAM. Annam (Trung Phan 
region): Tourane (Da Nang), Jul 1927, /. Clements & M.S. scala ones iyi Mindanao 
Island: Davao, Apr 1903, DeVore & Hoover 112 (US). INDO hera g, Pasir 
Putih, cultivated, 13 Feb 1981, Taylor NM-III P704 (A). ai one of Loemban Ria, Asahan, 5 Feb- 

12 Apr 1934, Bocea 8022 (US). Adian Rindang, vicinity of Hoeta Tomoeau Dolok, 17 Nov-10 Dec 
1935, Bocea 8710 (A). 
AUSTRALIA. New South Wales: Summer Hill,"spontaneous in garden,"May 1970, Michael M8 


5): 

NORTH AMERICA. UNITED STATES. ARIZONA. Navajo Co.: s.f. Hotevilla, Hopi Indian Reser- 
vation, 6 Nov 1950, Sauer 1343 E (WIS); 5.1, May-Aug 1959, Sauer 2530 (WIS). ALABAMA. Jefferson 
Co.: Birmingham, ruderal, 4 Oct 1968, Kral 33646 (SMU). CALIFORNIA. San Bernardino Co.: San 
Gabriel Mountains, San Antonio Canyon, Chapman Ranch above Mountain Baldy Village, 1300 m, 
Thorne & Thorne 191074 (RSA). Santa Clara Co.: Stanford University, 28 Oct 1896, Dudley 24899 (SMU). 
Palo Alto Harbor, ruderal, 11 Aug 1974, Thomas 17504 (DS).Palo Alto,along Alma Street, near Channing 
Street, 27 Sep 1961, Thomas 9791 (CAS, DAV).San Joaquin Co.: Calaveras River bottom, 17 Sep ; 
Stanford 170585 (DS). KENTUCKY: unknown locality, cultivated, 1817, Short s.n. (NA). MICHIGAN. 

Co.: near Lansing (ruderal), 13 Oct 1866, Bailey s.n. (BH). MISSOURI. Boone Co.: Columbia, 
“ruderal,"18 Aug 1933, Dimes 983 (UMO). East Higlands, ruderal,9 Aug 1933, Dimes 921 (UMO). NEW 
YORK. Albany Co.: near Londonville (ruderal), 12 Sep 1932, Muenscher 17858 (BH); 5.1.23 Sep 1 on 
Burnham ae (BH). PENNSYLVANIA. Allegheny Co.: N side of Pittsburgh, ruderal,4 Aug 1 
Buker s.n. (CM).Bucks Co.: West Bristol, ruderal, 17 Jul 1952, Long 75306 (CM).WISCONSIN. Dane ie 
Madison, "waste ground,” Oct 1938, Shinners s.n. (WIS). HAWAIL Honolulu Co.: Aina Haina, 27 Oct 
1978, Bush ate NA). 
AMERICA AND GREATER ANTILLES. MEXICO. CHIAPAS.: iin Ocosingo, 26 
Nov 1976, co pai 2869 (WIS). JALISCO.: Mpio. De Cuantilan, 19°28'13"N, 104°11'04"W, 750 m, 28 
Aug 1991 ue et al. 140 (SMU). MICHOACAN.: Patzcuoro, 2050 m, 20 se a Caballero 1106 
(WIS); ee 140 9, Caballero 1059 (WIS). OAXACA:.: sf. see Aug 59, Sauer 2529 (WIS). 
PUEB eee 30 Oct 1948, Sauer 1278 (GH, MO, NA, UC, WIS). CUBA. Ciudad de la Havana: 
nei Estacion Central Agronomica, cultivated, Baker 2843 (GH). aries Island, Vicinity of 
la ; May 1929,E.C. Leonard & G.M. Leonard 15302 (US). en ee Dep. Alta Verapaz: Coban, 
2N 950, Sauer 1265-F (WIS). Cubilquitz, 350 m, 1903, Tuerckheim s.n. (GH). De 
Be sf. Siem: 28 May 1948, ne i‘. a (MO, NA, WI); 5.1, Sauer 1759, 1183 
(DAV, NA); 5.1, Sauer 1184, 1178 (DS, GH, UC, NA, WIS); 5.1, 10 Oct 1948, Sauer 1267,1240 (MO, NA, WIS): 
5.1, 31 Oct 1948, Sauer 1240, 1279 (MO, WIS, NA);s.l.and same date Sauer 1276,1159, 1129, 1264, 1305, 
(DS, GH, NA, UC, WIS); s.l. and same date Sauer 1323, 1296 (DAV, NA, UC);5s.1., 20 Nov 1948, Sauer 1373, 
1374 (MO,NA, WIS);s.l.and same date Sauer 1367 (NA);s.1.,9 Dec 1948, Sauer 1380 (MO, WIS, NA):s.L, 
23 Dec 1948, Sauer 1240 (DAV, MO, NA, WIS); s.L, 1 Feb 1949, Sauer 1949 (MO, NA, WIS). s.f. San Juan 
Sactepequez, 18 Oct 1948, Sauer 1177 (NA, DAV); 5.1, 30 Oct 1948, Sauer 1322, 1269, 1286, 1176, 1321, 
1275, 1265 (NA, UC, WIS); 5.1.6 Nov 1948, Sauer 1265 (DAV,UN):s.1, - Sauer 1369, 1176 (DAV, UN). 
NICARAGUA. Rivas: Isla Ometepe, Volcan Maderas, 1 1°27-28'N, 85°31~32', 400-800 m, 24 Sep 1984, 
Robleto 1273 (MO). EL SALVADOR: San Salvador, cultivated, Jun i Calderon 739 (GH). 20 Dec 
1921-4 Jan 1922,“weed in garden,” Standley 19238 (GH, US). 
SOUTH AMERICA. VENEZUELA: Bolivar, on the Orinoco, 75 m, Feb-Mar 1921, L.H. Bailey & Z. 


\O 
No 


— 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 949 


Bailey 839 (BH). COLOMBIA. Dep. Antioquia: Robledo, 1560 m, 12 Sep 1965, Grisales 17 (US). Dep. 
Cesar: Becerril, Jul 1970, Ruddle s.n. (WIS).PERU. La Liberdad: Trujillo, Hacienda la Encalada, 2300 m, 
13 Feb 1957, Sagastegui 38 (US). BRASIL. Distr. Federal: Paranoa at Lagoa Paranoa Lake, 12 Dec 
1965, Irwin et al. 11239a (MO). Distr. Santa Catarina: Florianopolis, 20 m,8 Aug 1964, Klein 5353 (US). 


3. Amaranthus hybridus L., Sp. Pl. 990. 1753. Type: “Habitat in Virginia”; LINN 1117/19. 


Annual to 0.5-1.5(-2) m, usually erect. Stems weakly to densely pubescent above, 
multiseriate hairs being mixed with uniseriate hairs. Leaves rhombic-ovate to 
broadly-ovate-lanceolate. Inflorescence dark-green, either with many crowded 
slender, lateral branches, or lax, with only a few flexuous branches. Bracteoles 
2.5-4(-6) mm long, narrow, about 1.2-1.5 to 2 times longer than the tepals. Tepals 
5, almost equal, or one of them longer, 1.5-2.5(-3) mm long, with the mid-vein 
dark-green, with very variable form and length: a) narrowly-ovate to oblong- 
lanceolate, acute to subacute, erect, shorter than the fruit or b) oblong-lanceolate, 
obovate to spathulate, erect to outwardly curved, equal to longer than the fruit. 
Fruit usually circumscissile dehiscent, 1-1.5 times longer than wide, with the 
dehiscence line at the middle or in the upper half, more or less abruptly nar- 
rowed toward the stigma region and coarsely wrinkled above the dehiscence 
line. Infrequently plants with indehiscent or irregularly dehiscent fruits may 
be also encountered. Stigma branches are short, slender, erect (0.9-14 mm length 
and 0.1-118 mm width at the base). Type of pericarp surface variable: B to C 
(Fig. 1D). Seeds 1-14 x 1-1.35 mm, broad-elliptical to circular, lenticular. The tip 
of the hilum is placed below or at the same level as the tip of radicle. Seeds dif- 
ferentiated into central and a marginal zones, both sculptured. The cells of the 
exotesta are polygonal, 25-35 um, with prominent anticlinal walls and convex, 
finely sculptured periclinal ones. The border of the seed is acute. Color dark 
brown to black, sometimes with the marginal zone having a paler tint. 
Amaranthus hybridus isan extremely variable species, especially in Mexico, 
Central and South America and Africa (see also Sauer 1950). Coons (1975, 1978), 
after a meticulous study of their relationships, considered A. quitensis a syn- 
onym of A. hybridus. However she admited that one character, the inner tepal 
index, may be significant though not correlated with other characters. The de- 
scription by Kunth (1817) of the pistilate flowers of A. quitensis: “calyx 
quinquepartitus; laciniis lanceolato-oblongis, obtusis” is actually a better de- 
scription of A. hybridus. Thellung (1914) amended the description of A. quitensis: 
“tepals 5, about 2 mm long, spathulate with the apex broad-rounded (with 
mucro), with a green mid-vein continued into the mucro. Fruit shorter (rarely 
as long as) than the perianth.” We consider that both descriptions together are 
more likely to represent A. quitensis. These features induced Bolos and Vigo 
(1974) to include A. quitensis as a subspecies of A. retroflexus. However, the real 
affinities of A. quitensis are more with A. hybridus, in which we include it as a 
subspecies. The hybrid between subsp. hybridus and subsp. quitensis has a pol- 
len viability of 60% (Greizerstein & Poggio 1992), a value that shows both the 


950 BRIT.ORG/SIDA 19(4) 


affinity and the distinctness of the two taxa. Also, Greizerstein et al. (1997) 
showed significant differences in karyotype formulae and asymmetry indexes 
between the two taxa. Molecular evidence for the distinctness of the two taxa 
was provided by Chan and Sun (1997). We find the pollen of the two subspecies 
of A. hybridus easy to differentiate. 

Amaranthus hybridus (especially subsp. quitensis) stands apart from 
Amaranthus powellii and A. retroflexus in its ecology and in its flowering and 
fruiting times in temperate regions, being about 2-4 weeks later. Native to East- 
ern North America, Mexico, Central and South America, but nowa widespread 
and noxious ruderal and segetal weed. 

1. Cotyledons narrow-elliptic to elliptic with rounded or obtuse apex; most ee 
narrowly-ovate to oblong-lanceolate, acute or subacute, erect, shorter than the fru 
tectum with medium density of granules_ Ss A hybrids on hybridus 
Cotyledons lanceolate, with acute apex; most tepals spathulate to obovate, obtuse 
or truncated, almost erect or outcurved, longer (rare equal) than the fruit; tectum 
with high density of spinules A. hybridus subsp. quitensis 


3a. Amaranthus hybridus subsp. hybridus (Fig. 4). 


Amaranthus patulus Bertol. Comment. Itinere Neapol. 19. 1837. A. hybridus L. subsp. patulus 
(Bertol.) Carretero var. patulus, Collect. Bot. (Barcelona) 11:127. 1979. A. hybridus L. subsp. 
cruentus(L.) Thell. var. a (Bertol.) Thell., Fl. Adventice Montpell. 8:206. 1912. A. hybridus 
L. subsp. hypoc} Thell. “proles” cruentus (L)Thell. var, patulus (Bertol.) Thell., 
Ascherson & cata. ie ane FI. 5:244. 1914 

Amaranthus incurvatus Tim. ex Gren. & Godr, Prosp. Fl. France 8. 1846. A. hybridus L. subsp. 
incurvatus (Tim. ex Gren. & Godr.) Brenan var. incurvatus, Watsonia 4+:268. 1961 

Amaranthus chlorostachys auct., non Wild. 


Cotyledons narrow-elliptic to elliptic, 10-12 x 4 mm, with rounded or obtuse 
apex. Pollen grains 22-28 ym with 25-35 pores of 1.4-1.6 um in diameter. Den- 
sity of granules medium. (Fig. 2, E). 2n = 32, 34. 

The name A. hybridus was rejected as ambiguous by some authors because 
it was widely and persistently misapplied to A. powellii or A. hypochondriacus. 
As all these species were clearly typified, we cannot follow such a view. An- 
other name “A. chlorostachys” which was very much used to designate this taxon, 
is based ona single green plant, within a red-coloured population of cultigens 
from India (A. paniculatus WILLD 17521(B)). It appears that Willdenow de- 
scribed this mutant as a separate cultigen, which later authors, ambiguously 
but constantly synonymized either with A. hybridus or with A. powellii (espe- 
cially in Europe). 

Sauer (1950) distinguished a “northern race” and a “tropical race” of A. 
hybridus. We subscribe to this practical, non-systematic grouping, The northern 
race is the “typical” A. hybridus,a common weed in the temperate regions of the 
world that causes little taxonomic problems. Sometimes introgression with A. 
retroflexus, A. powellii or dioecious species has been documented, but usually 
individuals are easy to identify. The length of bracteoles and the inflorescence 


— 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 


ROARIUM 


3 Fr 73 


Herbarium - Universiey of California Riverside 


1 


wets hvbralas 


iy 


Mihat Costes 


dwar 


Fic. 4.4 thus hybrid t I hybridus, from McWilliams 15123 


= 


IA) 


952 BRIT.ORG/SIDA 19(4) 


are variable, continuous and not correlated. Consequently, further infraspecific 
separation of a var. (subsp.) “hybridus” with bracteoles about 2 times longer 
than the tepals and inflorescence dense with many lateral crowded branches, 
and a var. (subsp.) “patulus” with bracteoles about 1-L5 times longer than the 
tepals and an inflorescence less ramified and more lax, seems arbitrary. 

The variability of the “tropical race’—common in Mexico, South America 
and Africa—is on the other hand exceptional, suggesting in our view that some- 
times, additional taxa (unknown or other than the ones examined in this study) 
have contributed to the variability. Often these plants can be named A. hybridus 
only ina broad sense. If we use the tepal length character, such plants fall again 
into the two categories; a) plants with flowers that have the tepals longer than 
the fruit, and b) plants with flowers that have the tepals equal or shorter than 
the fruit. Within the first category, we encountered among plants from Mexico 
a distinctive variation that might deserve taxonomic recognition. It resembles 
subsp. quitensis but is obviously different, with tepals acute, mucronate, re- 
curved, and up to 2 times longer than the fruit. Such plants were identified by 
Sauer as “A. scariosus x A. hybridus ?” or “A. hybridus, atypical” and were also 
observed and cited by Sanchez-Del Pino et al. (1999). Examples: 


Chiapas: Municipio of Venustiano, Carranza, 25 Oct 1966, Laughlin 2699 (DUKE). Durango: Oct 1896, 
Palmer 759 (GH). Hidalgo: Rio Tula, 1700 m, 15 Mar 1965, Quintero 2673 (DS).Oaxaca: Valley of Oaxaca, 
1560 m,8 Jul 1897, Pringle s.n. (CM, 1A, MICH, MIN). 15 km SE of Oaxaca, 1740 m, 1 Oct 1962, Ugent & 
Flores 2628 (WIS); Upper slopes of Cerro San Felipe, 17°9'N 96°50'W, 3 Oct 1962, Ugent & Flores 2706 
(WIS). Chimalapa, 1100 m, 16°42'N 94°05'W, 23 Oct 1984, Maya 763 (MO);16°43'N 94°08'W, 14 Nov 
1984, Maya 899 (MO). Tamaulipas: San Jose, 20 Feb 1939, Le Sueur 112 (ARIZ). 


The plants from the second category are even more variable suggesting, in our 
view, the participation of other taxa than A. powellii, A. cruentus or (and) A. 
hypochondriacus. Besides the “typical” variation of Amaranthus hybridus, we 
have encountered the following situations: 


1) Plants reddish throughout. 

2) The inflorescence composed of lax monochasial cymes (ripidium) with 
the rachis and the secondary axes visible (usually cymes in Amaranthus are 
very contracted and such details cannot be easily observed). Such lax cymes 
may be arranged in thick spiciform inflorescences, resembling A. powellii or in 
very branched inflorescences, with many patent, lateral branches (as in typi- 
cal A. hybridus). 

3) The bracteoles extremely thin, acicular, spinescent, rigid, 2-4(-5) times 
longer than the flowers. 

4) The bracteoles arranged more or less at right angles to the main branches 
of the inflorescence, giving a general bristly appearance. 

5) The bracteoles strongly recurved. 

6) Variable number of tepals, 3-5, membranous, very unequal, without green 
mid-veins. 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 953 


7) Tepals spathulate to obovate. 

8) Tepals reduced, very small. 

9) Variable number of stames, 3-5. 
10) Fruit spherical, indehiscent, irregularly dehiscent or circumcissile with 
the pericarp smooth, the apex rounded (without a “beak”) and the stigma 
branches very thin and erect. 


The plants may exhibit one or more of the above characteristics, sometimes in 
combination with the usual features of A. hybridus. The variation is continu- 
ous, and at least for the moment, it was impossible to classify it into distinct 
types. As Sauer suggested (1950), collecting more material from these regions is 
needed for a better understanding of this taxon. 

Distribution and ecology.—Originally from Eastern North America, Mexico 
and Central America, but now widespread all over the world as a weed in culti- 
vated or waste places. 


eas specimens exer: ABRICN: elaiia ailimsen: vicinity of Tlimsen, 250 m, 
1932, Faure s.n. (US).ZIMBABWE ity (Harare), 3 Mar 1979 He 
ee (MO). Que Que Distr.: Silsbury, 15 Mar 1978, Chipunga 165 (MO). MALAWI: Ulongue, Zone A, 
17 Oct 1985, Patel & ee 2799 (M 
ASIA. PAKISTAN. Azad Kashmir: Mazaffarabad Distr., 4 Oct 1975, Chaudhri et al 320 (MO). 
Punjab: Rawalpindi ae Islamabad, University of Islamabad, 2200 m, 2 Sep 1975, Chaudhriet al. s.n. 
(CM). Swat: Marghazar, 1300 m, 14 Oct 1975, Shah et al 501 (A). CHINA. Huanan: Cili, 390 m, 9 Sep 
1984, Chow et al 395 (A). JAPAN. Honshu: Pref. Kyoto, Goma, Hiyoshi-cho, Funai-Gun, 200 m, 29 Sep 
1991, Murata & Takahashi 70295 (A). Miyagi: Miyagino-mushi, 3 Oct 1972, Naito 72103 (A, MQ). 
Musashi: Tokyo-to, Hondo, 22 Oct 1960, Furuse s.n. (A). 
AUSTRALIA AND OCEANIA. AUSTRALIA. New South Wales: Woolloomooloo, 33°52'S, 
51°13'E, 6 Apr 1976, Coveny 7788 (A, RSA). Summer Hill, May 1970, Michael M9 (WIS). Queensland: 
Brisbane, Kingaroy, 15 Apr 1947, Smith 3027 (A). 

EUROPE. SWITZERLAND. Basel: Klein Riechen, 10 Sep 1932, Aellen s.n. (WIS).SPAIN. Catalunya: 
Barcelona, Can Tunis, 23 Sep 1913, Sennen 1789 (GH). ITALY. Pro esses tee pemllia, 22 
Sep 1917, nee OONOr ees OM, GH). Prov. Turin: La Molineta, 11 Sep 1852, Perrier dela Ba 
(GH).ROMANIA. Mun. Buct Bucharest, 75 m,26 Sep 1971 see 3334 (CM). ee 
Campus seh pe of Agronomic Sciences, 12 Sep 1994, Costea s.n. (BUAG). 

NORT A. CANADA. Lambton Co.: Squirell Island, 2 Oct 1957, se O. eepele”? ge 

(WIS). caine nits ALABAMA. Dallas Co.: 5 mi E of Selma, 35 m, 23 Sep D 
(SMU, WIS). Lamar Co.: 7 mi N of Millport, 26 Jul 1987, Gasparini 156 (IBE). oe Pinal Co.: San 
Tan Village, near Sacaton, 29 Sep 1925, Peebles et al. 160 (ARIZ). New Jersey. Copper Point, 6 Nov 
ons nes MARZ), Lariat Benton Co.: Butler ne ee Sulphur Springs, 920 m, 2 
Sep MU, WIS). Crittenden Co. mphis, 70 m,16 Sep 1969, Demaree 
61082 ie SMU). iincoln aa a Id, 32 m, Demaree ae Hs Montgomery Co.: top of Cristal 
Mountain, 580 m, 24 Nov 1965, Demaree 53340 (SMU, WIS). Nevada Co.: Prescott, 300 m, 15 Aug 
1970, Demaree 62606 (DS, SMU). Pulaski Co.: Little Rock, 100 m, 20 Oct 1965, D 52919 (OKL, 
SMU). Saline Co.: Benton, 90 m, 19 Sep 1965, Demaree 52912 (MSC, OKL, SMU, WIS). CALIFORNIA. 
Los Angeles Co.: Bryant Ranch, 13 Sep 1932, Wolf 5215 (RSA). Riverside Co.: ranch near Tamecula, 
27 Sep 1940, Stubblefiel ee RSA). San Francisco Co.: San Francisco, Embarcadero at Mont- 
gomery Street, 22 Sep 1957, Howell s.n. (RSA, CAS, DAV).San Francisco, Fort Point, 29 Sep 1974, Norris 
256346 (RSA). Santa Barbara Co.: Santa Barbara, 7 Sep 1957, 2 Oct 1957, Pollard s.n. (CAS, SMU). 


— 


—a 


954 BRIT.ORG/SIDA 19(4) 


Ventura Co.: Ventura River's bed near Casitas Spring, 24 Sep 1965, Pollard s.n. (CAS). Horn Canyon 
Creek near Ojai Avenue, 2 Aug 1960, Pollard s.n. (DAV). CONNECTICUT. Southington, 16 Sep 1904 
Andrews s.n. (BH). FLORIDA. Alachua Co.: Gainesville, E side of SW 17" Street, 20 Jul 1970, Perkins 
663 (FLAS).W of Gainesville, ca. 1/2 miW of I-75 on N side of FL 26,12 Jul 1990, Gilliand 11 (FLAS, NA). 
Broward Co.: 2 mi W of Coral aoe 21 Feb 1970, McCart 11305 (FLAS). Lee Co.: Western Sanibel 
Island, 6 Dec 1972, Brumbach 812 AS). Indian River Co.: W of Wabasso Island, 7Aug 1968, D’Arcy 
2299 (FLAS).GEORGIA. Elbert Co.: ee perton, garden of Stinchcomb Church Road, 23 Aug 1977, 
Dunn 1243 (FLAS). Ogelthorpe Co.: Lexington, 29 Sep 1965, Demaree 53018 (WIS). Illinois. Cham- 
si ign Co.: Urbana, 2 Oct 1947, Jones 17668 (MIN).Union Co.: Panther’s Den, 7 Aug 1959, MacMahon 
5 (MIN). Randolph Co.: Old Kaskaskia, Kaskaskia Island, 21 Oct 1959, Fvers 63076 (MIN). Madison 
Collinsville, 9 Nov 1967, Demaree 57461 (SMU, WIS). Jackson Co.: 2 1/2 mi of Carbondale, 19 Oct 
1985, Handel s.n. (CM). River Forest, 8 Oct 1896, £.T Harper & S.A. Harper s.n. (ISC, WIS). lowa. Johnson 
Co.: Oxford Tw ie 80°N, O8°W, 24 Sep 1975, Kantor 605 (IA). 10 mi N of Johnston 20 Aug 1952, Ander- 
son s.n. (WIS). Linn Co.: Coggon, 15 Oct 1965, McWilliam 15123 (US).Muscatine Co.: near Fruitland, 
25 Sep 1909, oi mk s.n. (IA). Pine Mills, 28 Aug 1892, Reppert s.n. (IA). INDIANA. Jackson Co.: near 
seinen 985, Handel s.n. (CM). Ripley Co.: along Laughery Creek, E of Friendship, 13 Oct 
Deam 55896 (MIN). Steuben Co.: E side of James Lake, 20 Sep 1914, Deam 15484 (MIN).Cloud 
dia, between 5 & 6 Streets, 1 Sep 1940, Fraser 622 (KSC). Salem, 20 Jul 1930, Brooks 1445 
min ee Riley Co.: Manhattan, H.A.Goff Farm, 23 Sep 1969, Goffs.n. (KSC). Saline Co.: Salin 
31 Aug 1930, Hancin 695 (KSC).17 Aug 1931,B1 ake 11380 (MSC).Cambridge, 27 Sep 1896, Blanki a 
5.n. (ISC). MICHIGAN. Lanawee Co.: Hidden Lake Sn of Michigan State University, 15 Sep 1960, 
Freeman 18 (MSC). ee Grenada Co.: 2 mi N of Gore Springs, 12 Aug 1986, Morris 2471 
(IBE). Leflore Co.: 7.5 mi N of Green Wood, 27 nae 959, McDaniel 1276 (IBE). Jones Co.: 1 mi E of 
Laurel, 29 Sep 1978, ne 1279 (IBE). Madison Co.: Natchez Trace Parkwey, 22 Sep 1948, McDougall 
1/77 (US). Monroe Co.: vicinity of Aberdeen, 4 Sep 1994, MacDonald 7599 (IBE). Washington Co.: 
Just N Leland, 28 Aug 1990, Bryson 10271 (IBE). Missouri. Kansas City, 25 Sep 1913, Rose 15133 (US). 
Jackson Co.: 14 Oct 1943, Bush 316 (MIN). Jasper Co.: Webb City, 7 Oct 1949, Palmer 49851 (UMO). 
L nce Co.: Sycamore Hollow, 20 Sep 1950, Palmer 51143 (UMO). Lincoln Co.: N of Troy, 15 Sep 
es aa (WIN). St. Louis Co.: Saint Louis, 20 Sep 1 saline Muehlenbach 17995 (CM); 
51,21 Aug 1949, a. ee MIL, SMU, WIS). Nevada. Clark Co.: 1.5 mi S of Moapa, 450 m, Niles 
3100 (ARIZ). NORTH CAROLINA. Chemung Co.: Elmira, 21 Sep 194, a 2306 (BH). oe Co.: 
Wilton, 2 Sep 1937, a eee (NA). Iredell Co.: 0.8 mi W-NW of Scotts, then 2.6 mi S, 24 Oct 1958, 
Ahles 51921 (FLAS).Orange Co.: Korstian Division, E of Route 1712,400 m, 22 Sep 1988, Palmer 1 ed 
(DUKE). Wake Co.: 0.5 mi NW of Sandy Ridge or the aa of NC 50 and County Road 1 
Wilbur 45364 (DUKE). Wilson Co.: State Road 1649, iS of Lucuma, 22 Oct 1991, Wilbur ere 
(DUKE). NEW YORK. Bronx Co.: New York, near the ne Garden, 21 Oct 1986, Nee 3327( ae 
WIS). Kings Co.: Long Island, 25 Sep 1955, Brooks 3977 (BH). Tompkins Co.: ca. 2 mi S of Ithaca 
Sep 1943, Schuster 8217 (DUKE). Ithaca, 1 Oct 1941, Thorne 273 (IA). Ithaca, 6 Oct 1926, Bailey s.n. “i 
10 Sep 1917, Bailey s.n. (BH) 21 Aug 1916, Metcalf 6395 (BH). Ohio. Portage Co.: Kent, 20 Sep 19 
Hopkins s.n. (CM). Richland Co.: Mansfield, 12 Aug 1895, Wilkinson s.n. (BH). OKLAHOMA. Kay a 
near Tonkawa, 4 Aug 1913, Stevens 1841 (US).Cleveland Co.: Little River bottoms, 25 Sep 1920, Jeffs 
1309 (OKL). Muskogee Co.: s.|,, 28 Aug 1927, Little Jr. 2711, 2487, 2197 (OKL), Ottawa Co.: Ottawa, 29 
Aug 1913, Stevens 2557 (MIN). PENNSYLVANIA. Allegheny Co.: 4km N. of Tarentum, 2 Sep 1990, 
Zand 339 (BH, CM). Bearer Co.: 18.2 mi Wet Raccon St. Park., 9 Oct 1965, Farnsworth s.n. (CM). near 
Georgetown, along Ohio River shore, 1 Sep 1951, Henrici 15112 (CM). Bedford Co.: 1/2 mi S-SE 
Hyndman, 300 m, 2 Sep 1940; Berkheimer 2212 (CM). Berks Co.: 1/2 mi S of Bethel, 110 m, 19 Sep 
1948, Berkheimer s.n. (IA). 2 mi NE of fae Oct 1986, Nee 33266 (CM). Butler Co.: Petersville, 23 
Aug 1922, — $.n. ieee Centre Co.: Oak Hall, Mill Pond, 10 Sep 1939, Wahl 434 (BH). 
Delaware Co.: Upper Darby,0.2 mi SE of intersection between Marshall Road and 69" street Boule- 
vard, 30 Sep 1941, Wheeler pee 48 (RSA). Waele Co.: Penn, South Huntington Township, 


— 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 955 


40°14'N, 79°45'W, 270 m, 12 Aug 1974, Utech 94-1959 (CM). TENNESSEE. Davidson Co.: sf seat 
locality cultivated at Ames, lowa, 1 Oct 1965, McWilliams 1826 (WIS).L Lake Co.: Near Markham, 1 
Aug 1947, Sharp et al.6266 (WIS). Shelby Co.: Presidents Island, nen m,25 Oct 1952, Demure 
33190 (GH). pe Co.: Johnson City, 813 Forest Avenue, 15 Oct 1994, Churchill 94206 (MSC). 
TEXAS. Brazos Co.: Col Hills Woodlands (cultivated), 9 Sep 1942, ee (SMU). Sunset Co.: 
8 mi Tyler, 14 Oct 1944, ee 917 (BH, GH). VIRGINIA. Bedford Co.: 15 Sep 1869, Curtiss s.n. (GH). 
Quantico, 20 Aug 1915, Tidestrom 7582 (NA). Campbell Co.: intersection of Waterlik and Leesville 
Rds., 26 Aug 1978, Ramsey & Freer 26303 (SMU, WIS). Diggs Beach on St. Rte. 643, 8 Oct 1978, van 
Monttfrans 2106 (FLAS).Giles Co.: New Port, 580 m, 14 Aug 1958, IItis 19938 (WIS).Wisconsin. Richland 
1 mi NE of Richland Center, 20 Aug 1983, Nee 43787 (MO). Rock Co.: near Edgerton,5 Sep 1952, 
Sauer 1597 (WIS). WASHINGTON D.C. 10 Sep 1897, Steele s.n. (DUKE); s.1., 11 Aug 1949, Freeman 417 


oS 


). 
MEXICO AND GREATER ANTILLES. BERMUDA. St Island: South Road, 31 Aug 19 
Collins 189 (GH);s.1., 10 Mar 1908, Brown 538 (GH). MEXICO. Chiapas: Mpio. Pokolum, Tenejapa oe 
m, Breedlove 6091 (DS); 5.1.22 Nov 1964, Breedlove 7423 (WIS). Chihuahua: Mpio. Batopilas, Barranca 
de Batopilas, 2100 m, 10 Oct 1973, Bye 5391 (ECON). Nobogame, 28°30'N, 108°30'W, 1800 m, 10 Sep 
1987, Laferriere 545 (WIS). Hidalgo: San Bartolo, Tutotepec, 1000 m, 4 Nov 1973, Gimate 861 (ARIZ, 
UMO, CAS). Distr. soir Ba a campus a UNAM, 2300 m,4 Sep 1986, Nee 32980 (CM, WIS). 


Jalisco: peeere Oct 1896, Palmer H, US). 15 km W of Guadalajara, 20°43'N, 103°24'W, 
1700 m, 26 Sep 1978, een 625 (WIS). ae Colima, 19°39'N, 103°32'W, 1750-1800 m, 24 
Sep 1978, HH. llis et al 559 (WIS). Michoacan: N of Mason Nuevo, 2300 m, 31 Jul 1977, Bennett et al. 


721 (UMO). Distr. Ixtlan, Mpio. Comaltepec, S of sarees 17°33'N, 96°31'W, 2000 m, 8 Dec 1987, 
Lopez Lopez 5 (MO). Nuevo Leon: 35 mi of Monterrey, Hacienda Vista Hermosa, 700 m, 27 Jun 1939, 
White 1592 (ARIZ). Oaxaca: Nayarit, 11 Jul 1948, Sauer 28 (MO). Valley of Etla, Sep 1895, Aloarry 749 
(GH). Puebla: Puebla, Rancho Pasadas, 2194 m,25 Aug 1909, Nicolas 292 (CM).Veracruz: Distr. Papautla, 
Tajin, 3 Dec 1947, Gonzales 105 (BH). Mpio. Xalapa, Claviejo, Xalapa Botanical Garden, 19°30'30"N, 
96°56'30"W, 300 m, 18 Sep 1986, Nee 33066 (WIS, CM). Remundandero, Feb 1923, Purpus 8870 (GH). 
Mpio. Emiliano Zapata, Barranca de San Antonio, 590 m, Hernadez 62 (MO). between Veracruz and 
Villahermosa, km marker 180, 70 m, Croat 62122 (MO).Mpio. Coatepec, 3 km N of Coatepec, 19°29'N, 
96°57'W, 1250 m, 22 Aug 1986, Nee 32852 (WIS).Rancheria Palmas Cuatas, Ignacio de la Llave,6 m, 12 
Aug 1966, Martinez 21739 (DAV). BAHAMAS. Crooked | land: Landrail Landing, 6 Jun 1977, Corell & 
ieee (US). CUBA: Valley of San Angustina, 21 Mar 1903, Britton et al.512 (CM).GUATEMALA. 
Verapaz: s.f.Coban cultivated at Davis, California 29-31 Oct 1948, Sauer 1309 (WIS). Dep. 
foe .2 km N of el Progreso (W side of the Road) to Jalapa, 1040-1060 m, 27, 30,31 Dec 1975, 
lItis G-14 (WIS). Dep. Santa Rosa: Santa Rosa, 930 m, Aug 1892, ie hci est teh oie NICARAGUA. 
Dep. Esteli: Santa Cruz, 13°01N, 86°18'W, 950-1000 m, 9 Nov 1980, Moreno 4561 (MQ). Loma 
Ocotecalzado, 13°10'N,86°18'W, 1260-1300 m, Stevens et al 15589 Mon COST: ee Prov. Cartago: 
Cartago, 1400 m, Nov 1887, Cooper 5908 (GH, US). Prov. San Jose: San Jose, 1100 m,Jan 1896, Tanduz 
9856 (GH). PANAMA. Chiriqui: vicinity of Boquete, 1000-1250 m, 12-13 Dec 1966, Lewis et al 594 
(GH, NA, UC). Bambito, 1400 m, 12 Mar 1974, Tayson 7240 (US 
SOUTH AMERICA. COLOMBIA. Cauca: Cordillera Central, River Palo, between Tacueyo and La 
Tolda, 1780-1900 m, 14 Dec 1944, Cuatrecasas 19472 (GH). BRASIL: Bello Horizonte, 1000 m, 5 Mar 
1945, William & Assis 6461 (GH). Goias: Corumba de Goias, 16°S, 49°W, 1000 m, 3 Dec 1965, Irwin et al 
10991 (RB). Parana: Parque Nacional de Iguacu, Picado do Benjamin, 400 m, 26 May 1949, Duarte & 
Pereira 1911 (RB). Rio Grande do Sul: Minas, 16 Apr 1935, Hoffman 6 (RB). BOLIVIA. Yungas. 1890, 
Bang 231 (GH). Hacienda Simaco above the road to Tipuani, 1400 m, Jan 1920, Buchtien 5401 (GH). 


> 


BASIONYA 


3b. Amaranthus hybrids ee quitensis eu Costea & Carretero, comb. 
OV. b,, Bonpl. & Kunth, Nov. Gen. Sp. 2., folio: 
156; ed. 4:194. 1817. A. retrofle exus L. subsp. ee (Kunth) O. Bolos & Vigo, Butll. Inst. 


956 BRIT.ORG/SIDA 19(4) 


Catalana Hist. 38:89. 1974. A. hybridus L. var. quitensis (Kunth) Covas, Darwiniana 5:329-368. 
1941. Type: ECUADOR: “Crescit in ripa Muvii oe bae, alt 1030 hex,” (Regno Quitensi), 
6, 1802, Humboldt & Bonpl and 3082 (HOLOTYPE: 


v 


Amaranthus caudatus sensu Greuter et al., Med-Checkl. 1:46. 1984. p.p., non L.s. str. 


Cotyledons lanceolate, 12 x 2-2.5 mm, with acute apex. Pollen grains 25-30 um 
with 28-35 pores of 1.2-1.4 1m in diameter. The pollen differs to subsp hybridus 
by the tectum with high density of spinules. (Fig. 2, F). 2n= 32 

We share a different view from Coons (1975, 1978) over the boundaries of 
this taxon. The tepals can be variable, erect to recurved as in other species (for 
example Amaranthus retroflexus). More important are the length and shape of 
tepals. Therefore the type of this taxon is indeed Humboldt & Bonpland 3082 
deposited in Paris (P). 

The length of bracteoles is variable in subsp. quitensis too. Typically the 
bracteoles are 15-2 times longer than the tepals. The plants from Galapagos 
Islands have shorter bracts, 1-1.2 times longer than the tepals and also stand 
apart in that their {lowers have tepals and bracteoles with very large, ramified, 
green mid-veins with very narrow membranous margins. In the present account 
we do not include A. hybridus var. sangorache (Coons 1975, 1978) within A. 
hybridus subsp. quitensis more research being necessary in order to clarify the 
status of this taxon. 

Distribution and ecology.—A native riverbank pioneer in tropical South 
America, where it is a noxious weed, this plant is more dependent on a warm 
climate than subsp. hybridus. Even though it was recorded in Europe since the 
19th century, as a casual in most countries, it is naturalised only in the Azores 
and Balearic Islands (Akeroyd 1993). It has also been collected in Australia. 

|: SOUTH AMERICA. ECUADOR. Charles Island: Black Beach 

19 ie 1932, ell 9381 (CAS). Isabela Island: Tagus Cove, Jun 1899, Snodgrass & Heller 219 (GH). 
Santa Cruz: along “Old Trail" from Academy Bay toward Bella Vista, 5-100 m, 24 Jan 1964, Wiggins 
ise (CAS). Bella Vista, 200 m, 27 Jan 1967, Fosberg as (RSA). Academy sie Apr 1953, Bowman 
Bates ae | Chato, 31 Jul 1966, Colinvaux s.n. (CAS). PERU. Dep. Caj Valley of Rio Chamaya, 

W of Pucara, 5°55'S, 79°19'W, 1100 m, 18 Apr 19 Croat 58375 (CAS). oe Cusco: Prov. 
fai 1/2 mi E of Urubamba, 2800 m, /Itis & Ugent 1160 (CAS). Dep. Piura: 37,3 km of Pucara, 
5°47'S, 79°27W, 1550 m, Croat 58392 (CAS). BRASIL. Estado do Rio Grande do Sul: Serra Geral, 17 
Mar 1983, Silveira & Frosi 544 (RB). BOLIVIA, Andres Ibanez: 12km E of center of Santa Cruz, 17°46' 
to 47',63°04'W, 375 m, 4 Feb 1987, Nee 33988 (CM).W side of Santa Cruz, 17°47'S, 63°40'W, 420 m, 14 
Jan 1987, Nee 35481 (CM). 15 km of Cotoca, 17°42'S, 62°53'W, 325 m, 28 Jan 1989, Nee 37776 (WIS), 
Santa Cruz: Prov. Caballero: 2km NW of Rio San Isidro bridge in San Isidro, along to highway to 
Comarapa, 18°02'S, 64°27'W, 1575 m, 29 Dec 1995, Nee 46591 (ADA). Angostura, 550 m, 25 Jun 1966 
Stel ba s.n. (RSA). ca. 21 km SE of Palmar del Oratorio, 18°02'S, 63°01'W, 365 m, 22 Jan 1989, Nee 
37648 (WIS). Ichilo oo Vista, 17°27'S, 63°40'W, 370 m, 2 Aug 1987, Nee 33510 (CM). PARAGUAY. 
ha Tororo, Camino a Polilla, 25°55'S, 55°15'W, 10 Dec 1988, Degen 1035 (MO). Centro Forestier, 
Pta. Stroessner — Alta Parana 250 m, 24 May 1982, Stuts 370 (MO). URUGUAY. Dep. Montevideo 
Pocitos, Mar 1924, Herter 68154 (MSC, WIS). ARGENTINA. Prov. Buenos Aires: La Belgica, 17°34'S, 
63°13'W, Nee 33475 (ISC). Hudson, 15 May 1945, Hunzinker 661 (A);s.1., 15 May 1945, Hunzinker 2260 


te 


— 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 957 


GH). Pergamino, 22 Mar 1929, Parodi 8910 (GH). Prov. Catamarca: Ancasti, Rio Chico, 28 Nov 1950, 
Brizuela 759 (CM). Dep. La Paz, El Bello, 3 Mar 1950, Brizuela 1211 (MICH, CAS). Prov. Chaco: Dep San 
Martin, La Leonor, 26-27 May 1988, Schinini 26222 (GH). Prov. Formosa: Dep. Pilcomayo, Ruta 86 al 
km 55,9 Feb 1948, Morel 4594, 6542 (UMO). 3 km of Portenito, 6 Oct 1947, Morel 3857 (US). S of 
Laguna Primavera, 20 Dec 1949, Morel 9110 (CM). Prov. Misiones: Dep. San Pedro, Cataraguatay 
(Centro) 11 May 1949, Montes 1625 (CAS, WIS). Pasadas, 21 Jul 1945, Bertoni 1519 (A);5.1,,17 Nov 1907, 
Ekman 117,118 (US, GH). Isla Pindoi, Jul 1945, Grovetto 3435 (CAS). a ene El Potrero, 30 Mar 
1950, Brizuela, 1099 (US). Dep. Boqueron, 68 km NE de Filadelfia, 10 Dec kas et Cristobal 
44281 (MO). Dep. Candelaria, El Datil, 7 Feb 1949, Montenegro 349 oe ae Rosario de la Frontera, 
La Junta, 690 m, 29 Jan 1935, Carbone 12700 (GH). Prov. Tucuman: La Ramada, 450 m, 2 Apr 1933, 
Deirano 9666 (GH).Dep Rio Chico, Escaba,600 m,5 Dec 1913, Onetti 1653 (GH).Dep Seales, la Florida, 
270 m, Jun 1913, Monetti 11659 (GH). CHILE: s.f. Angol, grown at Cornell University, Jul-Aug 1937, 
Murray 36023 (10) (BH). 


= 


4. Amaranthus hypochondriacus L., Sp. Pl. 991. 1753. (Fig. 5). Type: “Habitat in Virginia’, 
LINN 1117/24. 


Amaranthus chlorostachys Willd. var. erythrostachys (Mogq.) Aellen, Hegi, Illustr. Fl. Mittel.-eur. 
ed. 2, 3(1-2):482. 1959. A. hybridus L. subsp. hybridus var. erythrostachys Mogq., DC., Prodr. 
13(2):259. 1849. A. Ny Detdats E var. hypochondriacus (L.) Robinson, Rhodora 10:32. 1908. A. 
Cie subsp. I (L.) Thell. proles erythrostachys (Moq.) ThelL., Ascherson 
& er, Syn. Mitteleur Fl. 5:241. 1914. 

ree ean L. Syst. Nat. ed. 10. 2:1269. 1759. 

Amaranthus frumentaceus Buch. Hamilt. ex Roxb,, FI. Ind. 3:613. 1832. 

Amaranthus anardana Buch. Hamilt. in Wall. ex Moq.-Tand., DC. Prodr. 13(2):256. 1849. 

Amaranthus leucocarpus S. Wats., Proc. Amer. Acad. Arts. 10:347. 1 

Amaranthus leucospermus S.Wats., Proc. Amer. Acad. Arts. 22:446. 1887. 


Like A. cruentus, but inflorescence usually stiff, with thick branches. Cotyle- 
dons as in A. cruentus but larger, 18 x 5 mm. Bracteoles 3-5 mm long, about 1.5 
mm longer (sometimes 2 times longer) than the tepals. Tepals 5, one equal to or 
longer than the fruit, the others 4 shorter, lanceolate, with the mid-vein brown- 
reddish. Fruit circumscissile, tapering gradually toward the stigma region, 1.5- 
2 mm long, about 2 times longer than wide, with the dehiscence line in the 
upper half. Stigma branches thick, spreading, about 1.6-18 mm long and 0.6- 
0.8 mm wide at the base. Type of pericarp surface variable: A to B. All the other 
morphological characteristics of the fruits and dark seeds correspond to those 
of A. powellii. The pale seeds are very much like those described for A.caudatus, 
differing in having the cells of the exotesta inconspicuous (because their anti- 
clinal walls are inconspicuous) and the periclinal walls evidently wrinkled. 
Pollen grains of 18-23 um, with 32-45 pores of 13-15 ym in diameter. Density 
of granules medium. 2n = 32, 34. 

Distribution and ecology.—This species is originally from North America, 
where its closest wild relative A. powellii is common, though at the same time it 
is evidently related to the cultivated A. cruentus. Sauer (1993) suggests that it is 
probable that it is of hybrid origin from those two taxa. This relationship is also 
supported by some molecular studies (Transue et al. 1994; Kirkpatrik 1995; Chan 


BRIT.ORG/SIDA 19(4) 


Herbarium - University of California Riverside 


Det. by Mihai Costea , 
CULTIVATED PLANTS 
5 


Amaranthus 


Regional Plant Introduction Station 
Iowa State University, Ames, Lowa 5001 
AMES 2155 81A0S.1 Type 1 
Dansago, Nepal 
Rodale een 

Lehmann 9/19/85 


coll. 


RETUM, WASHINGTON, D.C 


wy) 4 hus hybrid hypochondriacus (L.) Thellung 
TES NATIONAL ARB 
ASHINGTON, D, C Det.: Richard Splut | 
| ETHEL 
aN wi i ae ale USDA, Agncultural Research Service, Plant Serence Institute, Systematic Born 
Laboratory 


Fic. 5.A thus | hondri from Lehmann, AMES 2155 (NA) 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 959 


& Sun 1997). Like A. cruentus, it is cultivated as an ornamental, for grain and as 
a potherb. 


air ies ea examined: AFRICA. ie Nairobi Distr.: Nairobi, Ministry of Works 
ate, W Bahati, 1678 m, Ie Fe b 1969, Mwangangi 1331 (MO). 
AKISTAN B Sk 1U,2150m He 1936, Koelz 9627, 2928 (NA). Swat Distr.: 

Fatehpur, 1270 m, 17 Oct 1975, Shah et al 690 (A). INDIA: s.f. unknown locality cultivated at Kutztown, 
Pennsylvania, 12 Sep 1984, Strudwick & Reider RRC 1175 (MO). Sholur, 15 Mar 1963, Noble # M (NA 
WIS). Kasalhado, 14 Feb 1963, Noble #/ (NA, WIS). E Jakatala, Badaga, 1780 m, 14 Feb 1963, Noble #k 
(WIS).Northern Thuneri, 1900 m, 27 Jul 1993, Noble s.n.(WIS);s.1., 30 Aug 1964, Sauer 3952, 3942c (NA, 
WIS). Jammu and Kashmir: Srinagar, 1400 m, 20 Sep 1956, Polunin 56/819 (MO, NA, WIS). Tangmarg, 
1900 m, 16 Aug 1956, Polunin 56/300 (MO, WIS). Sind Valley, Rezan, 2400 m, 31 Aug 1956, Polunin 56/ 
617 (MO,WIS, NA). near Miragund, 1400 m, 15 Sep 1959, Polunin 56/804 (WIS). Upper Nilgiri Hills, Kil- 

Kotagari, 1800 m, 18 Sep 1962, Noble # e (WIS). Wangat Nullah at Khanan, 2000 m, Polunin 56/759 
(MO, NA). Tamil Nadu: near Coonoor, 21 Jan 1963, Noble # b (NA, WIS). Madras, Kurumba Villages, 
1370 m, 18 ee 1962, 16 Dec 1963, 20 Jan 1963, Noble s.n. (WIS). s.f. Kurumba, 6 Oct 1964, Sauer 
3949b (MO, WIS, ene |, 2 Nov 1964, Sauer 3950, 3954b (WIS, NA, MO). Uttar Pradesh: s.f. Mussorie, 
a by Lal (1961), cultivated in ee York, Ithaca, Aug—Nov 1969, Sauer 2845 (BH, NA, WIS). 
NE s.f. Marku Valley, Sirsagarhi, 10 Nov 1950, cultivated, Sauer 1495D (A).SRILANKA. Distr. North 
cua Province: Anurahdapura, = 61 mi from Kandy on the road to Jaffna South Kagama, 24 
Feb 1973, Townsend 73/117 (US). Central Province: Matale Distr., Sigiriya Wewa, 11 Mar 1973, 
Townsend 73/205 (US). CHINA. Prov. Heilongjiang: Ping Shan,45°57'N, 127°23'E, 370 m, 2 Sep 1993, 
NACPEC, HLJ-37 (NA). Border between Prov. Hopeh and Honan, Chicungshan, 400-900 m, 18 Jun 
917, Bailey s.n. (BH). s.f. Muping Sikong, 2500 m, 10 Nov 1950, Sauer 1489-E (GH). Szechwan: s/f 
Mowhsien, 2000 m, 10 Nov 1950, Sauer 1499-F, 1484-D (GH). Yunnan: Muli, Wachin, Schawan, 3000 
m, ‘side of field,” 1937, Yu 14481, 14482 (A). Ho- “pei: Nesrcpeu Cheng, 26 Sep 1948, Beach 238 (US). 


— 


EUROPE. NORWAY: O39: Oslo s.f. Bc ical 1, University I Norway iS Sep 1940, Gillett 40- 

5-2 (NA). SWEDEN. S kholm oe i eich of Stockholm, Grant 225 (WIS). ITALY. 
Lerten Giulia: s.f. Udine, B IGarden of Instituto Tecnico Zanon, Grant 426 (NA). FRANCE. 
oubs: Besancon, s.f. Botanical Garden of the U sity de la Ville, Grant 428 (NA). Loire- oe 


i Botanical Garden Nates, unknown date, Grant 223, 227, 2276 (NA). ee (WIS). HOLLAND. Noord- 
Holland: Amsterdam, s.f. Botanical Garden of Amsterdam, Grant 137 (WIS). Laine Nordrhein 
Westfalen: Bonn, s.f. Botanical Garden, University Bonn, Grant 428 (NA). AUSTRIA. Burgenland: 
Pamhagen, Aug 1954, Patzak s.n. set RUSSIA. s.f. unknown locality grown in Ithaca, New York, 31 
Jul 1936, Muenschers.n. (BH). ROMANIA. Cluj Co.: s.f. Cluj, cultivated at Ithaca, New York, 19 Jul 1970, 
Dress 9943 (BH).Mehedinti Co.: ee 966, Morariu s.n (BUCA). LITHUANIA. Ne 

s.f. Kaunas, cultivated in Ottawa, Canada, 1939, Zinck 39-97-7 (BH). IRAN: s.f. unknown locality a 
vated — aa Dale, een a fe 939, Cowgill 2355 (BH). 

H AMERICA. CA A. ONTARIO: Ottawa, cultivated at Dominion Botanical Garden, 

29 his 939, Lawrence 682 elie Guns is CALIFORNIA. San Bernardino Co.: San cae 
dino Valley, 300 m, 16 Sep 1907, Parish 6472 (MIN). Yolo Co.: Davis, on the campus, 10 Aug 1 
Kellner s.n. (DAV). FLORIDA. Citrus Co.: Inverness, 3385 South Highlands, 20 Jul 1 ee sn. 
(FLAS). KANSAS. Washington Co.: SE corner of the County, 13 Oct 1938, Holman s.n. (KSC). MICHI- 
GAN. Gratiot Co.: Alma, Sept 1935, Davis s.n. (BH). MINNESOTA. Dakota Co.: Inner Grove, ruderal, 
14 Sep 1930, Jones 513 (MIN). MISSOURI. Jakson Co.: Independence, 2 Sep 1895, Mackenzie 21111 
(MIN, ISC). NEBRASKA. Adams Co.: Hastings, 20 Oct 1934, Rose de Lima 300 (CM). NEW YORK. 
Onondaga Co.: Syracuse, SE corner of Onondaga Lake, 21 Aug 1916, Wiegand 6396 (MO, GH). 
Tompkins Co.: Ithaca, “weed in garden,” 14 Aug 1919, Bailey s.n. (BH); s.l., ruderal, 24 Aug 1925 
Muenscher s.n. (BH).Cinders, College of Agriculture, ruderal, 12 Sep 1923, Burnham 15065 (BH).PENN- 
SYLVANIA. Dane Co.: Murphy's Creek, 22 Aug 1945, Hale s.n. (WIS). Erie Co.: 10 Aug 1894, Miller s.n. 


960 BRIT.ORG/SIDA 19(4) 


(CM). Stoyestown, 20 Jul 1873, Patterson 6210 (CM). WISCONSIN. Rock Co.: Beloit, 9 Aug 1936, An- 
ert n. (WIS). Oneida Co.: Minoqua (ruderal), 24 Sep 1981, Harrington 36 (WIS). 
XICO. Chihuahua: Rio Mayo, eae 20 Sep 1935, Scott Gentry 1844 (NA).s.f. Rancho 
Tigre, 1 a Oct 1948 Sauer 1193, 1396 (BH, NA, WIS). Colima: Colima, Au a t 1897, Palmer 120 (DS). 
Distr. Federal: s.f. Mpio. Atlapulco, San Gregorio, 28 May 1948, Sauer 1177 (MO, GH, NA, WIS); s.,, 31 
Oct 1948, Sauer 1211 (GH, MO, NA, WIS); s.., 31 Jan 1949, Sauer 1404 oonene ees sf. 
Chilapa, 20 Aug 1948 — 21 Jan 1949, Sauer 1162—E£ (MO, GH, NA, WIS). Pemex N of Chilpancingo, 
“ruderal,” 20 Aug 1972, Dunn et al. 20477 (UMO). Jalisco: Tlajomulco, 25 Aug 1947, Sauer 1174 
GH, MO, NA, WIS). Mexico: Temascaltepec de Gonzales, Timbres, 1660 m, 15 Oct 1932, Hinton 2137 
(MO). Mpio. Michoacan: Zacapan, 25 Aug 1947, Sauer 1170 (BH, NA). Zacapu, 25 Aug 1947, Sauer 1395 
(DS, MO, NA, WIS). Opopeo, 30 Oct 1948, Sauer 1274 (BH). Oaxaca: Clavillones in Zimatlan area, 30 
Oct 1948, Sauer 1266 (MO, NA,WIS). Puebla: San Jeronimo Teocuizmalco, 31 Oct 1948, Sauer 1239 (NA, 
WIS). Sonora: Rio Mayo, cultivated, 20 Sep 1935, Scott Gentry 1843 (GH).Rio Mayo, Rancho el Moschite, 
1 Oct 1979, Nabhan GN 975A (GA). Tlaxaca: San Berabe Amaxac, 5 Oct 1949, Sauer 1535 (GH, MO, 
UC, NA, WIS). San Miquel del Milagro, 30 Oct 1949, Sauer, 1288, 1293, 1277, 1139 (GH, MO, NA, WIS). 
Veracruz: between Tequila and Orizaba, 15°45'N,97°5'W, 1600 m, Jan 1984, B. Benz & K. Benz 980 (WIS 


ed 


ies) 
ac 


a 


5. Amaranthus powellii S.Wats., Proc. Amer. Acad. Arts. 10:347. 1875. Typr: De- 
scribed from a plant cultivated at Harvard University “from seeds brought from Arizona by 
Col Powell,” Powell s.n. (HOLOTYPE: US 16163; ISOTYPE: MO) 
Erect annual 0.5-1.5(-2) m, with stem glabrous to puberulent mostly in the in- 
florescence region, with trichomes multicellular and uniseriate. Cotyledons 
narrow-elliptic, 10-12 x 3 mm, with acute to rounded apex and acute to cu- 
neate base; petiole about 5-6 mm long. Leaves, broadly-elliptic to rhombic or 
lanceolate, 3-8 x 2-6 cm. Bracteoles rigid, heavy, 4.5-6(-8) mm long, about 2- 
3(-4) times longer than the tepals. Tepals 3-5, evidently unequal; one tepal is 
2.2-3 mm long, the others 1.2-1.6 mm long, linear-lanceolate to elliptic, with 
the mid-veins inconspicuous (only in the longest tepal may be partially vis- 
ible). The longer (outer) tepal surpassing the fruit, the shorter (inner) tepals 
shorter than (or as long as) the fruit. Fruit dehiscent or indehiscent, elliptical to 
obovoidal, 1.5-2 times longer than wide, when dehiscent with the dehiscence 
line in the upper half, gradually narrowed toward the stigma region, the apex 
of the fruit truncated. Stigma branches thick, spreading from the base, 1.9-2.2 
mm long and 0.08-0.1 mm thick at the base. Seeds lenticular, differentiated into 
central zone and marginal zones. Colour black to dark brown, uniform. Pollen 
grains 18-23 um with 30-45 pores of 1.1-1.3 um in diameter. Density of gran- 
ules low to medium. 

Sauer (1967) and afterward other authors Jalas & Suominen 1980; Carretero 
1985, 1990; Akeroyd 1993 etc.) considered the “European” taxon A. bouchonii, to 
be conspecific with A powellii on the basis of their similarity. The study of 
isozymes (Wilkin 1992) failed to establish satisfactorily the distinctness of these 
two taxa, but the author maintained A. bouchoniiat the specific level. The same 
option was followed by Cacciato (1982), Hiigin (1986, 1987), Kerguélen (1993), 
Stace (1991, 1997). 

Studying European plants we found the indehiscent character of the fruit 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 961 


to be constant. For example, European plants of A. bouchonii cultivated for eight 
years in the vicinity of A. powellii, A. hybridusand A. retroflexus, in the Botani- 
cal Garden of the University of Agronomical Sciences Bucharest, maintained 
their character. Greizerstein and Poggio (1992); Greizerstein et al. 1997) found 
differencies in chromosome number, chromosomal asymmetry and total DNA 
that supports A. bouchonii as an independent taxon. There is also a tendency 
toward ecological separation of the two variants in Europe: A. bouchonti popu- 
lations occur primarily along riverbanks as pioneers, while A. powellii is a rud- 
eral or segetal weed. 

Based on our observations, in North America, plants with indehiscent fruits 
are more variable than in Europe. Some of them could be easily associated with 
A. powellii but others with A. hybridus. In both situations the same plant may 
bear only indehiscent fruit (especially A. powellii) or a mixture of circumscis- 
sile, iregularly dehiscent and indehiscent fruits (especially A. hybridus-like 
plants).S the dehiscence line is partially visible but the fruit does not 
open. The phenomenon occurs naturally. It is possible that the process of evo- 
lution of this taxon is taking place simultaneously in America and Europe, and 
that in Europe the varieties with indehiscent fruits have acquired more stabil- 
ity and consistency. It is also probable that the European A. bouchonii forms 
have reached America. 

After close morphological and anatomical examination we found some new 
characters that, together with the previous data, suggest the subspecies rank 
for the plants with indehiscent fruit. 


1. Inflorescence stiff and erect, + unbranched or with a few widely spaced, long 

branches. Fruit circumcissile, 2 times longer than wide, irregularly wrinkled above 

the dehiscence line, with the pericarp surface type A. Seeds with an indistinct oa 

row extending 1/3 to midway through the seeds, with the marginal zone alm 

mooth A. powellii subsp. powellii aa 1A;2 A,C;6) 

1. Inflorescence often not strictly erect, more lax, with many lateral branches. Fruit 

indehiscent or irregularly dehiscent, 1.5 times longer than wide + smooth with the 

type of pericarp surface variable: B to C. Seeds with a concavity above the hilum 
and the radicle, with the marginal zone evidently sculptured A. powellii subsp. 
bouchonii (Figs. 1 E, F; 2 B; 7) 


5a. Amaranthus powellii subsp. powellii (Fig. 6). 

Amaranthus chlorost ie Willd. var. ae ee ThellL., Vierteljahrssch. Naturf. Ges. 
Zurich 52:443. 1907. Amaranthus hybridus L. subsp. hypochondriacus (L.) Thell. var. 
a. ‘1 Thell. f. pseudo- einai til eee ee ner, Syn. Mitteleur. 

914. Amaranthus ee s Willd. var. (subsp.) powellii (S. Wats.) Priszter, An 


t. Vit. Budapest, 2 (2):144. 1953. see oe idus L. subsp. hybridus var. es 
ait es Thal eee oe Bot. 11(4):12 
Amaranthus chlo ensu Hayek, Prodr. Fl. en a 1:160. 1927.; Davis, Fl. Turkey 


2:341. 967.; Cacciato, FI. av 1:179. 1982 
Amaranthus hybridus” sensu Aellen in Tutin & al. (eds.), Fl. Eur, ed 1, 1:109 (1964); FL. Palaest. 


962 BRIT.ORG/SIDA 19(4) 


Fic. 6.A h lif sut powellii, f thet (P lis.n., US 16163). 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 963 


Amaranthus hybridus subsp. hybridus sensu Townsend, p.p., non L.s. str, Fl. Zambesiaca 9(1):49. 


: thus | hondri et al., Med-Checklist 1:47. 1984. p.p., non L.s. str. 

Amaranthus hypochowdriacus var. chlorostachys sensu Morariu, Fl. R-PR. 1:586. 1952 
Inflorescence stiff, erect, + unbranched or with a few widely sae long 
branches; the terminal branches are usually much longer than the lateral ones. 
Bracteoles usually 2-3 times longer than the tepals. Fruit 2 times longer than 
wide, coarsely wrinkled above the dehiscence line. Pericarp surface type A (Fig. 
1 A). Seeds elliptic to obovate, 1.25-13 x L- 1.1 mm, the marginal zone almost 
smooth. Cells of the exotesta with i ticlinal and almost smooth, 
plane periclinal walls (Fig. 2, A, C); border of the seed acute. 2n = 22,08 

Distribution end ecololgy.—Worldwide weed, native to North and South 

America. In Europe it is a very frequent ruderal and segetal weed frequently 
mistakenly called “A chlorostachys,” “A-hybridus” or “A hypochondriacus.” 


Representative specimens examined: AFRICA. ETHIOPIA. Shewa: Addis Ababa, near University 
College, 2300 m, 13 Dec 1965, Wilde Duyfies 9235 (MO). Alemaya, College of Agriculture, 2050 m 
Apr 1968, Westphal & Westphal Stevels s.n. (MO). UGANDA: Muzingura-Kashaarara, 00°20'S, 30°26 E, 
1900 m, 16 Dec 1987, Rwaburindore 2550 (MO). TANZANIA. Iringa: Mufindi, 8°31'S, 35°10'E, 30 May 

1989, oe 642 (MO). 

ASIA. PAKISTAN. Kurram Valley Distr.: Makai to Parachinar, 1700 m, 15 Oct 1975, Dar &al. 7 
(A). ey aan and Kashmir: Upper Nilgiri Hills, 1930 m, 23 Jun 1963, Noble s.n. (WIS) 

AUSTRALIA AND OCEANIA. AUSTRALIA. New South Wales, 15 Mar 1970, Michael Mé (WIS). 
NEW ZEALAND. Waitemata Co.: Marangi Bay, 36°44'S, 174°45'E, 30 Mar 1974, Bangerter 5157 (NA). 

NORTH AMERICA. CANADA. Lambton Co.: Forest, 2 Oct 1963, Geiser 3374 F (WIS). UNITED 
STATES. ARIZONA. Cochise Co.: Chiricahua Mountains, W side of mountain, on road to ane 
National Monument, 2150 m, Oct 1943, Kearney & Peebles 15119 (CAS). Coconino Co.: Home- 
stead, Flagstaff, 2100 m, 15 Aug 1935, Whiting 756/1471 (ARIZ). CALIFORNIA. Alpine Co.: ae 
1800 m, 10 Aug 1964, Howell 409170 (CAS). Inyo Co.: White Mountains, Antelope Springs, 1800 m, 2 
Aug 1985, Morefield 3110 (ARIZ). Lassen Co.: Sierra Nevada, Susanville, 1400 m, 8, 9 Jul 1974, Howell 
50500 (CAS). Monterey Co.: King City, 5 Aug 1963, Howell 39616 (CAS). 7 Aug 1963, Howell 40067 
(CAS). San Joaquin Co.: 5 mi W of Stockton, 14 Aug 1953, Sauer 1642 (WIS).Santa Barbara Co.: San 
Antonio Road near Hollister Ave., 19 Aug 1957, Pollard s.n. (CAS). Santa Clara Co.: San Francisco 
Creek, 29 Jul 1973, Wiggins 22055 (DS). Sonoma Co.: near Guerneville, 3 Sep 1951, Rubtzoff 734 (CAS 
Ventura Co.: Ojai Valley, Lama Drive, 8 Nov. 1967, Pollard s.n. (CAS, MIN). Ojai, Ojai Avenue, between 
Gorham Road and San Antonio Creek crossing, 16 Oct 1966, Pollard s.n. (CAS). along Front Street, 5 
Oct 1960, Pollard s.n. (CAS).Mirror Lake, 25 Jul 1959, Pollard s.n. (CAS).Southern Pacific Road, near San 
Juan Barranca crossing, 20 Sep 1960, Pollard s.n. (CAS).Ortonville, 19 Oct 1962, Pollard s.n. (CAS). Yolo 
Co.: Merrit Island, near junction 140 and 142, 3.5 mi S Clarksburg, 18 Aug 1969, Quick 69-20 (CAS). 
McKinley Co.: Zuni reservation, 13 Aug 1978, Nabhan 1037 (ARIZ). COLORADO. Adams Co.: Ben- 
nett, Kiowa Creek at route 36 bridge, 2 Sep 1974, Churchill s.n. (MSC). Mineral Co.: Don Juan Moun- 
tains, 5 mi bellow the summit Wolf Creek Pass, 2600 m, 28 Jul 1928, Wolf 3074 (CAS, DS, BH). IDAHO. 
Blaine Co.: Hailey, 23 Aug 1909, Woods & Tidestrom 2814 (US). Spencer, 26 Jun 1916, Rust 420 (CAS). 
Nez Perces Co.: Forest, 17 Jul 1896, 1100 m,A.A. Heller & E. G. Heller 3428 (US). IOWA. Palo Alto Co.: 
Lost Island Lake, 17 Jul 1934, Ada Hayden 4006 (lA). MASSACHUSETTS. Hampshire Co.: Haydenville, 
3 Sep 1975, Ahles 81290 (UMO, CM, IA). Bristol Co.: Dighton, 5 Oct 1956, Seymour 17042 (WIS).Worces- 
ter Co.: Lancaster, 12 Sep 1943, Seymour 6006 (WIS). Leicester, 7 Aug 1944, Gates s.n. (WIS). MICHI- 
GAN. Grawford Co.: 1 mi N of Seven Mile Road and 4 mi W of Beaver Creek Road, 14 Sep. 1992, 


— 


a 


964 BRIT.ORG/SIDA 19(4) 


Chittenden & Peil 1108 (MSC). Allegon Co.: Holland, Hope College, 16 Sep 1976, Gillis 13485 (MSC). 
Shiawassee Co.: 12458 Warner Rd., Laingsburg, 17 Aug 1984, Harwood 285445 (MSC).Mecosta Co.: 
along Michigan Northern Railroad track, 50 m W of center of village Stanwood, 29 Aug 1984, Rabeler 
832 (DUKE, MSC).MINNESOTA. Freeborn Co.: Albert Lea, 16 Sep 1979, ia cae (MIN).Heunepin 
Co.: Minneapolis, N end of University of Minnesota campus, 17 Sep 1970, Ownbey 4398 (MIN). Mower 
Co.: Brownsdale, 26 Aug 1981, Smith 5576 (MIN). Washington Co.: along the street Criox River, 26 
Sep 1988, Smith 15117 (MIN).MISSOURI. Jasper Co.: near Joplin, 17 Aug 1920, Ss et Mel 
" re: 7 Apr. 1958, Muehlenbach 1435 (UMO); s.,, 26 Jun 1971, Muechlenbach 3587 (MO, NA). N 
A. S of Carlin, 26 Aug 1984, Williams & Tiehm s.n. (CAS). Lander Co.: Trout ae 16 Aug 38 
Be Henning 67842 (DUKE). Washoe Co.: N of Sparks, near Wild Creek Golf Course, 1100 m, 
Sep 1984, Williams s.n. (CAS). NEW MEXICO. Dona Ana Co.: Organ Mountains, 1400 m, 23 Sep Ha 
Wooton & Standley s.n. (US). Grant Co.: Fort Bayard, 28 Aug 1905, Blumer 42 (GH). Valencia Co.: £1 
Morro National Monument, 4 Aug 1939, Vogt 20 (ARIZ). NEW YORK. Chemung Co.: Elmira, 20 Sep 
1941, Smith 2303, 2304 (BH).Madison Co.: Oneida Lake,6 Sep 1939, Allen 20005 (BH). Tompkins Co.: 
South Hill, Ithaca, 23 Aug 1933, Allen 18316 (BH). OREGON. Benton Co.: (without location) 1908, 
Gilkey s.n. (US). Polk Co.: along Hwy. 99 W,N of Hoffman Road, 16 Jul 1982, Hasse 2513 (ARIZ). PENN- 
SYLVANIA. Bedford Co.: Coleraine township school, Route #326, 3 Sep 1972, Duppstadt s.n. (MIN), 
near Cessna, 30 Aug 1941, Berkheimer 2934-B (CM). Berks Co.: 2 mi NE of Kutztown, 6 Oct 1986, Nee 
33265 (CM).1/2 mi NE of Trexler, 18 Aug 1953, Schaeffer 44917 (US). Lawrence Co.: 0.25 mi SE of New 
Bedford along Marr Road.8 Sep 1990, Bonnie & Isaac 354389 (C _ . ISLAND. New Port Co.: 
Block Island, Old Harbor, 19 Aug 1913, Fernald & Long 9463 (GH). TEXAS. Brewster Co.: Chisos Mts.,4 
Aug 1931, Mueller 7986 (US). UTAH. Iron Co.: Cedar City, 1900 m, ve Sep 1968, Howell & True 45349 
(CAS). Salt Lake Co.: Salt Lake City, 28 Aug 1939, WC. Muenscner ori, V.Muenscher, 15913 (BH). WIS- 
CONSIN. Columbia Co.: 1 mi W at Hwy 51, 22 and 60 intersection, 22 Sep 1964, Weis 30 (WIS). 1 mi 
SW of Columbus, 10 Sep 1956, I/tis & Koeppen 8174 (WIS).Dan Madison, Oct 1949, sat sn. 
(WIS). Green Co.: roadside in Juola, 29 Sep 1957, Fell 57-1400 nee Jefferson Co.: 1.5 mi NE of Fort 
Atkinson,6 Sep 1981, Nee 21934 (WIS). Milwaukee Co.: Milwaukee, Lake Park, 15 Aug 1 ee Aug 
1939, —_ 983,985 (MIL, WIS).Lincoln Co.: Near Round Lake, 22 Aug 1955, Seymour 16239 (WIS). 

aukee Grafton,8 Sep 1973, Strenger 37 (WIS).Portage Co.: 9 mi W of Plover River, 31 Jul 1964, 
rae a WYOMING. Goshen Co.: Southern Powder River Basin/SE Plains: Goshen Hole, 
Bump Sullivan R ir,ca.4 mi S of Yoder, 1300 m, 28 Aug 1994, ee ee Laramie Co.: 
near Hillsdale, 1 1800 m, 14 aug) 959, Porter & Porter 8019 (DS, CAS). Yakima Co.: Washington, Naches 
Heights, 31 Aug 1937, WC Muenscher & M.V. Muenscher 11292 (BH), 

MEXICO. Baja California: Rio Santo Domingo, 4 mi above Mission, 12 Sep 1930, Wiggins & 
Demaree 4789 (ARIZ). Santa Catarina, 1200 m, Broder 363 (DS). Chihuahua: Mojaracnis 25 Aug 1938, 
Knobloch 5513 (MSC). Temosachi, Nabogame, 28°00'N, 108°30'W, 1800 m,L 6 (ARIZ). Hidalgo: 
near Tolcayuca 2350, 23 io 1970, Baron 55 (CAS, UMO). Zempoala 2450 m, 28 Gen | 975, Ventura 
339 (UMO). Michoacan: 10 mi of Hidalgo, 7 Jul 1947, Sauer 1712 (MO).1/3 mi N on road to La Barca, 

19° 58'N, 102°16'W, 29 Jul 1 ee Sauer 491 (WIS).Morelos: Jiutepec, Jul 1965, Alcocer 8 (ISC). Sonora: 
Las Tierritas de El Temblor, Sierra de El Tigre, 18-24 Aug 1940, White 3393 (DS). 

OUTH AMERIC ERU. Dep. Arequipa: Prov. Caraveli, 20 Km N of Puerto Chala, 15 Jan 

1963, HH IItis, CM. Its, DU eae 578 (WIS). Dep. Lima: Prov. Huarochiri, Surco, 2000 m, 24 

Apr 1940,Asplund 11025 (US). BOLIVIA. Dep. Chuquisaca: Prov.Oropeza, Villa Maria, 12 Apr 1963, D. 

Ugent & Carderas 4948 (WIS). CHILE. Prov. Cautin: Temuco, 110 m, 2 Mar 1935, Montero 2303 (GH). 


— 


= 


5b. Amaranthus powellii sus) poucons ee an aie retero, comb. nov. 
(Fig. 7). BASIONYM: Amaran Thell. in Monde PI. 27 (160):4. 1926. A. hybridus 
L.subsp. bouchonii(Thell. O. Bolos & Vigo in Butl. Inst. Catalana Hist. Nat. 38:89. 1974. A. hybridus 
L. var. bouchonii (Thell.) Lambinon. Candollea, 52:239-279, 1997. TypE: ea Allee du 
Boutant, Chemin remblaye avec des balayures du port, 25 Sep 1925, A.B TYPE: ?) 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 965 


Herbarium — University of California Kiverside 
i i hoch TVA Casiva: Cine 


F 


é Sf 
Det. by Mihai Costea 


PHERBARIUM of The University of Texas at El Paso 
UTEP No # | 140! 
7 


FAMILY 
NAME 


powell Lonrs 
OCATION: Crecon, Henton Co., 


DATE; AUR. OF 1 ALTITUDE feet meter 
HABITAT: 
COLLECTOR: 1.60 Carraway anc o,/, Ne 
IDENTIFIED BY ROY, REFERENCE 
Fic 7 A L ri he L houchonit £ dis fomatha sm ohy 
7 \ } 


f Ly 


966 BRIT.ORG/SIDA 19(4) 


Inflorescence often not strictly erect, more lax with many lateral branches. 
Bracteoles shorter and thinner compared to subsp. powellii, 1.8-2.3 times longer 
than the tepals. Fruit indehiscent, about 1.5 times longer than wide, with the 
pericarp + smooth. Pericarp surface type B to C (Fig. l, E, F). Seeds usually broad- 
ovate to round (rare obovate), 0.9 x 1-12 mm, with the marginal zone sculp- 
tured and the margin of the seed usually rounded (Fig. 2 D). Cells of exotesta 
with prominent anticlinal walls, polygonal, 25-30 um length; periclinal walls 
plane, finely sculptured. The origin unknown with certitude but relationship 
with A. powellii subsp. powellii undeniable. 2n = 


Le specimens examined: EUROPE. near Prov. Noord-Holland: Beverwijk, 23 Aug 

3, Akkerman s.n. (UTEP). FRANCE. Puy-de e: Clermont-Ferand, unknown author, Sep 1960 
ie SMU). Gironde: sae Jul 1960, ee ). SWITZERLAND. Haut-Rhin: Sundhoffen, 9 
Sep 1962, a 5.n. (A). ITAL mbardy: Comosee, 23 Aug 1984, Neuffco 386 (UTEP). 

NORTH AMERICA. cafe STATES. CALIFORNIA. Los Angeles Co.: Arcadia, Winie Way oe 
Santa Anita, : May 1962, Schmid s.n. (DAV). Fresno Co.: Huntington Lake, Big Creek, 1400 
3Aug 1951, Pollard s.n. (CAS).Plumas Co.: Belden, 660 m,21 Sep 1959, Howell 43244 ee ena 
Co.: Riverside, Sanders 20048 (UCR, UTEP). Ventura Co.: Santa Monica Mountains, 1 mi SE of Camarillor, 
200 m, 19 Sep 1958, Raven & Thompson 13726 (CAS). OREGON. Benton Co.: 4.5 mi N of Corvalis on 
ae Drive,6 Aug 1979, Carraway & Verts 11200 (UTEP). por Lake Road,4 Oct 1959, Johnson 

93 (IA). Hood River Co.: 10 Sep 1924, Henderson 300 (MO).M ANA. Lake Co.: in a vicinity of 
Montana Biological Station, Yellow Bay, E shore of Flathead | me 850 1 m,5 Aug 1965, Thomas s.n. 
(DS). PENNSYLVANIA. Bucks Co.: West Bristol, along Pond Street, 20 Jul 1950, me 71768 (CM). 
Thurston Co.: 5 mi W of Olimpia, 15 Aug 1936, Meyer 786 (GH). WASHINGTON. Whatcom Co.: 
Lynden, 21 Aug 1939, Munscher 10558 (BH). Ten Mile, 31 Aug 1943, Munscher 16104 (BH). King Co.: 
Seattle, on the Gas of Washington campus, 17 Sep 1967, Simpson 370 (US). 


6. Amaranthus retroflexus |, Sp. PI. 991.1753. Typr: “Habitat in Pensylvania”. LINN 1117/22. 


Annual with erect stem, up to 1.5(-2) m. Cotyledons lorate (oblong, ratio 6:1, see 
Hickey 1978) 13 x 2-2.5 mm, with acute apex and cuneate base: petiole 5-6 mm 
long. Plant scurfy-villous in and for some distance below the inflorescence, both 
multiseriate and uniseriate trichomes present. Leaves ovate or rhombic-ovate, 
3-7 x 2-4 cm, usually coarsely pubescent beneath, at least along the veins. In- 
florescence usually with many short, thick and crowded branches, the termi- 
nal part about as long as the laterals. Bracteoles (2.5-)4-6(-8) mm, 1-3 times 
longer than the tepals. Tepals 5, + outcurved 2.5-3 mm long, linear-cuneate to 
spathulate, rounded, truncate or emarginate, usually mucronate at the tip, of - 
ten with inconspicuous or pale mid-veins. The tepals much longer than the 
fruit. Fruit circumscissile, about 2 times longer than wide, obovoid, dines 
narrowed toward the stigma region. Dehiscence line in the upper part of the 
ruit, coarsely wrinkled above the dehiscence line. Type of pericarp surface vari- 
able, A to B (Fig. 1 B). Seeds obovate 11-125 x 0.8-0.9 mm, lenticular. Between 
the hilum and the radicle a furrow extends almost to the middle of the seed. 
The seeds are differentiated into a central, smooth zone anda marginal, sculp- 


reo 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 967 


tured zone. Color dark-brown to black, uniform. Pollen grains 20-27 um, with 
30-42 pores of 1.1-1.4 wm in diameter. Density of granules high. 2n = 34. 

Usually A. retroflexus offers no identification problems, yet we have en- 
countered interesting specimens sharing the characteristics of A. retroflexus 
and A. powellii, but somehow different from both. These plants seem to belong 
to a distinct type that occurs in the dry mountains of Arizona and southern 
California. They resemble Amaranthus x tucsonensis recently described by 
Henrickson (1999) from Arizona. Henrickson suggests only one of the parents- 
A. hybridus. In our opinion, based on the description and illustration, the flow- 
ers are closer to A. retroflexus var. delilei (Richt. & Lor.) Thell. but the general 
habit indicates A. powellii. Unfortunately, we could not examine neither the type 
nor the specimens cited in the study. The plants we examined are fertile, sug- 
gesting that they are not Fl hybrids and further research will be necessary to 
clarify if they are only a variety of A. retroflexus or a stabilized hybrid. Examples: 
ARIZONA. Apache Co.: White Mountains, 10 mi S of McKays Peak, 2270 m, 31 Aug 1948, Gould & 
Robinson 4998 (ARIZ). Navajo Co.: Pinaleno Mountains, West Peak, 2600 m, 12 Sep 1989, Mc Laughlin 
5809 (ARIZ); s.1., 2660 m, 6 Sep 1944, Darrow et al. 1179 (SMU). Snow Flat, 2600 m,6 Sep 1944, Darrow 
et al. 1178 (SMU).N of Grasshopper Ruin, 28 Aug 1971, 1800 m, Bohrer 1496 (ARIZ). Gila Co.: Sierra 
Ancha Wilderness Area, 2000 m, 21 Sep 1991, /mdorf271 (ARIZ). CALIFORNIA. San Bernardino Co.: 
San Bernardino Mountains, 116°55'W, 34°11.25N, 1590 m, 18 Sep 1994, Sanders 15690 (UCR, ARIZ). 
NW Shore of Baldwin Lake, 116°49.11W, 36°16.53N, 2043 m, Sanders & Helmkamp 19489 (UCR). 


Distribution and Ecology.—This plant is a native of North America, but has be- 
come a cosmopolitan weed. 


APPENDIX | 


Representative accessions cultivated in Rodale Germplasm, Ames, lowa be- 
tween 1984-1989, deposited at NA 


Amaranthus caudatus 

USA. AMES 5125, 5127, 2191, 2212, 2213. PERU. P| 490437, Pl 490438, Pl 490439, P| 490440. INDIA. P| 
166107, Pl 175039, Pl 166045, P| 480576. NEPAL. AMES 2079, 2165, 2168, 2169, 2182, 2190, Pl 427224, 
P| 427225, P| 427228. 


Amaranthus cru 

CANADA. need rie 5327.USA. AMES 2264, 3216;California, San Rafael, AMES 5165.MEXICO. 
Chapingo, Inia, AMES 5548; Puebla, AMES 5638, 5640; Sonora, San Bernardo, AMES 5320. GUATE- 
MALA. Choatalum, AMES 5271,5275; San Pedro Corcha, AMES 5269; Aldea Choatalum, AMES 5272, 
5273, 5276, 5277 (type 2); Chimaltenango, AMES 5330; Guatemala City, AMES 5278, 5279 (type 1); 
Coban, AMES 5270, 5279 (type 2),5298;Chimaltenango, San Martin, AMES 5299; P| 433228, Pl aie 
Pl 451825, Pl 451826; AMES 5676; PERU. Ayacucho, AMES 5346. PORTUGAL. Evora, AMES 5 
CHINA. AMES 1981, 2016, 5313, 5327. INDIA. R-129 A-2250, Spjut 8809; Pl 288278, P| re . 
288281: AMES 2037, 2228; Coimbatore, AMES 1977, 5386. NEPAL. P| 490656; Pokhara, AMES 2089 
2092; Chauri Jari, AMES 2061, 2089. MALYSIA. AMES 2054. TAIWAN, AMES 1991, 1992, 2201. INDO- 
NESIA. AMES 2042, 2049, 2044. ETHIOPIA. AMES 2003. ZAMBIA. PI! 494768, P| 494769. GHANA. 


968 BRIT.ORG/SIDA 19(4) 


AMES 1959, 1961, 1963, 1968, 1969, 1970, 1971, 1978, 2006, 2008, 2011, 2012, 2013. BENIN. AMES 
5108.NIGERIA. AMES 1973, 2055, 2056, 2057, 2058, 2250. TANZANIA. AMES 1979, 2004. ZAIRE. North 
Shaba Province, AMES 5369. DAHOMEY. AMES 1964, 2000, 2249 (type 1 and 2). ORIGINUNKNOWN. 
AMES 1985, 2081, 2087, 5151, 5303, 5362, 5384, 5686. 


Amaranthus Aglare 
USA. AMES 2211, 3079, 3078, 5140; pe ike 8271, 8404; Pennsylvania, K-254; PI 477917. 
MEXICO. eins AMES 2260;P! 490753; San Miquel, Chiconcuac, AMES 5474; Huazulco, Morelos, 
AMES 5190, 5503, 5637; Oaxaca, AMES 5215; Jautetelco, Morelos, AMES 5503, 5505; Distr ses 
AMES ae care AMES 2085 (type 1 and 2). CHILE. AMES 5355. PERU. Lima, AMES 7 AF- 
GHANISTAN. AMES 5610. INDIA. Coimbatore, AMES 1976;P| 274276, P| 274278, P| 274279, ee 
P| 480787, P| 480790, P| 480791, Pl 480793, Pl 480796, P| 480797, Pl 480798, Pl 480799, P| 480800, PI 
480804, 480806, P| 480807, PI 480810, PI 480811, Pl 480818, Pl 480821, P| 480823, P| 480865, PI! 480870, 
480871.Uttar Pradesh, Pl 166045, Pl 337611, Pl 480814, Pl 480815, Pl 480819, Pl 480820, Pl 480866, 
Pl 480872, P| 480874, P| 480875, PI 490756, AMES 2036. Tamil Nandu Agriculture University, AMES 
2230, Pl 480802, P| 480805, PI 480822, Pl 480974. NEPAL. P| 427228, P| 490759; Kathmandu, AMES 
2071, 2077 (type 1 and 2), 2162; Patra Sanghu, AMES 2163;Tatopani, AMES 2064, 2160, 2167;Chhana, 
AMES 2173;Charyari, AMES 2061, 2155,2185; Nagma, AMES 2166, 2186; Tibrikot, AMES 2159; Chauri, 
Jahari, AMES 2086; Khallaged, AMES 2157, 2158, 2172; Batra, AMES 2178, 2184, 2194, 2253; Rodlkhola, 
AMES 2175; Delhlkot, AMES 2170;Ranka, AMES 2162; Mahadeb, AMES 2171 (type 1 and 2), 2077;Vid 
Mahuri, AMES 2176; Rotehala, AMES 2177; Tata Village, AMES 2062; Chauri Jari AMES 2067. INDONE- 
SIA. Ames 2043. TANZANIA. AMES 1979. NIGERIA. AMES 1975, 2055, 2256.GHANA. AMES 2001 
(Type 2), a ORIGIN UNKNOWN. AMES 5141, 5151, 5152, 5158, 5161, 5163, 2081, 2082, 2088, 
(Type 2), 5361, 5381, 5657, Pl 490757. 


ACKNOWLEDGMENTS 


We are very grateful to Gerold Htigin for his constant help along the last eight 
years. As well to Julio Iranzo who kindly made possible the SEM study. In addi- 
tion, we are much indebted to Jose Luis Carretero for his assistance and the useful 
discussions over the problems of the genus. Moreover, Mihai Costea owes thanks 
to his teachers Vasile Ciocarlan, Livia Ungurean, and Constantin Chirila. We 
are also obliged to Gheorghe Dihoru and Gavril Negrean for their observations 
and suggestions. Our sincere thanks go also to Jonathan Sauer and David Brenner 
who have reviewed the paper providing comments that have improved the qual- 
ity of the manuscript. Many thanks go to Darleene DeMason and Edward Plumer 
for the images. We are also very grateful to all the directors and curators of her- 
baria cited. The paper was written during first author’s Fulbright scholarship 
at University of California, Riverside. 


REFERENCES 


Ac.ten, P. 1959. Amaranthus L.|In: G. Hegi, Illustrierte flora von Mitteleuropa. (second ed.). 
Munchen. 3(2):465-516. 

Actten, P. 1964. Amaranthus L. In: T.G. Tutin, V.|H. Heywood, N.A. Burges, D.H. Valentine, S.M. 
Valters and D.A.Webb, eds. Flora Europaea. University Press, Cambridge. 1:109-110. 

Actten, P. 1970. Amaranthus bouchonii Thell. en France. Monde PI.65:367-368. 

Actten, P. 1972. Amaranthaceae. |In: K.H. Rechinger, ed. Flora lranica. Pp. 1-19. 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 969 


Acone, S.G. and PO. AyiecHo. 1991. The rate of outcrossing in grain amaranths. PI. Breed. 
107:156-160. 

Axeroyo, J. 1993. Amaranthus L. In: T.G. Tutin, N.A. Burges, A.O. Chater, J.R. Edmondson, 
V.H.Heywood, D.M. Moore, D.H. Valentine, S.M.Walters, D.A. Webb, eds. Flora Europaea 
(second ed.). University Press, Cambridge. 1:130-132. 

BaatHLott,W. and N.EHLer. 1977. Raster-elektronen-mikroskopie der epidermis-oberflachen 
von Spermatophyten, Wiesbaden. 

Boos, O.and J. Vico. 1979.Observaciones sobre la flora de los paisos catalanas. Collect. Bot. 
(Barcelona) 11:25-89. 

Brenan, J.RM.1961.Amaranthus in Britain. Watsonia 4:261—280. 

Brenan, J.PM.1981.The genus Amaranthus in Southern Africa. J.S. African Bot.47:451-492. 

Brenner, D.M. 1990. The grain amaranth gene pools. In: Proc. 4th National amaranth sym- 
posium: Perspectives on production, processing and marketing, Minneapolis. 23-25. 
Minnesota Ext. Serv., Univ. Minnesota, St. Paul. Pp. 193-194. 

Brenner, D.M., D.D. BalTENsPERGER, PA. KuLAKow, J.W. LEHMANN, R.L. Myers, M.M. Staseert, and B.B. 
SLEUGH. 2000. Genetic resources and breeding of Amaranthus. P|. Breed. Rev. 19:227-285. 

Burkiit, H.M. 1985. The useful plants of West Tropical Africa. Royal Botanic Gardens Kew. 
1:48-53. 

Cacciato, A. 1982. Amaranthus L.|n:B. Anzalone, A. Becherer, F. Ehrendorfer, H.Merxmiller, H. 
Metlesics, F. Rasetti, T. Reichstein, |. Segelberg, eds. Pignatti S., Flora d'Italia. Edagricole, 
Bologna. 1:177-182. 

Carretero, J.L.1979.El genero Amaranthus en Espana. Collect. Bot. (Barcelona) 11:105-142. 

Carretero, J.L. 1985. Consideraciones sobre las Amaranthaceas Ibericas. Anales Jard. Bot. 
Madrid 41:271-286. 

Carretero, J.L.1990.Amaranthus L.|In:S.Castroviejo, M.Lainz,G.Lopez Gonzales, P. Montserrat, 
F. Munoz Garmendia, J. Paiva and L. Villar, eds. Flora Iberica. Plantas vasculares de la 
Peninsula lberica e Islas Baleares, Platanaceae-Plumbaginaceae. Jardin Botanico, Madrid. 
2:554-569. 

Cuan, K.F. and M. Sun. 1997. Genetic diversity and relationships detected by isozyme and 
RAPD analysis of crop and wild species of Amaranthus. Theor. Appl. Genet. 95:865-873. 

CHEREPANOV (CzerEPANOV), S.K. 1995. Vascular plants of Russia and adjacent states (the former 
USSR). University Press, Cambridge. Pp. 6-9. 

Ciements, S.E. 1992. Chenopodiaceae and Amaranthaceae of New York State. Contr. Flora 
New York State. 10(485). Univ. State of New York, Albany. 

Coons, M.P.1975.The genus Amaranthus in Ecuador. Ph.D.diss. Indiana Univ. Bloomington. 

Coons, M.P. 1978. The status of Amaranthus hybridus L.in South America. The taxonomic 
problem. Ci. Naturaleza. 19:66-71. 

Coons, M.P. 1982 Relationships of Amaranthus caudatus. Econ. Bot. 36:129-146. 

Costa, M. 1997a. The genus Amaranthus L., section Amaranthus in Romania. Acta Horti. 
Buc. (University of Bucharest) 1995/1996, 105-125 (in English). 

Costea, M. 1997b. Morphology of fruit in some species of the genus Amaranthus L. Acta 
Horti. Buc. (University of Bucharest) 53:135-149 (in Romanian). 


970 BRIT.ORG/SIDA 19(4) 


Costea, M. 1977c. Morphology of seed in some species of the genus Amaranthus L. Acta 
Horti. Buc. (University of Bucharest) 53:24-37 (in Romanian). 

Costea,M.1998a.Amaranthus L., Subgenus Albersia (Kunth) Gren. & Godr.in Romania., Rev. 
Rom. Biol. (Romanian Academy of Science) 43:95-112 (in English). 

Costea, M. 1998b. Monograph of the genus Amaranthus L.in Romania. Ph.D. diss., Univer- 
sity of Bucharest, College of Biology (in Romanian). 

Costea, M. and D.A. DeMason. 2001. Stem in Amaranthus L.-taxonomic significance. Bull. 
Torrey Bot. Club. 128:254-281. 

Covaco, A. 1976.Les Amaranthus de Madere et de Acores. Bol. Soc, Port, Cienc. Nat. 16:79-89. 

Covas, G.1939.Los generos de Amaranthaceas Argentinas. Rev. Argent. Agron, 6:282-303. 

Covas, G. 1941.Las Amaranthaceas Bonariensis. Darwiniana 5:329-368. 

Covas,G. 1992. Clave para la identification de los amarantos cultivados y especies silvestres 
utilizables como hortalizas of forrajeras. Amarantos Novedades e Informaciones. 12:9-12. 

Dostat, J. 1950. Amaranthus in Kvetena CSR a ilustravny klik k urceni vsech cevnatych 
rostlin. Praha. Pp. 441-448. 

Duke, JA. 1961. Amaranthaceae. |n: Flora of Panama. Ann. Missouri Bot. Gard. 48:348-392. 

EHRENDORFER, F. 1973. Liste der gefaspflanzen Mitteleuropas (2-end ed.). Stuttgart. 

Evasson, U. 1987. Amaranthus L. In: GH. Harling and L. Andersen, eds. Flora of Ecuador 
28:20-36, ae Council to! Notaie Pelee ot Botany. 

Eviasson, U. 1988.Floral fA tt 
in the New World and he ee Islands. J. Linn. Soc., Bot. 96: Zape204. 

EroTman, G, 1966. Pollen morphology and plant taxonomy. Angiosperms. Alquimist & 
Wiksell, Stokholm. 

ESPARZA-SANDOVAL, S., G. ALEJANDRE-ITURBIDE, and Y. Herrera-Arrieta. 1996. Foliar anatomy and 
morphology of seeds in some Mexican species of Amaranthus. Phytologia 81:273-281. 

Espitia-Rancel, E. 1994. Breeding of grain amaranth. In: O. Paredes-Ldpez, ed. Amaranth: 
biology, chemistry, and technology. CRC Press, Boca Raton, FL. Pp. 23-38. 

Georce, A.S. and G. Bent. 1981. Amaranthaceae. In: J. Jessop, ed. Flora of Central Australia, 
Reed, Sidney, N.S.W, Australia. Pp. 76-85. 

Gravis, A.and C. CoNsTANTINESCO. 1907.Contribution a l’anatomie des Amaranthaceae. Arch. 
Inst. bot. Univ. Liege 4:1-65. 

Greizerstein, E.and L. Pocaio. 1992. Estudios citogenetico de seis hibridos inter-especificos 
de Amaranthus Darwiniana 31:159-165. 

GreIZERSTEIN, E.,C.A. NARANJO, and L. Pocscio. 1997. Karyological studies in five wild species of 
Amaranths. Cytologia (Tokyo) 62:115-120. 

GreuTer, W.1981.Med-Checklist Notulae 3.Willdenowia 11:3-43. 

GreuTer, W., H.M. Buroet, and G. Lone. 1984. Med-checklist. A critical inventory of vascular 
plants of the circum-mediterranean countires. Conservatoire et Jardin Botaniques de 
la Ville de Geneve, Geneve. 1:46-48. 

Gusev, J.D. 1972. The survey of the genus Amaranthus in U.S.S.R. Bot. Zhurn. S,S,S.R. 57: 
457-464. 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 971 


Hauptu, H. and S. Jain. 1984. Allozime variation and evolutionary relationships of grain 
amaranths (Amaranthus spp.). Theor. Appl. Genet. 69:153-165. 

Havetu, H.and S. Jain. 1985.Genetic variation in outcrossing rate and correlated floral traits 
in a population of grain amaranth (Amaranthus cruentus L.). Genetica 66:21-27 

Henrickson, J. 1993. Amaranthaceae. In: J.C. Hickman, ed. Jepson manual: higher plants of 
California. Univ. California Press, Berkeley. Pp. 130-134. 

HENRICKSON, J. 1999. Studies in New World Amaranthus (Amaranthaceae). Sida 18:783-807. 

Hickey, L.J. 1978. A revised classification of the architecture of dicotyledonous leaves. In: 
CR. Metcalfe and L.Chalk,eds. Anatomy of Dicotyledons (2-end ed.). Oxford. Pp. 25-39. 

DEN . Wee: Die venieltng von Amaranthus- Arten in der stdlichen und mittleren 

len gebieten.Phytocoenologia 14:289-379. 

Huain, G. 1987. Einige Bemerkungen zu wenig bekannten Amaranthus-sippen 
(Amaranthaceae) Mitteleuropas.Willdenowia 16:453-478. 

HuNzINker, A.T. 1987.Taxonomia de las especies de amarantos cultivados y de los silvestres 
relacionados. In Goldberg, ed. Actas de las Primeras Jornadas Nacionales Sobre 
Amarantos, Faculdad de Agronomia ,La Pampa, Argentina. 

JAIN, S.K., H.Hauetti,and K.R.Vaiova 1982. Outcrossing rate in grain amaranths. J. Hered. 73:71-72. 

Javas, J. and J. Suominen. 1980. Atlas florae Europaeae, Vol. 5, Helsinki, Societas Biologica 
Fennica, Vanamo. Pp. 90-98. 

KAUFFMAN, C.S.and P.Waconer. 1984.Grain amaranth: An overview of research and produc- 
tion methods. Rodale Research Center. New Crops Dept. Report NC-84/6. Rodale Press 
Inc., Emmaus, PA. 

KAUFFMAN, C.S. and L.F. Weser. 1988. Grain amaranth. In: J. Janick and J.E. Simons, eds. Ad- 
vances in new crops: Proc. of the first national symposium new crops, research, devel- 
opment, economics. Indianapolis, 23-36 Oct. Timber Press, Portland. 

KAUFEMAN, C.S. 1992. Realizing the potential of grain amaranth. Food Rev. Int. 8:5-21. 

KERGUELEN, M. 1993. Index sinonimique de la flore de france. Muséum National d'Histoire 
Naturelle, Paris. Pp. 13. 

Kirkpatrick, B.A. 1995. Interspecific relationships within the genus Amaranthus 
(Amaranthaceae). Ph.D. diss. Texax A & M Univ., College Station. 

Kvoprer, K. and J. Roget. 1989a. Beitrage zur systematik, morphologie und anatomie der 
gattung Amaranthus L. 1. Karpomorphologie und-anatomie ausgewahlter vertreter. 
Gleditschia 17:3-21. 

Ktoprer, K. and J. Roset. 1989b. Beitrge zur systematik, morphologie und anatomie der 
gattung Amaranthus L. 2. Samenmorphologie und-anatomie ausgewahlter vertreter. 
Gleditschia 17:171-182. 

Kovacney, |. 1977. Amaranthus species in Europe. Priroda (Sofia, Bulgaria) 26:32-39. 

Kowal, T. 1954. Cechy morfologiczne | anatomiczne nasion rodzaju Amaranthus L. oraz 
klucze do ich oznaczania. Monogr. Bot. (Warszawa) 170-193. 

KutaKow, PA., H. Hauptu, and S. Jain. 1985. Genetics of grain Amaranths. Mendelian analysis 
of six colour characteristics. J. Hered. 76:27-30. 


972 BRIT.ORG/SIDA 19(4) 


Kutakow, PA. 1987.Genetics of grain Amaranths. ll. The inheritance of determinance, panicle 
orientation, dwarfism, and embryo color in Amaranthus caudatus. J. Hered. 78:293-297. 

Kutakow, P.A.and S.K. Jain. 1990. Grain amaranth—crop species, evolution, and genetics. In: 
Amaranth: perspectives on production, processing, and marketing. Minnesota Exten- 
sion Service, St. Paul. Pp. 105-114. 

KUNTH, C.S. 1917. Nova genera et species plantarum. Voyage de Humboldt et Bonplanct. 
2:194-195. 

LAMBINON, J., J-E. De LANGHE, L. Detvosatte, and J. Duvicneaub. 1992. Amaranthus L. In: Nouvelle 
flore de la Belgique, du Grand Duché de Luxembourg, du Nord de la France et des 
Regions voisines. Pp. 258-164. 

Lanoue, K.Z., P.G. Wotr, S. BRowninG, and E.E. Hoop. 1996. Phylogenetic analysis of restriction- 
site variation in wild and cultivated Amaranthus species (Amaranthaceae). Theor. Appl. 
Genet. 93:722-732. 

Moraariu, |. 1952. Genus Amaranthus L. In: Savulescu, ed. Flora R.RR. Romanian Academy, 
Bucharest. 1:587-607 (in Romanian). 

Mosyakin, S.L. and K.R. Rosertson. 1996. New infrageneric combination in Amaranthus 
(Amaranthaceae) Ann. Bot. Fennici 33:275-281. 

Murray, MJ. 1940. Colchicine induced tetraploid in dioecious and monoecious species of 
Amaranthaceae. J. Hered. 31:477-485. 

Nowicke, J.W. 1993. Pollen morphology and exine ultrastructure in Caryophyllales In: H.D. 
Behnke and T.J.Mahbri,eds.Evolution and Systematics in Caryophyliales. Springer Verlag. 
Pp. 165-221, 

Pat, M. and T.N. KHosHoo, 1972. Evolution and improvements of cultivated Amaranths. V. 
Inviability, weakness, and sterility in hybrids. J. Hered. 63:78-82. 

Pat, M. and T.N. KHosHoo. 1973. Evolution and improvements of cultivated Amaranths VI. 
Cytogenetic relationship in grain types. Theor. Appl. Genet. 43:343-350. 

Priszter, S. 1949. Amaranthus—viszgalatok. 1.Amaranhtus- hibridek Magyarorszagon. Index 
Horti Bot. Univ. Budapest 7:116-149. 

Priszter, S. 1953, Revisio critica specierum generis amaranthi in Hungaria. Agrartud Egyet. 
Kert Szdlész. Foisk. Evk. 2:121-262. 

Priszter, S. 1958. Uber die bisher bekannten bastarde der gatung Amaranthus. Bauhinia 
1:126-135 

Rosertson, K.R. 1981. The genera of Amaranthaceae in the southeastern United States. J. 
Arnold Arbor. 62:267-313. 

SAMMOUR, R.H.,M.A. HAmMoup, and S.A.A. Atta. 1993. Electrophoretic variations in Amaranthus. 
Bot. Bull. Acad. Sin. 34:37-42. 

SANCHEZ-Det PINO I., H.F. Otvera, and J. Vatpes. 1999. La familia Amaranthaceae en la flora 
halofila y gipsofila de Mexico. Anales Inst. Biol. Univ. Nac. Auton. Mexico, Bot. 70:29-135. 

Sauer, J.D. 1950. The grain amaranths: a survey of their history and classification. Ann. Mis- 
souri Bot. Gard. 37:561-362. 

Sauer, J.D. 1955. Revision of the dioecious amaranths. Madrono 13:5-46. 


COSTEA ET AL., AMARANTHUS HYBRIDUS SPECIES COMPLEX 973 


Sauer, J.D. 1967. The grain amaranths and their relatives: a revised taxonomic and geo- 
graphic survey. Ann. Missouri Bot. Gard, 54:103-137. 

Sauer, J.D. 1976. Grain amaranths. In: N.W. Simmonds, Ed. Evolution of crop plants. Long- 
man. Pp.4-7. 

Sauer, J.D. 1977. The history of the grain amaranth and their use and cultivation around 
the world. In” Proceedings of the first amaranth seminar, Rodale Press Inc., Emmaus, PA. 
Pp.9-15. 

Sauer, J.D. 1993. Historical geography of crop plants. A select roster.C.R.C. Press, Boca Raton, 
FL. Pp. 9-14. 

Scuinz, H. 1933. Amaranthaceae In: A. Engler & K. Prantl, eds., Die naturlichen 
pflanzenfamilien. (2-end ed.) Leipzig. 2(16c):7-85. 

Stace, A. 1991.Amaranthus L. In: New flora of British Isles. Cambridge. Pp. 186-190. 

Stace, C.A. 1997. Amaranthus L. In: New flora of British Isles, second ed. Cambridge. Pp. 
150-155. 

Standley, PC. 1914. New or notable species of Amaranthus. Bull. Torrey Bot. Club 41: 
505-510. 

STANDLEY, P.-C. 1917. Amaranthaceae in North American flora. 21:95-169. 

StanbLey PC. 1937. Amaranthaceae. In: Flora of Peru. Fieldiana, Bot. 13:478-518. 

Stanotey, PC. and J.A. Steyermark. 1946. Amaranthus In: Flora of Guatemala. Fieldiana, Bot. 
24:152-157. 

THELLUNG, A. 1907. Beitrage zur Adventivflora der Schweiz 5. Vierteljahrsschr. Naturf. Ges. 
Zurich 52:434-473. 

THELLUNG, A. 1914. Amaranthus in P. Ascherson and P. Graebner, eds., Synopsis der 
Mitteleuropdischen Fora 5:225-356. 

THELLUNG, A. 1919. Beitrage zur adventivflora der Schweiz 3. Vierteljahrsschr. Naturf. Ges. 
Zurich 64:684-815. 

THELLUNG, A. 1926. Amaranthus bouchonii Thell. spec. (?) nov. Monde Plantarum 27:4-5. 

TIMONIN, A.K. 1986a. On the taxonomic value of stomatal characteristics and evolution of 
the stomatal complex in flowering plants. Biul. Mosk o. va. ispitatelei prirodi, T 91 ser. 1. 

TIMONIN, A.K. 1986b.On some peculiarities in the formation of stomatal apparatus in cases 
of their polymorphism in dicotyledons. Biologischie Nauki 2:70-75 (in Russian). 

Townsenp, C.C. 1974. Amaranthaceae. |n:E. Nasir and A\l. Ali, eds. Flora of West Pakistan No. 
71. Royal Bot. Gard, Kew. 

Townsenp, C.C. 1985. Amaranthaceae. In: R.M. Polhill, ed. Flora of Tropical East Africa. A.A. 
Balkema, Rotterdam, Boston 

Townsenb, C.C. 1988.Amaranthus. In: E.Launert, ed. Flora Zambesiaca 9(1): 45-60. London. 

Transue, D.K., D.J. FAIRBANKS, L.R. RoBinson, and W.R. ANbersEN. 1994. Species identification by 
RAPD analysis of grain amaranth genetic resources. Crop. Sci. 34:1385-1389. 

Tucker J.M. and J.D. Sauer. 1958. Aberrant amaranth populations of the Sacramento-San 
Joaquin Delta, California. Madrofo 14:252-261. 

USDA, ARS. 2000. National genetic resources program. Germplasm resources informa- 


974 BRIT.ORG/SIDA 19(4) 


tion network-(GRIN). [Online Database] National germplasm resources laboratory, 
Beltsville, Maryland. http://www.ars-grin.gov. 

Wituams, J.T.and D. Brenner. 1995, Grain amaranth (Amaranthus species). In: J.T.Williams, ed. 
Cereals and pseudocereals, Chapman and Hall, London. Pp. 129-186. 

Witkin, P1992. The status of Amaranthus bouchonii Thell. within Amaranthus L. section 
Amaranthus: new evidence from morphogy and isoenzyme. J. Linn. Soc., Bot. 108: 
253-267. 

Witson, C.L. 1924. Medullary bundle in relation to primary vascular system in 
Chenopodiaceae & Amaranthaceae. Bot. Gaz. 78:175-99. 


NOTES ON SOME LITTLE KNOWN AMARANTHUS 
TAXA (AMARANTHACEAE) IN THE UNITED STATES 


Mihai Costea Andrew Sanders 
University of Agronomical Sciences Herbarium 
Dept. of Botany Dept. of Botany and Plant Science 
Bd. Marasti, 71331, Sector 1 University of California 
Bucuresti, ROMANIA Riverside, CA, 92521-0102, U.S.A. 
coste_amihai@hotmail.com andrew.sanders@ucr.edu 
Giles Waines 


Dept. of Botany and Plant Science 
University of California 
Riverside, CA, 92521-0102, U.S.A. 
giles.waines@ucr.edu 


ABSTRACT 


A L Lt: ndi 


bility in the United States are analyzed ene classi- 


ariab 
cal morphological a some new Scanning piace Microscope characters. Two new combinations 
within A. blitum are proposed, A. blitum subsp. Sieraces ane A. blitum subsp. emarginatus var. 


pseudogracilis. For the first time, A. graecizans L gin Neh America. Some 
le 


nomenclatural issues involving A. blitum L. and A. graecizans | 


RESUMEN 


Se analiza Amaranthus blitum L y su variabilidad intraespecifica en los Estados Unidos, utilizando 


los caracteres elaeleos ya algunos nuevos obsepvados con ne Microscopio Electrénico de Barrido. Se 
litum subsp. oleraceus y A. blitum 


ae sea shes var. peeudogracilis, Por primera vez se confirma la presencia de Amaranthus 
graecizans en Norte Am a. Se analizan también algunos problemas de nomenclatura de 
Amaranthus oe y eee 


INTRODUCTION 


+] 


Many of the roughly 75 species of the genus Amar. native to the Ameri- 
cas. Of these, 10 are dioecious (Sauer 1955) and about 40 monoecious. Approxi- 
mately 10 of these American species are now widespread weeds, some of them 
being listed among the worst weeds of the world (e.g, A. retroflexus, A. hybridus, 
A. powellii, and A. viridis—Holm et al. 1977; Holm et al. 1997). In return, the 
Americas have received few species of weedy amaranths from the Old World, 
though A. blitum (subsp. blitum) and A. graecizans are examples. Among the 
species introduced to North America, A. blitum is especially worthy of atten- 
tion because in Europe, Africa and Asia it is often a troublesome weed in irri- 
gated crops (Holm et al. 1977; Hugin 1986, 1987; Costea 1998a, b). In the United 
States, Teitz et al. (1990) stated that in “recent years A. lividus ( A. blitum) has 


SIDA 19(4): 975 - 992. 2001 


976 BRIT.ORG/SIDA 19(4) 


become the most serious weed problem in vegetable production in Ohio” in crops 
such as lettuce, radish, celery and carrot. Furthermore, A. blitum isa valuable leaf 
vegetable crop and is cultivated as such in Asia, Africa, and the Pacific Islands. 

After a survey of the 40 of the most important herbarium collections in 
the United States, as well as the floristic literature, for a revision of the A. 
hybridus complex, it became obvious that A. blitum is inadequately understood 
in North America. This taxon is often confused with A. viridis and its infraspe- 
cific variability has received no attention. 

Likewise, A. graecizans is poorly understood in North America. The name 
Amaranthus graecizans has been widely used in North America (e.g., Kearny 
& Peebles 1960; Hitchcock & Cronquist 1973; McGregor 1986), but has always 
been misapplied to A. albus or A. blitoides. In view of this past nomenclatural 
confusion, the name A.graecizans is probably regarded with suspicion by many 
botanists in North America. We show that this species has been introduced into 
the United States, and suggest that it may still exist. 

The purpose of this paper is not a comprehensive review of the status of 
these taxa in the United States, but rather to provide the necessary information 
from which a better understanding of them can develop. To facilitate correct 
identification, detailed descriptions are provided for each taxon in this species 
group using both traditional characters and new ones such as trichomes, sculp- 
ture of the seed coat and pollen morphology. 

The seeds are usually differentiated in a central, convex zone and a mar- 
ginal, plane zone, exceptions from this rule being rare (A. blitum subsp. 
oleraceus). The SEM characters of seeds pertain to the ornamentation of the 
exotesta in the marginal zone, as observed under 300x, or more magnification. 
The sculpturing of the seeds is described using the terminology proposed by 
Barthlott and Ehler (1977). The appearance of the anticlinal (prominent or in- 
conspicuous) and periclinal (flat, concave or convex with the sculpture of the 
epicuticular waxes smooth or punctiform) walls of the epidermal cells is noted. 

The pollen grains are pantoporate, apolar, small (with D = 18-28 wm) and 
generally have more than 18 sunken pores, uniformly distributed and having 
the apertural membrane granulated. The tectum has granules or spinules. See 
also Eliasson (1988), Nowicke (1993), Costea (1998a,b). Pollen characters include: 
the diameter of the pollen grain; number and diameter of pores; density of gran- 
ules or spinules on the tectum according to the following scale: high = 20-30 
granules or more/| um?: medium = 10- 19 granules/um? and low = 2-9 gran- 
ules /1 pm2. The SEM observations were carried on with a Hitachi S-4100 SEM 
at ISKV, using a Bio-Rad Sputter-Coatter SC-500. 

Some nomenclatural issues with respect to these introduced taxa are dis- 
cussed as well, including two new combinations 


— 


COSTEA ET Al 977 


NOMENCLATURE 


Amaranthus blitum 

Amaranthus blitum and A. lividus have been long known to be synonyms 

(Hooker 1885; Thellung 1914) and the choice between the two names generated 

an interesting nomenclatural problem, since both taxa were described by 

Linneus in 1753. Both Moquin-Tandon (1849) and Thellung (1914) considered 

A. blitum to include A. graecizans, based on the synonyms they cite under A. 

blitum. Because of this confusion Brenan (1961) stated that the name A. blitum 

should be rejected as nomen confusum, and he used instead A. lividus. More re- 
cently, Brenan and Townsend (1980) proposed explicitly that A. blitum be placed 
on the list of nomina rejicienda as nomen ambiguum. The Committee for 

Spermatophyta recommend that A. blitum not be rejected (Brumitt 1984). This 

decision was made on the basis of the fact that “since the last century, and in 

the present century (A. blitum) was used in the correct sense of A. lividus or has 
not been used at all.” 

In order to establish the priority of two possible legitimate names of equal 
priority it is necessary to determine which author was the first to unite them. 
That choice is definitive (Art 57. 2 Berlin Code, Art 11.4 Tokyo Code). 

Thellung (1914) selected the name A. lividus in which he included: 

Amaranthus blitum L. = elit a lividus proles ascendes—wild plants with 

rostrate stems and small leaves. 

i lividus L. = Aaranta: lividus proles lividus (“typicus”)—culti- 
vated forms with Os ascending or erect stems and large leaves, en- 
tire plant red-colored. 

Amaranthus a a Amaranthus lividus proles oleraceus—cultivated 
plants, very much like proles lividus but green-colore 

According to Filias et al. (1980) the choice between the two names was made 

before Thellung (1914) by Hooker in Flora of British India (1885), but in favor of 

A. blitum. Therefore, Amaranthus blitum is the valid name and Recommenda- 

tion 14a of the Tokyo Code (1998) should be followed. Consequently according 

to Filias et al. 1980) the wild, small-leaved form retains the name A. blitum L., 

being typified by the specimen on the Linnean Herbarium 1117/4. The culti- 

vated forms—A. oleraceus and A. lividus—are together A. blitum var oleraceus 

(L.) Hooker fil, this name being typified by the specimen on sheet 1117/13. 

It is interesting that even though the name A. blitum has predominantly 
been used in recent years, the name A. lividus it is still employed, even in some 
recent treatments of the genus (e.g., Iownsend 1988; Akeroyd 1993). 
Amaranthus graecizans 
Linnaeus’ phrase name for A. graecizans, A. floribus triandris conglomeratis 
axillaribus, foliis lanceolatis obtusis, is appropriate but the geographic source 


978 BRIT.ORG/SIDA 19(4) 


he reported, Virginia, is incorrect. Uline and Bray (1884) reported from America 
the following related taxa: A. blitoides S. Wats., A.albus L., A. graecizans L. and 
A. blitumL., included by Moquin- Tandon (1849) in the former genus Pyxidium. 
The two authors observed that the American plants of what was called A. 
graecizans were in fact the same as A. albus (supposed to have an American 
origin) and thus they synonymised the two names, choosing as valid, the first 
one, which was A. graecizans. The nomenclature confusion at that moment was 
even more complex because as we have shown, at least before Thellung (1914), 
Moquin-Tandon (1849) was using the name A. blitum to designate what we call 
now A. graecizans and other authors (e.g., Hooker 1885) to include both taxa. In 
such circumstances it is easy to understand why Uline and Bray (1884) were 
actually able to recognize clearly only A. blitoides “by its four or five sepals, thick 
at the base, and by the prostrate habit.” From that point, the idea that A. 
graecizans = A. albus persisted in many North American floras even as late as 


1986 (McGregor 1986). 
Amaranthus graecizans was probably frequently misapplied to A. blitoides 
§ I y freq y PP 
due to the resemblance between A. graeci grecizansand A. blitoi 


In both editions of the checklist of the vascular plants of the United ae 
Canada and Greenland (Kartesz 1980; Kartesz 1994) A. graecizans is no longer 
listed among the species of the genus Amaranthus. 


1. Amaranthus blitum L., Sp. Pl. 990. 1753. Tyee: “Habitat in Europa temperatiore”, LINN 


1117/4 (the correct specimen, Fillias et al. 1980) 


Annual with stems up to lm, procumbent, accadue to erect, glabrous or with 
sparse, multicellular, uniseriate hairs. Leaves, 2-8 x 1-5 cm, rhombic-ovate, 
short-cuneate to truncate at base, shallowly emarginate to obtuse-retuse at apex, 
somewhat fleshy, green or reddish. Flowers both in axillary cymose clusters 
and in a terminal variable inflorescence: short, dense and thick to long thin, 
branched and flexuous. Bracteoles triangular-ovate, shorter than the tepals, with 
the mid-vein green, usually with two lateral branches. Tepals 3, equal, 1.5-2 mm 
long and 0.2-0.4 mm wide, linear to (rarely) spatulate. One tepal (rarely 2) with 
green mid-vein enlarged above the middle, the others entirely membranous, 
with the mid-vein hardly noticeable. Fruit indehiscent, 1-1.5 mm longer than 
wide, ellipsoidal to globose, gradually or abruptly-narrowed toward the stigma 
region. Locule only slightly larger than the seed. Chromosome number: 2n=34. 

On the basis of morphology and ecology, three subspecies can be recog- 
nized. Although the habitus of plants, morphology of stem and size of leaves 
usually varies between subspecies (see descriptions), the most reliable charac- 
ters are the size of seeds and fruits. Also the infraspecific taxa of A. blitum can 
be recognized from the plantlet stage which is uncommon for amaranths, well 
known to be very difficult to separate in this phenophase. 


— 


COSTEA ET Al 979 


1. Cotyledons with rounded to truncate apex, 9-18 x 3-6 mm. Pollen grains with 
pores of 2.4—3.3 um.Fruit of 1.7-2.6(-3) mm long, with the pericarp 4-layered. Seeds 
of 1.1-1.8 mm in diameter 2 
2. Cotyledons of 15-18 x 5-6 mm. Bracteoles with the midvein branched. Seeds of 
1.2-1,.6(-1.9) x 1.2-1.6(-1.9) mm with inconspicuous sculpture and rounded 


margins subsp. oleraceus (Figs. 4, 5C) 
2. Cotyledons : 9-10 x 3 - 3.3 mm. Bracteoles with the mid-vein unbranc hed. 
Seeds of (1.1-)1.2 x 1-1.1 mm with evident sculpture and acute mar 


subsp. blitum (Fs 1,5A, 5D) 
1. Cotyledons with acute apex,6-7 x 3-6 mm. Pollen grains with pores of 1.6-1.9 

Fruit of 1.2-1.8 mm, with the pericarp 3-layered. Seeds of 0.8-1.1 mm in 

diameter subsp. emarg inatus (Figs. 2, 3, 5C, 5E) 


bo 


la. seieicp yas blitum subsp. blitum (Fig. 1). Albersia blitum (L.) Kunth, Fl. Berol., ed. 
1838. Euxolus blitum (L.) Gren., Mem. Soc. Emul. Doubs. ser 3, 10:652. 1869. [Flora 
oe oe 1869). 


Amaranthus ascendens Loisel., Not. Fl. France 141.1810. en. blitum L. var. (“b*) ascendens 
(“adscendens”) (Loisel.) DC., Cat. Pl. Horti Monsp. 4. 1813. Amaranthus lividus L. proles 
ascendens (Loisel.) Thell., Ascherson & Graebner, Syn. ee FI. 5:321. 1914. steels 
lividus L. var. ascendens (Loisel.) Thell. ex Hayw. & Druce, Advent. Flora Tweedside 177. 
Amaranthus on L. subsp. ascendens (Loisel.) Thell. ex Wachter, Heukels, Geill. a 
Neder, ed. 11, 169. 193 

Amaranthus eye auct., non L. 


The plants are normally green. The stem procumbent to ascending and without 
prominent ribs. Cotyledons lanceolate, 9-10 x 3-3.3 mm, with rounded apex and 
cuneate base; petiole 4-5 mm. Fruit, 2 mm long, when fresh with the pericarp 
almost smooth, when dried irregularly wrinkled in the zone with the seed, the 
rest of the fruit being smooth. Stigma branches, thin (0.9-1.1 mm long and 0.2- 
0.25 mm wide at the base) often no longer observable when the fruit is dried. 
Seeds, (1.1-)L.2 x 1-l.1 mm, broad-ovate to circular, differentiated into central 
and marginal zones. The marginal zone sculptured with the cells of the exotesta 
polygonal, 25-40 ym length, with the anticlinal walls prominent and the 
periclinal walls plane and minutelly punctiform sculptured (Fig. 5A). The margin 
of the seed acute. Color dark—brown to black with the marginal zone usually 
paler. Pollen grains 19-23 um with 32-45 pores of 2.4-3.3 um diameter (Fig. 5D). 

Distribution and ecology—Native to Mediterranean region, Eurasia and 
North Africa where it was cultivated as a potherb until the 18th century, when 
it was largely replaced by Spinacia oleracea. In these regions it is a frequent 
weed in vegetable gardens, and in waste places. In the United States this name 
was usually misapplied to A. blitum subsp. emarginatus which is more frequent. 
Subspecies blitum appears casua lly in waste places in urban settings. The only 
herbarium specimen seen from the United States was: New York. Central Park, 
1861, Austin s.n (GH). 


1b. Amaranthus blitum subsp. emarginatus (Mog. ex Uline & Bray) Carretero, 


980 BRIT.ORG/SIDA 19(4) 


Herbarium — U 


Amaranthus blitum L. subsp. blitum 


Det. by Mihara Costea 


Fic. 1. Amaranthus blitum subsp. blitum. 


COSTEA ET AL 981 


Munoz enue & Pedrol, Ann. J. Bot. Madrid 44:599. 1987. sella 
emarginatus Mogq. ex Uline & Bray, Bot. Gaz. (Crawfordsville) 19:319. 1894. TYPE: INDONESI 
ee cee L. ?, Java, 1842-44, Zollinger 1646” (LECOTYPE: P; ISOLECTOTYPE: - 
G, STR; cf. Htigin 1987. 461) 


Euxolus viridis L. var. (e) polygonoides Moq., DC., Prodr. 13(2):273. 1849. Amaranthus ascendens 
Loisel. var. polygonoides(Mogq.) Thell., Mem. Soc. Sci. Nat. Cherbourg 38:215.1912. Amaranthus 
lividus L. proles polygonoides (Moq.) Thell., Ascherson e Graebner, Syn. sence FL. 5:321. 
1914. Amaranthus lividus L. var. ee aiagaal dee ex Druce, Bot. Soc. Exch. Club Brit. 
Isles 5:574. 1920. Amaranthus lividus L. subsp. ae (Moq.) am ex Probst, 
Wolladventivfl. nee 74,1949, hee ascendens subsp. po Bai eciage ell. 
ex Priszter, Agartud. Egyet. Kert- Szologazdasagtud. Karanak Evk. 2:221. 1953. Amaranthus 
blitum subsp. pol les (Zollinger ex Mog.) Carretero, Ann. J. Bot. Madrid 41:276. 1985. 


fo) 


Similar to Amaranthus blitum subsp. blitum, differing in the following charac- 
ters: slender stems, up to 70 cm long, prostrate to ascending. Cotyledons nar- 
row-elliptic, 6-7 x 2-2.3mm with acute apex and base; petiole about 4 mm long, 
Leaves not fleshy, long-cuneate at base and deeply emarginate to bilobed at apex. 
Tepals 2(-3)0.75-1.5(-L8) mm long, obtuse. Fruit longer than the tepals, glo- 
bose to ovoid (1.0-)1.2-1.8(-2.0) mm long, 1-15 mm longer than wide, evidently 
larger than the seed. Seeds 0.8-L1 x 0.8-1.1 mm, circular, with the hilum promi- 
nent on the outline of the seed. The marginal zone more evidently sculptured 
than in subsp. blitum (Fig. 5B). Pollen grains as in subsp. blitum but pores of 
14-18 wm in diameter (Fig. 5E). Chromosome numbers: 2n=34. The appropri- 
ate rank of this taxon requires further investigation. Htigin (1986, 1987) and 
Costea (1998a,b) treated it as a species. 

The habit of the plants is variable: 


1. Prostrate plants, leaves (0.75—)1—2(-3.5) ¢ infl irely formed from 
axillary cymes (terminal part absent or reduced var. emarginatus 
Ascending more ith leaves 2—4(-6) cm long, inflorescence mainly 
terminal, long, thin, -lexuous var. pseudogracilis 


a. Amaranthus blitum subsp. emarginatus Mog. ex Uline & Bray var. emarginatus 


b. Amaranthus blitum subsp. emarginatus var. pseudogracilis (Thell.) Costea, 
comb. et stat. nov. (Fig. 3). BAsionyM: Amaranthus lividus L. aaa polygonoides f. 
pseudogracilis Thell. Ascherson & Graebner, Syn. Mitteleur. Fl. 5 1914. Amaranthus 
emarginatus subsp. pseudogracilis (Thell.) Htigin, Willd. 16: 463. oF = “im Botanischen 
Garten Strasburg,” 8.8 1904, A. Ludwig 12130 (B) specifying “Unter dem falschen Namen 
Euxo 


us caudaus.” 

Amaranthus viridis auct., non L. 
Distribution and Ecology.—Native to and very widespread in the tropics, but 
also introduced into the warm temperate regions of North America and Eu- 
rope. The two wild subspecies are partially sympatric, but are separated eco- 
logically, subsp. emarginatus being more demanding of warm climates than 
subsp. blitum. The varieties of subsp. emarginatus are further ecologically sepa- 


982 BRIT.ORG/SIDA 19(4) 


UNIVERSITY OF 


EXAS 


HERBARIUM 


Hert l ity of California Ri id 


Ul 


Simarantbus blitum [. subsp ginatus (Moq 2 & Bray 
Carretera, Munos Garmendia & Pedrol 


A rn bij l * . 
Fic. 2? emarainatus var emaraqinga THe 
r J ~/ 


COSTEA ET Al 


Fig. 3.A I 


thus blitum 1 


t 


Det. by Mihai Costea 


The New York Botanical Garden 
Amaranthus lividus 


NEW YORK: Bronx 


U.S.A: Co., Bronx, grounds 
New York Botanical Garden. 


Decumbent annual herb, forming dense mats to 2 m 
diameter; stems smooth, slightly succulent, slightly 
plants predominantly green, wit reddish tinge 
part Male flowers very inconspicuous. Uri 


emarainatus var. nseudoaracilis 
J P 


984 BRIT.ORG/SIDA 19(4) 


rated in that variety emarginatus grows preferentially on sandy alluvium on 
the banks of rivers, on lakeshores and beaches, while variety pseudogracilis is 
found mostly asa weed in irrigated vegetable crops and nurseries. This subspe- 
cies is the most widespread form of A. blitum found in the United States. 


Representative specimens examined for var.emarginatus: FLORIDA. Dade Co.: Key Biscayne, Cape 
Florida State Park, 15 Mar 1969, Gillis 7688 (A). Homestead, Bay Front Park, 5 Nov 1977, Correll and 
Popenoe 49092 (A). Hillsborough Co.: 12 mi SE of Tampa (“beach area’), 28 Oct 1960, cae 0605 
(GH). Leon Co.: Tallahassee, Lake Ella, 10 Jul 1986, in 9737 (BRIT). Near F.S.U. campus, 6 Oct 
1994, Anderson 15283 (BRIT). 30 Oct 1976, Brumbach 9087 (GH). Manatee Co.: 20 mi E even 
27 Apr 1975, Genelle and Fleming 2040 (RSA, FLAS). Prete Parish Calcasieu: 4 mi S of Lake 
Charles, 13 Jul 1969, Thieret 31617 (SMU).Massachusetts. Hampshire Co.: University of Massachu- 
setts campus, Amherst, 17 Sep 1966, Ahles 64683 (SMU). NEW YORK. Bronx Co.: New York, 26 Sep 

1937, Moldenke 10534 (VDB). Madison Co.: shore of Oneida Lake, Lewis Point, 14 Sep 1938, House 
26133 (GH).NORTH CAROLINA. Duplin Co.: 2 miW of Greenevers (weed in broccoli) 15 Oct 1991, 
Wilbur 59784 (DUKE). PENNSYLVANIA. Bradford Co.: shore of Susquehanna River at Homets Ferry, 
3 Sep 1962, Wahi 20220 (GH). 14 Aug 1955, Wah! 16564 (GH). Travis Co.: at Hornsby Bend Sewage 
treatment Plant (Montopolis Quadrangle), 19 Apr 1986, Carr 7281 (TEX). 

Representative specimens examined for var. pseudogracilis: FLORIDA. Alachua Co.: Gaines 
ville, University of Florida Campus, 13 Oct 1967, D'Arcy 2197 (FW). Lee Co.: Eastern Sanibel, 20 Oct 
1976, Brumbach 9077 (BRIT).21 Nov 1972, Brumbach 81 tae: TEXAS. Orange Co.: Orange, 55 May 
1937,Cory 22306 (A). MISSOURI. Cape Girardeau C Girardeau weed in green house, Brooks 
/868 (VDB). PENNSYLVANIA. Philadelphia Co.: seen 2 Jul 1942, Long 58418 (GH). NEW 
YORK: grounds of New York Botanical Garden, 13 Oct 1984, Nee 30150 (TEX). 


1c. Amaranthus blitum subsp. oleraceus (L.) Costea, comb. nov. (Fig. 4). BASIONYM: 
Amaranthus oleraceus L., Sp. PL, ed. 2, 1403. 1763. Type: LINN 1117/3. 

Euxolus oleraceus (L.) Mog, DC, Prodr. 13(2):273. 1849. Albersia oleracea (L.) Boiss., Fl. Or. 4:99]. 

Albersi var. oleraceus(L.) Hooker fil., Fl. Brit. Ind. 4:721. 1885. pie ae A 

L. proles oleraceus(L.) Thell., Ascherson & Graebner, Syn. Mitteleur. Fl. 5:321.1914. Amaranthus 


ascendens Loisel. var. oleraceus(L.) Thell. ex Priszter, Agartud. Egyet. Kert- Szologazdasagtud. 
eoraus ig 2:22], 1953. Amaranthus lividus subsp. oleraceus (L.) S06, Acta Bot. Sci. Hung. 
10:376. 1 


Foner ie L., Sp. Pl. 990. 1753. Euxolus lividus (L.) Moq., DC. Prodr. 13(2):273. 1849. 
Amaranthus lividus L. proles lividus (typicus”) (L.) Thell., Ascherson & Graebner, Syn. 
Mitteleur. FL. 5:321. 1914. 

We present this taxon for comparison and because it isa potentially valuable veg- 
etable, though it appears not to have yet been used for this purpose in North America. 

It is very much like subsp. blitum but differs in the following characters: 
the plants are green to purple. The stem is erect, very thick, strongly ribbed. 
Cotyledons narrow, ovate to lanceolate, 15-18 x 5-6 mm, with rounded or trun- 
cate apex and acute base; petiole 10-12 mm long. Fruit 2.3-3.5 mm, when dry 
with the pericarp coarsely wrinkled. Stigmas branches well-developed (2 mm 
long and 0.5-0.6 mm wide at the base) and persistent when the fruit is dried. 
Seeds 1.2-1.6(-1.9) « 1.2-1.6(-1.9) mm, circular, not differentiated into central and 
marginal zones. The sculpture of the margin is inconspicuous at <100 x mag- 
nification. The cells of the exotesta in the marginal zone often inconspicuous 


COSTEA ET AL 


Amaranthus blitum L. subsp. oleraceus (L.) Costea (in ed.) 
Det. by Mihai Costea 


Fic. 4. Amaranthus blitum subsp. oleraceus. 


986 BRIT.ORG/SIDA 19(4) 


(the anticlinal walls are inconspicuous) with the periclinal walls less puncti- 
form sculptured than in subsp. blitum (Fig. 5 C). Margin of the seed rounded. 
Color dark-brown to blackish, uniform. Pollen as in subsp. blitum. 

Originated as a selection of A. blitum subsp. blitum and exists only as a 
cultivated vegetable. The only herbarium specimens of A. blitum subsp. 
oleraceus seen from USS. were those of the accessions PI 606281 and P1606282 
cultivated at Ames, lowa (originally from Bangladesh). Chan (1996), Chan and 
Sun (1997) refer to these accessions as “unidentified species—A MES 5366.” Based 
on molecular techniques the authors separate it both from A. blitumand A. tri- 
color. Unfortunately the accession PI 288277 used by the authors (Chan 1996; 
Chan & Sun 1997) as “A. lividus” (= A. blitum) and the others from India—Ames 
5123, 5146, 5315, 5387, PI 271465, PI 288277 and PI 608661—do not belong for 
sure to A. blitum. All these plants are related to A. tricolor as circumscribed by 
Aellen (1959). These results show that the complex A. tricolor is still poorly 
understood and a revision of the group is necessary. For future studies we rec- 
ommend the germplasm collection of the US. National Plant Germplasm Sys- 
tem h ttp://wwwa rs-grl N.gZOV. 


2. Amaranthus viridis L. Sp. Pl. ed. 2, 1405. 1763. Typr: “Habitat in Europa, Brasilia,” 
LINN 1117/15. 


Amaranthus gracilis Poiret, Lam., Encycl. Suppl. 1:312. 1810. 


Annual, with stem up to 1 m tall, ascending to erect, glabrous to sparsely pu- 
bescent above, the trichomes being multicellular and uniseriate. Cotyledons 
lanceolate, 12 x 2.5mm, with acute apex and acute to decurrent base: petiole 5- 
6 mm long. Leaves 3-8 x 2-5 cm, rhombic-ovate, entire to shallowly emarginate 
at apex. Inflorescence mostly terminal, long, thin, flexuous, ramified or not. 
Bracteoles about Imm long, acute, triangular-ovate, shorter than the tepals. 
Tepals 3, equal, 0.8-15 = 0.3-4 mm, bent along the fruit, narrow-spatulate to 
narrow-obovate, with the mid-vein green, enlarged above the middle. Fruit inde- 
hiscent, 1.2-1.5 mm, ellipsoidal to globose, usually gradually narrowed toward 
the stigma branches. Asa rule, the pericarp is strongly wrinkled, muricate, but 
sometimes almost smooth. Stigma branches thin (0.9-1.1 mm long and 0.05-0.075 
mm wide at the base). Seeds 1-1.2 x 0.9-1.Imm, circular with an inconspicuous 
hilum, differentiated into central and marginal zones. The margin of the seed 
is acute. The sculpture of the seeds is variable, two “types” being possible: 


1) Seeds entirely verrucose. 
2) Seeds with inconspicuous sculpture, resembling A. blitum subsp. oleraceus. 


On the basis of the verrucose seeds, Kowal (1954) circumscribed the Section 
Puncticulate that includes A. viridis and A. acutilobus. Because this character- 
istic varies within a species, it is evidently that the section Puncticulate created 
by Kowal can not be recognized. 


COSTEA ET AL 


Fic. 5. A-C.S test tk d gi A.A hus bli bs} blitum (scale b 50 1 ); B.A. blitum 
subsp | 25 um); C.A hus bli bsp. of (scale bar 50 Lim). D—E. Pollen (scale bar 
? 5 um) D.A blit bsp. blitum: E.A hus bli bsp. emarginatus F-G. Surface of exotesta on the 
seed margin 50| \-A h g bst graecizans; G | th 9 j bst sylvestris 


988 BRIT.ORG/SIDA 19(4) 


Pollen grains 18-20 um with 32-40 pores of 1.9-3.2 um diameter. Density 
of granules medium. Brenner observed (unpublished) that plants grown in the 
green house produce abundant nectar. 

The name A. viridis is sometimes misapplied to A. blitumin North America. 
For the nomenclature and typification of A. viridis see Thellung (1914), Merril 
(1936), Coons (1975, 1981). Even though related to A. blitum, A. viridis is un- 
questionable a distinct species. 

Distribution and ecology.—Native to South America according to the ma- 
jority of authors, but “possibly of Asian origin” (Townsend 1988). This is practi- 
cally a cosmopolitan noxious weed in the tropical and subtropical regions of 
the world. It has been introduced into the warmer temperate regions of Europe, 
North America, as Africa and Australia. 


R ined: MISSISSIPPI. Washington Co.: ee eland, 1 Sep 1990, 
Bryson 10315 IBE). SOUTH CAROLINA. Richland Co.: Columbia, 22 Aug 1971, Leonard et al. 4939 
A TEXAS. Arkansas Co.: Rockport, 5 Jun 1958, D.S.&H.B. Correll 1 ae (LL). Brazoria Co.: San 

rnard ae 11 Aug 1969, Fleetwood 9539 (TEX). Brazos Co.: College Station Texas, 29 Nov 1949, 
ane 91 (TEX). Calhoun Co.: Matagorda Isle, 19 Jul 1973, Hartman et Smith 3648 (TEX). Harris 
Co.: Houston, 10 Jun 1958, Traverse 747 (LL). Travis Co.: 30°16'43"N, 97°29'00"W, 140m, 16 Jun 1 996, 
Carr 15530 (TEX). Austin, 23 May 1986, Carr 7466 (TEX). Webb Co.: Laredo, 8 Apr 1965, Cuesta 22 (LL 


Ww 


3. Amaranthus graecizans L., Sp. Pl. 990.1753. Type: “Habitat in Virginia’, LINN H17/3. 


Annual with erect to ascendant stems, up to 80 cm, in the upper part with tri- 
chomes like A. blitoides[uniseriate trichomes consisting from 2(-3) cells with 
the terminal cells spherical, much larger than the basal one], in addition multi- 
cellular uniseriate trichomes are present. Leaves 2-4(-5) x 1-3 cm linear, lan- 
ceolate to rhombic—ovate or rhombic-elliptic, without a conspicuous hyaline 
margin. Bracteoles equal to or longer than the tepals with the mid-vein rami- 
fied. Tepals 3, L5-2 mm long, elliptic to lanceolate-oblong, gradually or abruptly 
narrowed into a mucro, with the mid-vein ramified. Fruit usually circumscis- 
sile (sometimes indehiscent), (2-)2.2-2.5(-2.7) mm, broad-ellipsoidal to globose, 
gradually narrowed toward the stigma region, with the dehiscence line in the 
lower part, strongly-wrinkled. Stigmas 2-2.25 mm long and 0.4-0.6 mm thick 
at the base. Seeds circular, 1.0-1.3 x 1.0-1.3 mm, lenticular, with a short furrow 
above the hilum, differentiated into central and marginal zones. Margin of the 
seed acute. Pollen grains 21-25 um with 19-32 pores of 1.5-1.7 um in diameter. 
Density of granules medium. Chromosome number: 2n=32. 

Amaranthus graecizans differs from A. albus in not having spinescent 
bracteoles and by its larger seeds. From A. blitoides it can be distinguished by 
the erect or ascendant stems, possession of only 3 tepals, the absence of a hya- 
line margin on the leaf blades and by its smaller seeds. 

Native in the Mediterranean region, including North Africa. The two wide- 
spread subspecies differ in leaf shape and seed sculpture. 


COSTEA ET Al 989 


— 


. Leaf blade lanceolate, usually 1-1.5 cm broad; marginal zone of the seeds with 
prominent cells whose periclinal walls are smooth (Fig.5F) A. graecizans 
subsp. graecizans 

. Leaf blade rhombic-ovate to ovate-elliptic, up to 4 cm broad; marginal zone of the 

seeds with inconspicuous cells, whose periclinal walls are minutely punctiform 
sculptured (Fig. 5G) . gracizans subsp. sylvestris 


“In 


3a. Amaranthus graecizans subsp. graecizans 
Amaranthus chan am., Encycl. 1:115. 1783., nom. illeg. 
3b. A g bsp. sylvestris (Vill.) Brenan, Watsonia 4:273. 1961. 
— 5) 


A thus angusti sLam. var. sylvestris (Vill.)Thell. in Schinz & Keller, Fl. Schweiz ed. 4, 
1:222. 1923. “Aanananlle sylvestris Vill. Cat. Pl. Jard. Strasbourg 111. 1807. Amarante. 
graecizans var. sylvestris (Vill.) Ascherson in Schweif., Beitr. Fl. Aethiop. 176. 186 


Distribution and ecology.—A. graecizans subsp. graecizans is found in the 
warmer regions of Europe, through most of Africa and in tropical Asia, espe- 
cially India. Plants belonging to subsp. sylvestris have generally the same range 
as subsp. graecizans but penetrate into the cooler regions of Europe, southwest 
Asia and northwest India. Another subspecies—subsp. thellungiacus (Nevski) 
Gusev, having leaf-blades narrowly linear or lanceolate to rhomboid spathu- 
late and long-aristate tepals and bracteoles, awns 0.3-1 mm, divergent, has a 
more restricted range: Middle Asia, India and also sporadically introduced in 
Africa (Townsend 1988). 

In the United States we have seen only 2 specimens belonging to subsp. 
sylvestris, both from NEW JERSEY. Camden Co. Camden, “on ballast,” 9 Aug 
1879, Parker s.n.(RSA); same location, 30 Aug 1879 Parker s.n.(RSA). It is almost 
certain that this plant is more widespread than these collections suggest, being 
probably under collected or misidentified as A. blitoides (especially with var. 
reverchonii) or A.albus. Also, there are numerous herbaria that we did not sur- 
vey for this species group as it was not a primary focus of our research. 


ACKNOWLEDGMENTS 


We are indepted to Gerold Hugin for his constant help along the years. Many 
thanks go to Julio Iranzo who made possible the SEM study, to Darleen DeMason 
and Edward Plumer for the photos. We also thank to David Brenner for the sup- 
ply of material and the useful discussions. The paper was written during first 
author’s Fulbright scholarship at University of California, Riverside. 


REFERENCES 
Axeroyo, J. 1993. Amaranthus L. In: T.G. Tutin, N.A. Burges, A.O. Chater, J.R. Edmondson, 
V.H.Heywood, D.M. Moore, D.H. Valentine, $.M.Walters, D.A. Webb, eds. Flora Europaea 
(second ed), Vol. 1, University Press, Cambridge. Pp. 130-132. 


990 BRIT.ORG/SIDA 19(4) 


g i Lo subsp. svt Noll B 
Det by Mihiat Costes 
Coll. C. F. PARKER : a4 1sr 3) 
| 


Fic.6.A th I sylvestris, Parker s.n. (RSA) 


COSTEA ET AL 991 


Actten, P. 1959. Amaranthus L. In: Hegi G., Illustrierte flora von Mitteleuropa (second ed.). 
Munchen. Pp. 3(2):465-516 

ALLEN, P. 1972. Amaranthaceae |n: K.H.Rechinger, ed. Flora Iranica. Pp. 1-19. 

BartHLott, W.and N. EHter. 1977. Raster-elektronen-mikroskopie der epidermis-oberflachen 
von Spermatophyten, Wiesbaden. 

Brenan, J.P.M.1961.Amaranthus L.in Britain. Watsonia 4:261-280. 

BreNAN, J.P.M.and C.C Townsenb. 1980. Proposal to reject Amaranthus blitum L.under art.69 
in favor of A. lividus L. Taxon 29:695-696. 

Brummit, R. 1984. Report of the committee for Spermatophyta 27. Taxon 33:297-301. 

Cuan, KF. 1996. Phylogenetic relationships and genetic diversity detected by rapid and 
isozyme analysis of crop and weedy species of Amaranthus. Ph.D. diss. Univ. Hong Kong, 
Hong Kong. 

CHAN, K.F. and M. Sun. 1997. Genetic diversity and relationships detected by isozyme and 
RAPD analysis of crop and wild species of Amaranthus. Theor. Appl. Genet. 95:865-873. 

Coons, M.P.1975.The genus Amaranthus in Ecuador. Ph.D. diss.Indiana Univ. Bloomington. 

Coons, M.P. 1981.Hybridization between Amaranthus viridis L and A. blitum L.Experientiae 
(Vicosa) 27:179-194. 

Coons, M.P. 1981. The status of Amaranthus viridis L. and A. blitum L. (Amaranthaceae) in 
South America. Experientiae (Vicosa) 27:159-178 

Costea, M. 1998a.Amaranthus L., Subgenus Albersia (Kunth) Gren.& Godr.in Romania., Rev. 
Rom. Biol. (Romanian Academy of Science) 43:95—-112 (in English). 

Coste, M. 1998b. Monograph of the genus Amaranthus L.in Romania. Ph.D. diss. Univer- 
sity of Bucharest, College of Biology, Bucharest (in Romanian). 

Eviasson, U. 1988. Floral morphology and taxonomic relation among genera of Amaran- 
thaceae in the New World and the Hawaiian Islands. J. Linn. Soc., Bot. 96:235-283. 

Fiuias, FR. Gauttiez and M. Gueves. 1980. Amaranthus blitum vs. A. lividus (Amaranthaceae). 
Taxon 29:149-150. 

HitcHcock, C.L.and Cronauist A. 1973. Amaranthus In: Flora of Pacific NorthWest; University 
of Washington Press, Seattle and London. Pp. 102. 

Hom, G.L., DL. Plucknett, JV. PANCHo, J.P Hersercer 1977. The world’s worst weeds. Distribu- 
tion and biology. East-West Center Book, Univ. Press of Hawaii, Honolulu, Hawaii. 

Hoim, G.L., J. Dott, E. Hotm and J. PancHo. 1997.World Weeds. Natural histories and distribu- 
tion. John Wiley & Sons, Inc. 

Hooker, J.D. 1885. Flora of British India vol.4, London. 

le © Vee: Die VelDrSInnng von AGO Arten in der sudlichen und mittleren 

gebieten. Phytocoenologia 14:289-379. 

HUGIN, G. 1987. einige bemerkungen zu wenig ewan Amaranthus -sippen 
(Amaranthaceae) Mitteleuropas. Willdenowia 16:453-478. 

Kartesz, J.T.and R. Kartesz. 1980.A synonymized checklist of the vascular flora of the United 
States, Canada and Greenland. Vol 2. Biota of North America; The University of North 
Carolina Press Chapel Hill. Pp. 28. 


992 BRIT.ORG/SIDA 19(4) 


Kartesz,J.1.1994,.A synonymized checklist of the vascular flora of the United States, Canada 
and Greenland. Timber Press Oregon. 1:35-37; 2:28-29. 

Kearny, T.H.and Peebles R. 1960.Amaranthus In: Arizona flora. University of California Press, 
Berkeley and Los Angeles. Pp. 265-268. 

Kowal, T. 1954. Cechy morfologiczne | anatomiczne nasion rodzaju Amaranthus L. oraz 
klucze do ich oznaczania. Monogr. Bot. (Warszawa) 170-193. 

McGrecor, R.L. 1986.Amaranthus |n:T.M. Barkley, ed.G.F.PA. Flora of Great Plains. University 
Press of Kansas. Pp. 180-184. 

Merritt, E.D. 1936. On the application of the binomial Amaranthus viridis L. Amer. J. Bot. 
23:609-612. 

Moauin-TAnoon, A. 1849.Amaranthaceae In:A.P.De Candolle.Prodromus sistematis naturalis 
regni vegetabilis. Paris. 13(2):23 1-424. 

Nowicke, J.W. 1993. Pollen morphology and exine ultrastructure in Caryophyllales In: H.D. 
Behnke & TJ. Mahbri, eds. Evolution and systematics in Caryophyllales. Springer Verlag. 
Pp: 165-221. 

Sauer, J.D. 1955. Revision of the dioecious amaranths. Madrono 13:5-46. 

Teitz, A. and et at. 1990. The dormancy of livid amaranth (Amaranthus lividus L.) seeds. 
Seed Sci. & Techn. 18:781-789. 

THeLLUNG, A. 1914. Amaranthus In: P. Ascherson & P. Graebner, eds. Synopsis der 
Mitteleuropaischen flora 5:225-356. 

Townsenb, C.C. 1988. Amaranthus In:E. Launert, ed. Flora Zambesiaca. London. 9(1):45-60, 

Utine, E.B. and W. Bray. 1884. A preliminary synopsis of the North American species of 
Amaranthus. Bot. Gaz. (Crawfordsville) 19:313-320. 

USDA, ARS. 2000. National genetic resources program. Germplasm resources informa- 
tion network-(GRIN). [Online Database] National Germplasm Resources Laboratory, 
Beltsville, Maryland. http.//www.ars-grin.gov. 


CAREX CUMBERLANDENSIS, A NEW SPECIES OF 
SECTION CAREYANAE (CYPERACEAE) FROM THE 
EASTERN UNITED STATES OF AMERICA 


Robert F.C. Naczi Robert Kral 
laude E. Phillips Herbarium Botanical Research Institute of Texas 
pune of sc ae and Natural Resources 09 Pecan Street 
Delaware State University Fort Worth, TX 76102-4060, U.S.A. 


Dover, Be 19901-2277, nv 
rnaczi@dsc.edu 


ue T. Bryson 
SDA-ARS 
Southern Ne ee oe Unit 


ner ie ee USA. 
cbryson@ars.usda.gov 


ABSTRACT 


C | Eonee:| 4 ee the 


from ic fi n Alabama, Arkansas, Georgia, Indi- 


ana, Kentucky, Mississippi, North Carolina, Ohio, ee South Carolina, Tennessee, Virginia, 
and West Virginia. Previously, specimens of C. cumber 


a 


andensis had been confused with C. digitalis 
and C. abscondita. Carex cumberlandensis has vegetative shoots much taller than the culms, very 
narrow staminate spikes, short staminate scales, and obtuse staminate scales, all of which distin- 


guish it from C. digitalis. Unlike C. abscondita, C. cumberlandensis has the lower spikes on long and 
drooping or nodding peduncles, distichously imbricate perigynia, the bract blade of the distalmost 
lateral ike (12-)17-51 times as long as wide, and always densely caespitose habit. The chromosome 
number of C. cumberlandensis is 2n = 23 0 


RESUMEN 


Se describe como nuevo Carex cumberlandensis de los bosques mesofilos de Alabama, Arkansas, 
Georgia, Indiana, Kentucky, Mississippi, Carolina on Norte, Ohio, Pennsylvania, Carolina del Sur, 


Tennessee, Virginia, y West Virginia. Previamente abian con uundide especimenes de G eu DRET 
g y rg 
la eaalale con ie aires yC. abscondita. C 1 


to) 
aT ‘ 7 b 


todo lo eal lo distingue de € divitalis. Al contrario que C. abscondita, C. cumberlandensis tiene elas 
espigas inferiores nutantes 0 colgantes en largos pedunculos, utriculos imbricados disticamente, la 
lamina de la bractea de la espiga lateral mas distal (12-)17-51 veces mas larga que ancha, y siempre 
un habito densamente cespitoso. El namero cromosomico de C. cumberlandensis es 2n = 23 11. 


INTRODUCTION 


Carex section Care yanae Tuck. ex Kuk. is a group of sedges endemic to eastern 
North America that is most diverse in mesic deciduous forests of the southeast- 
ern United States. Tuckerman (1843) first used the name “Careyanae” for this 


SIDA 19(4): 993 - 1014. 2001 


994 BRIT.ORG/SIDA 19(4) 


group, but without description or designation of rank. Kikenthal (1909) vali- 
dated the name at the rank of section, but circumscribed Carex sect. Care yanae 
to include species currently placed in Carex sect. Laxiflorae Kunth and Carex 
sect. Siderostictae Franchet ex Ohwi. Most authors since Kiikenthal (e.g. 
Mackenzie 1935; Fernald 1950; Bryson 1980; Manhart 1986; Gleason & Cronquist 
1991) have included sect. Careyanae ss. in sect. Laxiflorae. Bryson (1980) and 
Manhart (1986), based on morphological and biochemical evidence, respectively, 
found sect. Laxiflorae s.l. to consist of two subgroups that correspond to sect. 
Careyanae s.s.and sect. Laxiflorae ss. A phylogenetic analysis of sect. Laxiflorae 
and closely related sections revealed that sect. Careyanae ss.and sect. Laxiflorae 
s.s. are not each other's sister groups and are best treated as separate sections 
(Naczi 1989, 1992). In addition, molecular evidence supports the separation of 
sect. Careyanae and sect. Laxiflorae (Starr et al. 1999). Thus, various lines of 
evidence indicate sect. Care yanae is worthy of recognition. 

Morphologic apomorphies that diagnose sect. Care yanae include perigynia 
acutely triangular in cross-section and perigynia with numerous (usually 40 
or more), longitudinal sclerenchyma traces (“nerves” of earlier authors, e.g. 
Fernald 1950; Gleason & Cronquist 1991) that are impressed in living perigynia 
but raised in dried perigynia (Naczi, 1989, 1992, 1997). Additional character states 
shared by members of sect. Care yanae, though not unique to them, include gla- 
brous leaf and bract blades, long-sheathing proximal bracts, usually unisexual 
spikes with the terminal one staminate, and glabrous perigynia with entire 
apices. This section contains eight species and three nonautonymic varieties 
(with synonyms from recent floras and taxonomic works in brackets or paren- 
theses): C. abscondita Mack. [C. abscondita var. glauca (Chapm.) Fernald, C. 
abscondita var. rostellata Fernald, C. magnifolia Mack, C. austrocaroliniana L.H. 
Bailey, C. careyana Torr, C. digitalis Willd. var. asymmetrica Fernald, C. digi- 
talis var. digitalis, C. digitalis var. macropoda Fernald, C. laxiculmis Schwein. 
var. copulata (L.H. Bailey) Fernald[C. copulata (L.H. Bailey) Mack.], C. laxiculmis 
var. laxiculmis(xC. absconditiformis Fernald), C. plantaginea Lam.,C. platyphylla 
J. Carey, and one undescribed species (the subject of this paper) 

During the course of studies on sect. Careyanae, we noticed specimens in 
several herbaria labeled as C. abscondita or as C. digitalis that do not match the 
descriptions or types of these names. Field work and additional herbarium work 
revealed these specimens to represent an undescribed species here named C. 
cumberlandensis. In this paper, we describe this species and clarify its relation- 
ships with other taxa in sect. Care yanae. 


Carex eee Naczi, Kral, &@ Bryson, sp. nov. (Figs. 1, 2). Typr: USA. 
ALABAMA. WINSTON Co. ca. 10 mi N of Double Springs, Sipsey River Recreation Area ol 
7 i National Forest, E of river and S of bridge, T9S, R8W, NE1/4 of section 8, 4 May 
1991, Naczi 2868 (HOLOTYPE: an ISOTYPES: AUA, GH, MICH, MO, NCU, NY, US, USCH, 
’DB, WIN, ctb). 


NACZI ET AL., 995 


A Carex digitalis surculis vegetativis quam culmis 1.4-3.3plo altioribus, spicis terminalibus 0.6-1.3 
mm latis, squamis staminatis brevioribus, squamis staminatis obtusis differt; a Carex abscondita 


habitu semper dense caespitoso, laminis bractearum subtenentes spicas laterales summas (12-)17- 
51plo longioribus quam latioribus, pedunculis proximalibus plerumque cernuis vel nutantibus, 
pedunculis spicarum proximalium (28-)44-84(-91) mm longis, spicis perigyniis paucioribus 
instructis, perigyniis distichis differt. 

Perennial herb, densely caespitose. Rhizomes short, 0.4-2.2 mm long between 
shoots or branches of the rhizomes, 1.5-1.9 mm thick, covered with cataphylls 
1.3-1L8 mm long. Shoot bases surrounded by cataphylls, stramineous, lacking 
purplish coloration. Reproductive shoots erect or ascending; culms 7.3-27.4 cm 
tall, 0.4-0.7 mm wide at mid-height, smooth throughout, obtusely trigonous. 
Cataphylls glabrous, stramineous to whitish, multicostate. Leaves of reproduc- 
tive shoots with blades 0.4-52 cm long, 1.2-5.8(-6.6) mm wide, the widest (3.2-) 
3.7-5.8(-6.6) mm wide, green, flat to barely plicate, margins smooth or antrorsely 
scaberulous, adaxial surface smooth or sparsely antrorsely scaberulous on main 
veins, abaxial surface smooth; leaf sheaths 2.1-10.6 cm long, tight, glabrous; 
adaxial face of sheaths with hyaline and glabrous band, hyaline band with apex 
slightly concave to slightly convex; ligules lingulate with apex obtuse. Vegeta- 
tive shoots 14.6-47.0 cm tall, (1.4-)L7-3.3 times as tall as culms; pseudoculms 
3.4-7.8 cm tall, 0.10-0.18 of vegetative shoot height. Infructescence a single ter- 
minal spike and 2-3 lateral spikes, usually with the distal 2-3 spikes overlap- 
ping, rarely with all spikes separate; proximalmost bract with blade 7.1-25.2 
cm long and (0.5-)0.9-1.8 times as long as culm; bract blade of distalmost lat- 
eral spike 0.9-15.4 cm long, 1.3-3.5 mm wide, (12-)17-5] times as long as wide, 
and much exceeding terminal spike. Spikes 3-4 per culm, simple, single at nodes, 
erect; terminal spike 5.1-21.2 mm long, the longest (7.6-)9.6-21.2 mm long, 0.6- 
1.3mm wide, entirely staminate, 4-24-flowered, on erect and smooth peduncle 
2.1-44(-71) mm long, usually overlapping distalmost lateral spike and slightly 
exceeding it or slightly exceeded by it, rarely exceeding distalmost lateral spike 
and separate from it; lateral spikes 53-17 mm long, 2.8-4.1 mm wide, entirely 
pistillate, 4+-8(-9)-flowered, the perigynia distichously imbricate, the intern- 
ode between the proximalmost perigynia 1.8-3.0 mm long, on smooth peduncles, 
the proximal ones usually pendulous; the longest peduncle drooping or nod- 
ding, (28-)44-84(-91) mm long. Staminate scales (2.6-)2.8-3.5(-3.8) mm long, 
L1-L7 mm wide, narrowly elliptic to elliptic, obtuse, center green and I|-nerved, 
margins hyaline and whitish or whitish with ferruginous speckles and streaks. 
Pistillate scales 1.3-2.2 mm long, 1.4-2.1 mm wide, 0.41-0.55 times as long as 
perigynium, broadly ovate, obtuse, center green and |-nerved; margins entire, 
hyaline, whitish to tawny, infrequently with ferruginous speckles and short 
streaks. Anthers 3, 12-18 mm long. Styles jointed with summits of achenes, 
portions distal to achenes withering with age, portion proximal to achene (style 
base) persistent and becoming achene beak; base 0.1-0.2 mm wide. Stigmas 3, 


BRIT.ORG/SIDA 19(4) 


996 


egetative shoots (shorter 


h g pair of immature v 


a. Habit. b. Offshoot fi | 


a 
and distal shoot cataphyll (right). Scale bars equal 1 cm, with th let 


the holotype. 


NACZI ET AL., 


C 

Fig. 2. Ce a.Tip of leaf blade, abaxial surface. b. P i f leaf midblade, abaxial surface (left) and 
aanlalsiitat: aright). G ve j d, adaxial suriade d. Distal portion of cul 

Pair with 1 iwi lieiacl L £ hich all Ait H h hk hor 
(left), J Vf 2 \ I y ee ir EE “Be ay 4 I ; L me | £. f f \ | | £. fal 7 
g. Stamen. h. Pistillate scales, f d adaxial (lower). i Apex of lateral spike showing a 
perigyniunt wih subtenainig Beale and) distally, a ale SE an undeveloped ida a Perigynia, front 

ie t ght) k.A | (lef ars equa al2 

+h In th £ teh L £, 

VVERET the SRGIE LIE SGHIC TUE each illustration 


group G hi andi k. Drawn - Robert Kral cat the 
holotype. 


998 BRIT.ORG/SIDA 19(4) 


0.8-13 mm long, withering with age. Perigynia 2.9-3.8 mm long, 1.5-2.1 mm 
wide, 1.6-2.3 times as long as wide, 1.4-1.5 times as long as achene bodies, as- 
cending, acutely triangular in cross section, abaxial faces slightly convex to flat, 
adaxial face slightly concave to flat, nerves shallowly impressed in living 
perigynia and slightly raised in dry perigynia, nerves 43-54, glabrous, green to 
brown, dull, ellipsoid or obovoid, gradually tapered from widest point to trun- 
cate base, gradually or abruptly tapered to subacute or acute apex, beakless or 
with minute beak; beaks 0-0.2 mm long, 0-7% of perigynium length, bent 
abaxially, smooth, entire. Achenes 2.2-2.9 mm long, 1.4-L8 mm wide, tightly 
enveloped by perigynia, obovoid to ovoid, acutely trigonous, faces slightly con- 
cave to flat, brown, basally abruptly contracted to stipe, apically abruptly con- 
tracted to beak; stipe 0.1-0.2 mm long, vertical or slightly bent; body 1.9-2.7 
mm long, with widest point 0.9-1.5 mm from body apex; beak 0.1 mm long, 
bent abaxially 30-60 from vertical. 

Previously, herbarium specimens of Carex cumberlandensis have been iden- 
tified as C. digitalis and C. abscondita. However, C.cumberlandensis and C. digi- 
talisare only superficially similar. The most obvious similarity for them is their 
sharing of proximal spikes borne on long peduncles, a plesiomorphy in the sec- 
tion (Naczi, unpubl. phylogenetic analysis of sect. Careyanae). Thus, the simi- 
larity between C. cumberlandensis and C. digitalis is not indicative of a close 
relationship between these two species. Four features distinguish C. 
cumberlandensis and C. digitalis. Carex cumberlandensis has vegetative shoots 
much taller than the culms [tallest vegetative shoot per specimen/tallest culm 
per specimen = (1.4-)1.7-3.3], very narrow staminate spikes (0.6-1.3 mm wide), 
short staminate scales [2.6-3.5(-3.8) mm long], and obtuse staminate scales. 
Carex digitalis differs from C. cumberlandensis in having vegetative shoots 
shorter than to slightly exceeding the culms [tallest vegetative shoot per speci- 
men/tallest culm per specimen = 0.5-1.3(-18)], wider stami spikes [(1.0-)1.2- 
2.7 mm wide], longer staminate scales (3.6-5.5 mm long), and acute staminate 
scales. A scatter plot of measurements for two of the differences between C. 
cumberlandensis and C. digitalis, vegetative shoot height/culm height vs. width 
of staminate spike, provides complete separation of these two species (Fig. 3, 
with specimens measured for this plot indicated in the asterisked citation of 
the holotype of C. cumberlandensis, asterisked citations of representative speci- 
mens of C. cumberlandensis, asterisked specimens cited in Appendix A, and 
specimens cited in Appendix B). 

Carex cumberlandensis is most similar to C. abscondita, which is the spe- 
cies with which C. cumberlandensis is most often confused. These two species 
share vegetative shoots much taller than culms, very narrow staminate spikes, 
short staminate scales, and obtuse staminate scales, the same four features that 
distinguish C. cumberlandensis from C. digitalis (Fig. 3). Each of these features 
is a synapomorphy within sect. Careyanae and possessed by only C. 


tm 
—— 


NACZI ET Al 999 


5 T T 
= " C. abscondita 
cr) 7 OC. cumberlandensis 
es aC. digitalis 4 
& 
3 = 
O _ 
a (@) 
= 3 = : O 4 
cab) a9 O a 
ae se .. 
5 Oo 
8 af 00.8 : 
ep) e) 80. Phe) " 
oD) Oo 
2 °O 44 44,8 
© 4F rN Ad AA 7 
® ana 2% Aaa 
® A 
> 

0 | | 

0 1 2 3 


Staminate Spike, Width (mm) 


Fic. 3. Scatter plot o specimen vs. width of longest staminate 


spike per specimen a vGarey abscondita G cumberlandensis, and C. digitalis. 


cumberlandensis and C. abscondita. Thus, C.cumberlandensis and C. abscondita 
appear to be sister species (Naczi, unpubl. phylogenetic analysis of sect. 
Careyanae). 

Carex cumberlandensis differs from C. abscondita in several ways. First, C. 
cumberlandensis is always densely caespitose, whereas C. abscondita is often 
loosely caespitose. The growth form of C. abscondita appears to vary clinally, 
with plants from the southern portions of its range having a looser habit than 
those from the north. In fact, plants of C. abscondita from the northernmost 
portion of its range (northern Virginia and northward) are usually densely cae- 
spitose. Second, the proximal peduncles of C.cumberlandensis are much longer 
than those of C. abscondita [longest (per plant) peduncle of a proximal spike 
(28-)44-84(-91) mm long in C. cumberlandensis vs. (7.0-)15-42(-49) mm long 
in C. abscondital. Third, the peduncles of C. cumberlandensis usually droop or 


1000 BRIT.ORG/SIDA 19(4) 


nod (the proximal spikes are usually pendulous), but those of C. abscondita are 
usually erect or, at most, spreading (the spikes have their apices pointing up- 
ward). The distichously imbricate perigynia of C.cumberlandensis versus spi- 
rally imbricate ones of C. abscondita constitute a fourth difference between the 
two species. Fifth, C. cumberlandensis has fewer perigynia per spike than does 
C. abscondita [longest spike per plant with 4-8(-9) perigynia, including unde- 
veloped or aborted ones, in C. cumberlandensis vs. (6-)8-13 perigynia in C. 
abscondital. Finally, the value of the length/width ratio of the blade of the bract 
of the distalmost lateral spike is greater in C. cumberlandensis than it is in C. 
abscondita [12-)17-51 for C. cumberlandensis vs. 5.6-17(-26) for C. abscondital, 
Onascatter plot of greatest number of perigynia per spike vs. length of longest 
peduncle of lateral spike, specimen clusters for C. abscondita and C. 
cumberlandensis are well resolved (Fig. 4, with specimens measured for this 
plot the same as measured for Fig. 3). Although measurements overlap for each 
of these features, simultaneous consideration of them allows unambiguous 
identification of all specimens except for a few depauperate ones. Specimens 
that are somewhat difficult to identify based on these two features can be de- 
termined with certainty through consideration of additional features, such as 
habit (densely caespitose vs. loosely caespitose), arrangement of the perigynia 
(distichous vs. spiral), and length/width ratio of bract blades. 

Thus, Carex cumberlandensis is a species with several diagnostic features 
that distinguish it from morphologically similar taxa. Despite the slight over- 
lap of measurements of some key characters among C. cumberlandensis, C. 
abscondita, and C. digitalis, we have seen no specimens that simultaneously 
overlap in more than one or two key characters between C. cumberlandensis 
and C. abscondita or C. cumberlandensis and C. digitalis. 

The chromosome number of C. cumberlandensis is 2n = 23 Ul, determined 
from metaphase | of pollen mother cells (methods in Naczi 1999). Though the 
studied plants were from two populations, they yielded the same count [vouch- 
ers: Alabama, Winston Co., ca. 10 mi. N of Double Springs, Naczi 2966 (DOV, 
MICH); Georgia, Stephens Co., ca. 3 mi. N of Toccoa, Naczi 2965 (DOV, MICH)I. 
Carex abscondita (one plant studied) has a chromosome number of 2n = 24 II 
(Naczi, 1999). Because Carex species commonly exhibit intraspecific variabil- 
ity inchromosome number, it is premature to conclude the chromosome num- 
bers of C. cumberlandensis and C. abscondita do not overlap. 

Carex cumberlandensis inhabits mesic forests. The forests are deciduous or 
sometimes mixed with hemlock. These forests usually have diverse herbaceous 
floras. Frequent vascular plant associates of C. cumberlandensis include 
Anemone acutiloba (DC.) G. Lawson, Carex amphibola Steud., C. laxiflora Lam., 
C. rosea Willd., Circaea lutetiana (L.) Asch. & Magnus, Galium circaezans 
Michx., Hexastylis arifolia (Michx.) Small, Houstonia purpurea L., Lindera ben- 
zoin(L.) Blume, Podophyllum peltatum L., Polystichum acrostichoides (Michx.) 


NACZI ET Al 1001 


@ 15 / 
x 
for 
Y) - = 
© 
o |_| 
a i | || Hm 
= 10F = : 7 
x om 8n8 oO (ele) 
is ee 8 O ¢D) 00 
& » 0000 @® 
s = = ©O WMG@Dq@®d O 
D 5 |- @mo a 
ro! 
= Oo 
Z| 
= 
® =" C. abscondita 
© OC. cumberlandensis 
O 0 | ! | | 
0 20 40 60 80 100 


Longest Peduncle of Lateral Spike, Length (mm) 


Int nf | £ . ele ee pert Pantha fineliual J | d L 4 4 <~\ 


a 
Fic. 4, ICALLET PIOl Ul Yea leat MUTE Ul yp Uy t \ y } t yy Vo. 
length of | Juncle of | i d C. cumberlandensis 


Ln £ ia I Ait 


be J J 


Schott, Sanicula canadensis L., Scutellaria elliptica Muhl., and Tiarella ordifolia 
L. The substrates of C. cumberlandensis are slightly acidic to circumneutral 
loams, sandy loams, loamy sands, and clays (Naczi, unpubl. data). 

Carex cumberlandensis ranges from southwestern Pennsylvania south to 
Georgia and west to Arkansas (Fig. 5). The range of this species is entirely within 
unglaciated terrain, though C. cumberlandensis closely approaches the south- 
ern limits of glaciation (Flint 1971) in southern Ohio, northern Kentucky, and 
southern Indiana. A substantial disjunction appears to exist in the range of C. 
cumberlandensis. A population in Franklin County, Arkansas (Naczi 2155) is 
about 400 km (250 mi) from the nearest known population in the main por- 
tion of the species’ range (Naczi 4846 & Reznicek, Tennessee, Dyer County). 
The Arkansas population, in the Boston Mountains subdivision of the Ozark 
Mountain physiographic region (Robison & Allen 1995), is the only one known 


1002 BRIT.ORG/SIDA 19(4) 


e 
 ) 
. os is 500 km ; 
e.2% . 
ec, e 
e \ 
e C] e 
°° * 
e 
e 
* e 
*” 3" 
80°W 
Carex 
cumberlandensis 


west of the Mississippi River. Several other flowering plant species of forests 
and forest edges exhibit a similar disjunction between the Ozark Mountains 
and regions east of the Mississippi River, including Carex ouachitana Kral, 
Manhart, & Bryson (Naczi & Bryson 1990; McNeilus 1992), C. willdenowii Willd. 
(Naczi et al. 1998), Castanea ozarkensis Ashe Johnson 1988; Nixon 1997), Cotinus 
obovatus Raf. (Little 1977), Fothergilla major Lodd. (Smith 1988; Meyer 1997a), 
Leptopus phyllanthoides (Nutt.) G.L. Webster (Clark 1967; Smith 1988), Magno- 
lia tripetala L. (Little 1977, Meyer 1997b), Monarda virgata Raf. (Scora 1967: 
Smith 1988), Neviusia alabamensis A. Gray (Long 1989), and Trillium pusillum 
Michx. var. ozarkanum (Palmer & Steyerm.) Steyerm. (Cabe & Werth 1995). 
Carex cumberlandensis occurs in several physiographic provinces, but most 
of its range lies in the Appalachian Plateaus (including the Cumberland Pla- 
teau), Interior Low Plateaus, and Piedmont. It is most common in eastern Ken- 
tucky, central and eastern Tennessee, and northern Alabama. Carex 
cumberlandensis is rare and local on the Coastal Plain. For the Coastal Plain, 
only eight populations are known, from Alabama (Butler, Lowndes, Russell, and 
Wilcox counties), Georgia (Stewart County), Mississippi (Choctaw and Win- 
ston counties), and Tennessee (Dyer County). The specimens cited below are a 


NACZI ET Al 1003 


representative sample of about 280 records (about 810 total specimens) of C. 
cumberlandensis we have studied. Collections of C.cumberlandensis date from 
at least 1844 (Rugel 108, Tennessee, Sevier County). Long-standing confusion of 
C.cumberlandensis with C. abscondita and C. digitalis accounts for why the rela- 
tively well-collected, widespread, and common C. cumberlandensis had escaped 
recognition for so long. 

Because of past confusion of C. cumberlandensis with C. digitalis and C. 
abscondita, the distributions of the latter two species need clarification. The 
range of C. digitalis includes the entire region inhabited by C. cumberlandensis 
and is quite extensive: southern Nova Scotia, central Maine, and southern 
Ontario west to southern Wisconsin and south to central peninsular Florida 
and eastern Texas (Bryson 1980; Oldham & Zinck 1997). Bryson (1980) provides 
a map of the range of C. digitalis, including its varieties. Since C. 
cumberlandensis has been confused most often with C. abscondita, we provide 
a map of the range of C. abscondita (Fig. 6) and cite representative specimens of 
it (Appendix A). Carex abscondita occurs mostly east and south of the range of 
C.cumberlandensis: eastern Massachusetts, southwestern Kentucky, and south- 
eastern Missouri south to northern Florida and eastern Texas (Figs. 6, 7). Carex 
abscondita is most frequent on the Coastal Plain. Since C. abscondita is absent 
from Indiana, Ohio, and West Virginia, reports of this species from these states 
(e.g. Mackenzie 1935; Deam 1940; Fernald 1950; Manhart 1986; Cusick 1988; 
Gleason & Cronquist 1991; Ohio Division of Natural Areas and Preserves 2000) 
almost certainly refer to C. cumberlandensis. In addition, though C. abscondita 
occurs in Kentucky and Tennessee, many of the previous reports of this species 
from these states (e.g. Manhart 1986; Cusick 1989; Chester et al. 1993; McKinney 
et al. 2000) refer to C. cumberlandensis. 

Though C. abscondita is absent from most of the range of C. 
cumberlandensis, the two species are sympatric in southeastern and southern 
portions of the range of C. cumberlandensis (Fig. 7). Infrequently, plants of these 
two species do grow syntopically. We have observed C. cumberlandensis and C. 
abscondita growing together in Butler, Coosa, Tuscaloosa, Wilcox, and Winston 
counties, Alabama; Jasper and Monroe counties, Georgia; and Winston County, 
Mississippi. In localities of syntopy, C. abscondita tends to grow in moister soils 
than C. cumberlandensis. We have never observed hybrids or intermediates 
between C. cumberlandensis and C. abscondita. 

We have selected cumberlandensis as an epithet for this new species be- 
cause the Cumberland Plateau of eastern Kentucky, eastern Tennessee, and 
northern Alabama is the center of its distribution, the area in which it is most 
common, and the region in which it is a very characteristic element of the un- 
derstory of mesic forests. 


1004 BRIT.ORG/SIDA 19(4) 


500 km 
® 
YY 
e o vy 
2. yemas |S *o 
oe 
+ 29 ke: 
: & ie . 
3, ‘ 31 N 
80°W 
Carex 
abscondita 


Fic. 6. Geographic distribution of Carex abscondita. 


IDENTIFICATION KEY FOR MEMBERS OF CAREX SECTION CAREYANAE 


This key is designed for identification of complete and ample specimens bear- 
ing mature and healthy perigynia. Occasionally, insects induce galls in the 
perigynia of members of Carex section Careyanae. Because these galls distort 
the morphology of the perigynia, one should avoid considering affected 
perigynia when identifying these taxa. Relative to healthy perigynia, perigynia 
with galls are usually paler (especially basally), more elongate, more excurved, 
much less likely to contain mature achenes, and shed earlier. 

The terminal spike is entirely staminate. The lateral spikes are entirely pis- 
tillate, except in C. laxiculmis (with the basal scales of the lateral spikes sterile 
or subtending staminate flowers). In addition, plants of some of the taxa (espe- 
cially C. digitalis var. macropoda) occasionally have a few staminate flowers 
terminating the otherwise pistillate lateral spikes. 


NACZI ET AL., 


1005 


2 500 km 
Carex cumberlandensis 


Carex abscondita 


Fic. 7. Regi f 
g 


C abscondita 


Length of the peduncle for lateral spikes is measured from the base of the 
peduncle (at the node of its subtending bract) to its summit (at the base of the 


lowest scale in the spike surmounting the peduncle). Thus, peduncle length 
includes the portion of the peduncle included in the bract sheath 
. Shoot bases brownish, with purplish col 


loration cor ly absent 
2. Foliage green or glaucous; leaf blades siiccerine! shoots 1 - 3 5 cn as es 
as bract blades; eae a 9 4cm wide; bract 


proximal portions of culms 4.5-24 c 


3. Proximalmost scale of each lateral pk subtending a perigynium. 
4. Terminal spikes 0.6-1.4(-1 wide;staminate scales obtuse, those from 
middle region of terminal spike =765CmNe mm long; vegetative shoots 


much taller than culms, tallest vegetative shoot (1.4—-)1.7-3.7(-4.9) times 
as tall as tallest culm 


5. Perigynia distichously imbricate; longer lateral spikes with 4-8(-9 
perigynia; peduncles of proximal spikes usually drooping or nodding, 


) 


nt 


1006 BRIT.ORG/SIDA 19(4) 


the longest (per plant) pedunc 


le (28-)44-84( g;bra 
of distalmost lateral spike (12-)17-51 times as long as wide; dense 
caespitese arex cumberlandensis 
5 g Cae | eon ee | | | 7 +t 3 per j nia; 


eae iace: of proximal spikes usually aree the oe (per plant) 
peduncle (7.0-)15-42(-49) mm long; bract blade of distalmost lateral 
spike 5.6-17(-26) times as long as wide; loosely or densely caespitose 
Carex abscondita 
4. Terminal spikes (1.0-)1.2-2.7 mm wide; staminate scales acute, those from 
middle region of terminal spike 3.6—-5.5 mm long;vegetative shoots shorter 
than or slightly taller than culms, tallest vegetative shoot 0.5—1.3(-1.8) times 
as tall potallest culm. 


le | 


t blade ees lateral spike; 


“Jonge fer slain seduncle of terminal spike (6.3-)8.1-15.9 cm long; 
widest leaf blade 2.0-2.9(-3.5) mm wide; each perigynium face 7-10- 
nerve Carex digitalis var. macropoda 


6. Terminal spike usually surpassed by bract blade of distalmost lateral 
spike; longest (per plant) peduncle of terminal spike 0.9-7.2(-11.4) cm 
long; widest leaf blade 2.7-4.5(-5.3) mm wide; each perigynium face 
(8-)11-15-nerved. 


7. Perigynia 2.5-3.3 mm long, apex barely excurved Carex digitalis 
var. digitalis 
7. Perigynia 3.2-4.2 mm long, apex noticeably excurved Carex digitalis 


var. asymmetrica 
3. Proximalmost scale of each lateral spike sterile or subtending a staminate 


flower 
8. Foliage usually glaucous; longest (per plant) terminal spike (1.0-)1.2-2.5 
m long; widest leaf blade 6.4-11.8 mm wide Carex laxiculmis 


var. laxiculmis 

8. Foliage usually bright green; longest (per plant) terminal spike 0.6-2.0 
2.3) cm long; widest leaf blade 5.3-8.3 mm wide Carex laxiculmis 
pulata 


2. Foli | af blades of vegetative shoots 3.8-9.0 times as wide as bract 
piace widest in binde .1-2.5 cm wide; bract blades from middle and proxi- 
mal portions of culms 2.0-6.2 cm long Carex teens 


. Shoot bases purplish, with purplish coloration ranging from slight tinging of brow 


background in basal 1.5 cm of shoots to strong staining that obscures brown b ‘ 
ground and extends to 10 cm high. 
9. Widest leaf blade 1.0-2.5 cm wide; peduncles of lateral spikes usually erect or 
spreadin 
10. pace fom middle and proximal portions of culms bladeless or with blades 
0.1-1.9cm long; se 3.7-4.9 mm long; longest (per plant) lateral spike 
with 9-13 perigyni Carex plantaginea 
10. Bracts from Edd and proximal portions of culms with blades 2.1-9.2 cm 
— be 5,.0-6.6 mm long; longest (per plant) lateral spike with 4-9 
Carex careyana 
9. ie leaf blade 0.3-0.6 cm wide; peduncles of lateral spikes usually drooping 
arex austrocaroliniana 


NACZI ET Al 1007 


Representative specimens of Carex cumberlandensis. eae marked with an asterisk are those 
measured for scatter plots (Figs. 3, 4).U.S.A, ALABAMA. Butler Co.: ca.0.5 mi.N of Oaky Streak, 17 
May 1995, Naczi 4686 (DOV). Cleburne Co.: ca. 10 mi. ae Borden Springs, 23 Apr 1990, Kral 77370 
(VDB). Colbert Co.: ca.4 mi S of Allsboro, 27 May 1986, Bryson 4399 (MICH, ctb*). Coosa Co.: 1.8 mi S 
of Marble Valley, along N side of Paint Creek, 29 Apr 2000, Naczi 8526 & Ford (DOV™, MICH, ctb). 
Jackson Co.: 9 mi.S of Huntland, by Alabama 65, 28 Apr 1972, Kral 45814 (MO, VDB, ctb*). Jefferson 
Co.: just W of Tafford, 2 May 1980, Kral 64863 (VDB). Lawrence Co.: Bankhead National Forest, Th- 
ompson Creek, 1 Jun 1978, Bryson 2099 (MO,TENN, ctb*). Limestone Co.: NE of Athens, ca. 1 mi.W of 
I-65, Hays Mill Road, 19 May 1995, Kral 84718 (DOV, VDB). Lowndes Co.: 4.5 mi. E of Braggs, Alabama 
21,6 Apr 1993, Kral 81988 (DOV, VDB). Madison Co.: Monte Sano Mountain, 9 May 1981, Manhart 
265 & Bryson (MICH, TENN, ctb), 22 May 1983, Bryson 3529 (MICH, ctb),6 May 1991, Naczi 2876 (DOV*). 
shall Co.: ca. 1 mi.W of Grant, in Fletcher's Hollow, 19 Apr 1935, Harper 3348 (GH, MO, NY, PH*, 
US). Russell Co.: 2 mi. SW of Holy Trinity, 3 May 1996, Naczi 5209 (DOV, VDB, ctb). Talladega Co.: NW 
of Chandler Springs, off Alabama 77,8 May 1984, Kral 71344 (VDB). Tuscaloosa Co.: ca. 8 mi. ESE of 
Windham Springs, E side of Black Warrior River, 22 May 1996, Naczi 5459 (DOV, ctb). Walker Co.: ca. 
30 mi NW of Northport, 4 Jun 1971, Kral 42893 (VDB, ctb*); Sipsey, along W side of Sipsey Fork of 
Black Warrior River, 2 Jun 1997, Naczi 6442 (DOV). Wilcox Co.: ca.4.5 mi.SW of Camden, E of route 41, 
24 May 1993, Naczi 3096 (DOV). Winston Co.: ca.5 mi. W of Grayson, Bankhead National Forest, 
Sipsey River Recreation Area, 5 May 1978, Bryson 1892 (TENN, VDB, ctb). ARKANSAS. Franklin Co.: 
ca.3 mi.E of Cass, Redding Recreation Area, 13 May 1989, Naczi 2155 (DOV*, ctb). GEORGIA. Jasper 
Co.: 3.5 mi.E of Ocmulgee, 6 Apr 1949, Smith 4873 & Duncan (NA);ca. 14 mi.SW of Monticello, 19 May 
1997, Naczi 6054 (DOV, MICH, VDB, ctb). Monroe Co:: ca. 2.5 mi SW of Russellville, 25 Apr 2000, Naczi 
8398 & Ford (DOV*, VDB, ctb). Stephens Co.: ca. 3 mi.N of Toccoa, along Toccoa Creek, 3 May 1991 
Naczi 2855 (GA, DOV*, MICH). Stewart Co.: ca.5 mi. NE of Omaha, 22 May 1997, Naczi 6121 (DOV, ctb). 
Talbot Co.: ca.4 mi NE of Pleasant Hill, 25 Apr 2000, Naczi 8425 & Ford (DOV,VDB, ctb). Upson Co.: ca. 
3 mi SW of Thomaston, 25 Apr 2000, Naczi 8414 & Ford (DOV). Walker Co.: NW of junction of routes 
136 & 27, along route 136,9 May 1981, Manhart 243 (MICH, TENN, US, VDB); In the Pocket of Pigeon 
Mountain, 1 May 1982, Manhart 444 & Muzzy (DOV*, FLAS, TENN, US, ctb). INDIANA. Brown Co.: 
Brown County State Park, 16 Jun 1982, Manhart 515 & Muzzy (MICH, MO, TENN, US). Crawford Co.: 
ca.0.5 mi. N of English, 26 May 1911, Deam 8463 (NY); 1.5 mi.S of Grantsburg, slope of Little Blue 
River, 9 Jun 1919, Deam 27807 (GH); ca. 1 mi S of Wyandotte Cave, 13 May 1922, Deam 35617 (PH*). 
Harrison Co.: 2.5 mi. SE of Corydon, 26 May 1919, Deam 27673 (NY). Jennings Co.: ca. 0.75 mi. 
above Vernon, along N fork of Muscatatuck River, 23 Jun 1912, Deam 11305 (GH, NY). Lawrence Co.: 
ca. 2 mi. NE of Tunnelton, bluffs of Guthrie Creek, 17 May 1918, Deam 24852 (US). Orange Co.: ca. 1 
mi. SE of Orangeville, 1 Jun 1926, Deam 43219 (NY, PH*); 6 mi. E of Baden Springs, 16 Jun 1982, 
Manhart 518 & Muzzy (MO, US). Perry Co.: ca. 2.5 mi. SW of St. Croix, 29 Apr 1927, Deam 44130 (NY); 
4.9 mi.N of Bristow, along E side of route 145,27 May 1990, Naczi 2503 (DOV*, PH, US, VDB, ctb). Scott 
Co.: 1 mi. N of Lexington, 20 May 1919, Deam 27584 (NY). KENTUCKY. Adair Co.: ca.9 mi. S of Co- 
lumbia, along W side of route 704, 16 Jun 1998, Naczi 7387 & Reznicek LOW Barren Co.: ca. 3 mi. 
SSW of Haywood, Brigadoon State Nature Preserve, 14 Jun 1997, Naczi 6460 & Heeg (DOV, KNK, ctb). 
Bell Co.: ca.5 mi. NW of Pineville, along E side of route eee aed 1993, Naczi 3149 (DOV, MICH, VDB, 
ctb). Boyd Co.: S of Ashland, above Hurricane Fork Road, 24 May 1993, Cusick 30901 (MICH). a 
Co.: Central Kentucky Wildlife Refuge, 13 May 1992, Neca 1 & Gladu (KNK). Casey Co.: 2.9 
S of Ellisburg, 18 Jun 1998, Naczi 7451 & Reznicek (DOV). Clay Co.: Benge, S of route 472 and S of Mil 
Creek, 28 May 1993, Naczi 3135 & Reznicek (DOV, MICH, VDB, ctb). Clinton Co.: ca. 3.3 mi. WNW of 
Albany, 23 May 1998, Naczi 7186 & Ford (DOV). Cumberland Co.: 3.0 mi.SW of Amandaville, along W 
side of route 704,9 May 1991,Naczi 2893 (DOV*, MICH, US, VDB, ctb). Estill Co.: ca.8.5 mi SW of Irvine, 
12 May 1999, Naczi 7784 (DOV). Fleming Co.: ca.4 mi.E of Hillsboro, 28 May 1998, Naczi 7325 & Ford 
(DOV, WIN, ctb). Hardin Co.: 3.2 mi.W of Summit, Meeting Creek (Hoover Branch) Gorge, 26 May 


1008 BRIT.ORG/SIDA 19(4) 


1979, Cranfill 4336 (MICH). Harrison Co.: ca. 1.5 mi. NE of Sunrise, Quiet Trails State Nature Preserve, 
21 Jun 1997, Naczi 6628 (DOV, KNK, ctb). Hart Co.: ca. 2.3 mi. NNE of Canmer, 17 Jun 1998, Naczi 7425 
& Reznicek (DOV, MICH, ctb). Jackson Co.: near South Tree Recreation Area, 16 Jun 1982, Manhart 
503 & Muzzy (MO). Knott Co.: SE of Cody, 20 May 1994, McKinney 61 et FENN). Knox Co.: ca. 3 mi 
NNE of Barbourville, 29 May 1993, Naczi 3143 (DOV). Laurel Co.: ca.5.5 mi. WNW of See io lia 
1997, Naczi 6521 & Heeg (DOV, ctb). Lawrence Co.: NE a Fallsburg, ae route 707,24 May 1993, 
Cusick 30922 (CM, MICH). Leslie Co.: along highway 421,W from Hyden, ca. 7 mi. E of Clay County 
line, 21 May 1992, McKinney 5035 (EKY, KNK). Lewis Be Laurel Fork, 19 May 1988, Cusick 27393 (NY, 
ee stele se Barthell SW agian: Rock Creek (west), 19 May 1989, Campbell s.n. (KY). 
rtin Co. of Add, along Pack Branch above Martin County Lake, 1 Jun 1993, Cusick 30967 
eo er ae ca.8 mi. SE of Tompkinsville, 23 May 1998, Naczi 7208 & Ford (DOV*, WIN). Ohio 
Co.: L.B. Davidson Wildlife Management Area, 4 May 1992, McKinney 4953 (EKY, KNK). Pendleton 
Co.: 2 road mi. ESE of eastern edge of Falmouth, 3 Jun 1994, Naczi 4053 & Thieret (DOV, MICH, PH, VDB, 
ctb). Powell Co.: ca. 2 mi. SSE of Slade, Natural Bridge State Nature Preserve, 22 Jun 1997, Naezk 6663 
(DOV, KNkK). Pulaski Co.: General Burnside State Park, 16 Jun 1982, Manhart 510 & Muzzy (MICH, 
TENN). Rockcastle Co.: 2.5 mi.N of junction of routes 490 and 89, 16 Jun 1982, Manhart 509 & Muzzy 
(MICH, TENN). Rowan Co.: N of Triplett, Burnt House Branch, 15 May 1987, Cusick 26398 (VDB). Rus- 
sell Co.: SW end of Cave Springs Road (between forks), 20 Jun 1993, Campbell s.n. (KY). Warren Co.: 
ca. 11.5 mi. NNE of Bowling Green, along S side of Clay Lick Creek, 15 Jun 1997, Naczi 6500 & Heeg 
(DOV). Wayne Co.: 1.75 mi. N of Kentucky-Tennessee line, along route 167,15 Jun 1982, Manhart 
499 & Muzzy (MICH, TENN). Whitley Co.: ca.8 mi.E eae 29 May 1993, Naczi 3150 (DOV, 
MICH, US, VDB, ctb). MISSISSIPPI. sone Co.: ca.4 mi. NNE of Ackerman, near Horse Branch, 4 
81,McDaniel 25006 (ctb*). Tishomingo aig Creek Quad., Burns Hollow, 15 May 1979, 
Rogers 46684 (TENN). Winston Co.: 3.5 mi. NW of Louisville, Tombigbee National Forest, 26 Apr 1977, 
Byson 1523 (ctb*);ca.6.3 mi. NNW of Louisville, 29 May 1997, Naczi 6277 et al. (DOV). NORTH CARO- 
LINA. Alexander Co.: 5 mi. £ of Vashti, 13 May 1956, Radford 11194 (NCU). Anson Co.: 8 mi. S of 
Lilesville, near Pee Dee River, 20 May 1961, Radford 43609 (NCU, US). Lee Co.: 2 mi.W of Moncure, 
near Deep River, 3 Jun 1960, Radford 43209 (NCU, TENN). Madison Co.: ca. 5 mi. W of Hot Springs, 
along Mine Hollow Branch, 22 May 1971, Leonard 4788 (MICH, NCU, NY, VDB, WVA). Montgomery 
o.: Uwharrie National Forest, along W fork of McLean Creek, Park's Place Road, 18 May 1970, Le- 
onard 3201 & Radford (MO, NCU). Orange Co.: Chapel Hill, [no date], Ashe 2062 (OS, PH, US). Polk Co.: 
R.R. Melrose, 19 May 1899, Churchill s.n. (GH). Stanly Co.: NE of Richfield, along Yadkin River, 6 May 
1951, Godfrey 51129 & Fox ae Yadkin Co.: near Yadkin River a route 67, 30 May 1 isa 
Manhart 323 & Smith (MICH).OHIO: Adams Co.: 3/5 mi.S of Lynx,6 May 1991, Reznicek 8 bervavans ek 
(MICH). Gallia Co.: ca. 10 mi. se of Gallipolis, a of upper part of ae Run, 25 May 1 
Reznicek 8170 & Reznicek (MICH). Scioto Co.: Nile Twp., W side of Rock Lick, 20 May 1985, Carr a 
(MICH). Vinton Co.: Knox Twp.,sect.31,13 Ma ee as PENNSYLVANIA. Blair Co.: 
E of Altoona, Brush Mountain, 15 Jun 1929, Bright 1905 (CM), Bright 1906 (CM), Bright 1908 (CM). 
SOUTH EAROONE ar Co.: Sumter National Forest, E- ae slopes at Johns Creek, 21 Jun 
1992, Hill 2 Ison (GH, ancaster Co.: 13.7 mi. NNW of Lancaster, overlooking Catawba 
River, 2] Apr 1958 Bunton arene ).McCormick pee km W of Edgefield County line, 
Sumter National Forest, NW of junction of S-204 and SC 28, 18 Apr 1996, Nelson 17142 (MICH, USCH). 
Oconee Co.: ca. 4 mi.W of Walhalla, 3 May 1990, Naczi 2358 (DOV*). Pickens Co.: ca. 5 mi.S of Rocky 
Bottom, 3 May 1989, Naczi 2092 (DOV™, ctb). Union Co.: ca. 9 mi. WSW of Union, 8 May 1993, Horn 
6588 (MICH). TENNESSEE. Anderson Co.: Norris Dam State Park, near Clear Creek, 12 May 1987, 
McNeilus 87-794 (MICH, TENN). Bledsoe Co.: NW of Pikeville, 16 May 1934, Harger 7827 (GH, TENN). 
Blount Co.: Great Smoky Mountains, White Oak Sink area, 12 May 1980, Bryson 2945 & Bryson (ctb*). 
Campbell Co.: 1 mi.S of Morley on route 25W, 13 May 1980, Bryson 2956 & Bryson (ctb*). Cannon 
Co.: N side of North Short Mountain Road, 3 May 1989, Orzell & Bridges 9438 (MICH*). Cheatham Co.: 
Pegram, 16 Jul 1939, Svenson 10346 (GH). Clay Co.: Along Turkey Creek, 29 Apr 1991, McNeilus 91-190 


NACZI ET Al 1009 


(TENN). Coffee Co.: W outskirts of Manchester in Oakwood subdivision, 6 May 1989, Kral 76195 
(VDB).Cumberland Co.: 3.0 mi.on Hinch Mountain Road S of its junction with old route 127,8 May 
1986, Wofford 86-43 & Ma (ctb*). Davidson Co.: ca.5 mi.NE of junction of Tennessee 12 and Marrow- 
bone Creek Road, 16 May 1971, Kral 42612 (MO, VDB, ctb). DeKalb Co.: E of Se ae he upper 
King Creek by road to Fall Creek Falls, 3 May 1986, Kral 73349 (VDB). Dickson Co.: 2.8 of White 
Bluff by route 70, 13 May 1975, Kral 55379 (MICH). Dyer Co.: ca.6 mi. NW of saa N of route 
182,27 Jun Hee ee spre neeei (DOV, MICH). Franklin Co.: Sewanee, Lost Creek Cove, 24 Jun 
1938, Svenson 9689 (GH, VDB). Grundy Co.: Fiery Gizzard Gorge, along Big Gizzard Creek, 15 May 
1965, Clark 1771 (NCU*). Hardin Co.: 6.8 mi. SE of Savannah on Tennessee 69, 2 May 1970, Kral 38916 
(VDB).Hickman Co.: 4—4.5 mi.W of Centerville, by Tennessee 50,4 May 1993, Kral 82267 (DOV, TENN, 

VDB). Humphreys Co.: ca.5 mi. NNW of Bucksnort, by Tennessee 230, 19 May 1989, Kral 76304 (VDB). 
Jackson Co.: NW of Gainesboro, above Cumberland River, 14 May 1992, Kral 80515 & Rust (DOV, 
VDB). Lawrence Co.: ca.8 mi. WSW of Lawrenceburg, Knob Creek, by Tennessee 242, 5 May 1990, 
Kral 77462 en Lewis Co.: Meriwether Lewis Park, 17 May 1971, Kral 42648 (MO,VDB).Loudon Co.: 
ca. 1.5 mi. E of Centerville Store, along Big Hill Road, 4 May 1993, McNeilus 93-406 (MICH, TENN). 
Macon Co.: S of and below Lafayette by route 10,5 May 1975, Kral 55211 (MICH).Maury Co.: along 
Big Bigby Creek at E side of Sandy Hook, 19 May 1993, Kral 82294 (DOV, VDB). Monroe Co.: SE of 
Pumpkin Center, between Chilhowee Lake and junction of routes 129 and 72,15 May 1980, Bryson 
2987 & Bryson (ctb*). Moore Co.: ca. 3 mi.W of Motlow State Campus and SW of Tullahoma, 26 Apr 
1989, Kral 76096 (VDB).Morgan Co.: Rugby, 18 Jun 1938, Svenson 9331 (GH,NY).Overton Co.: 3.4 mi. 
E of Alpine, 31 Apr 1971, Kral 42328 (VDB, ctb). Perry Co.: SE of Linden on old SR 13, 0n E side of 
Buffalo River, 4 May 1993, Kral 82278 (DOV, TENN, VDB). Putnam Co.: ca.8 mi. SE of Cookville, 10 May 
1988, McKinney 2843-3 (ctb*). Scott Co.: Along Low Gap Road, 24 May 1994, McNeilus 94-304 (TENN). 
Sequatchie Co.: NW of Dunlap, along Little Brush Creek, 29 May 1990, Naczi 2512 & Kral (DOV*, 
MICH, VDB). [Sevier Co.]: between Sevierville and the French Broad River, May 1844, Rugel 108 (MICH, 
NA, VDB). Stewart Co.: Land Between the Lakes, along Bee Creek, tributary of Bear Creek, 14 May 
1982, Patrick 4256 & DeSelm (VDB). Sumner Co.: 2.5 mi. S of Westmoreland, Taylor Hollow, 17 May 
1979, Carter 1816 (ctb). Union Co.: ca. 7 mi.W of Maynardville, Big Ridge State Park, 30 May 1995, 
Naczi 4747 & Naczi (DOV, MICH, VDB, ctb). Van Buren Co.: 6-7 mi. E of Spencer, by Tennessee 30, 1 
May 1971, Kral 42364 (VDB).Warren Co.: Rock Island, Rock Island State Park, 27 May 1993, Naczi 3115 
& Reznicek (DOV, MICH, VDB, ctb). Wayne Co.: ca.5 mi.W of Iron City, just E of Cypress Creek bridge, 5 
May 1990, Kral 77494 (VDB).White Co.: ca. 3.9 mi.E of Sparta, along highway 70,12 May 1986,McKin- 
ney 2135 (VDB). VIRGINIA. Buchanan Co.: NE of Slate, 20 May 1993, Wieboldt 8545 (DOV). Franklin 
Co.:ca. 10.5 km SWof Ferrum, 11 May 1995, Wieboldt 9202 (DOV). Rockbridge Co.: Natural Bridge, 28 
May 1909, Bartram s.n. (PH*). Scott Co.: ca.6 mi. NE of Gate City, W of route 669 crossing of Copper 
Creek, 21 Jun 1993, Naczi 3171 et al. (DOV, MICH). Wise Co.: ca. 1.75 mi. SW of Norton, Powell Moun- 
tain, 2 Jun 1988, Wieboldt 6675 (MICH). WEST VIRGINIA. Nicholas Co.: U.S. highway 41, above 
Meadow River, 28 May 1987, Brant 1116 & O'Donnell (MO). Ritchie Co.: ca. 2 mi. NW of Ellenboro, 8 
May 1985, Cusick 24199 & Ortt (MICH, NCU). Wayne Co.: mouth of Big Creek, 6 May 1938, McFarland 
4355 & Plymale (NA). 


APPENDIX A 


Representative Specimens of Carex abscondita. Specimens marked with an asterisk are those 
measured for scatter plots (Figs. 3,4).U.S.A. ALABAMA. Autauga Co.: ca.5.5 mi ENE of Autaugaville, 
28 May 1997, Naczi 6274 (DOV*). Baldwin Co.: 1 mi. SE of Spanish Fort, 24 Apr 1964, McDaniel 4257 
(BE). Barbour Co.: Blue Springs, Choctawhatchee River, 7 Jun 1976, Kral 58187 (DOV,VDB). Bibb Co.: 
near Lightsey’s Mill, Cahaba River bottoms,6 Jun 1935, Harper 3395 (GH,NY);ca.6 mi SE of Centreville, 
5 May 1987, Naczi 1527 (DOV*). Butler Co.: ca.0.5 mi. N of Oaky Streak, 17 May 1995, Naczi 4689 
(DOV). Coosa Co.: 1.8 mi S of Marble Valley, along N side of Paint Creek, 29 Apr 2000, Naczi 8529 & 
Ford (DOV*). Dale Co.: Just NW of Ozark, 24 May 1975, Kral 55862 (TENN). Dallas Co.: ca.9 mi. SE of 


1010 BRIT.ORG/SIDA 19(4) 


Selma, 16 May 1995, Naczi 4653 (DOV).Geneva Co.: 5 mi. NW of Hartford, bottoms of Choctawhatchee 
River, 18 May 1966, McDaniel 7552 (IBE, VDB). Jackson Co.: 1 mi.W of Scottsboro, 3 May 1939, Svenson 
10193 (GH, MO, NY, PH, TENN). Lee Co.: Auburn, 10 May 1901, Earle s.n. (NY). Madison Co.: Ardmore, 
Limestone Creek, 19 May 1995, Kral 84677 (MICH, VDB). Marion Co.: ca.3 mi.N of Guin, 25 Apr 1941, 
Harper 3845 (GH, NY). Mobile Co.: Mobile, 23 Apr 1968, LeLong 4386 (USAM). Monroe Co.: 1.8 mi N 
of routes 47-83 junction in Midway, 28 Apr 1991, Naczi 2819 (DOV*). Morgan Co.: 1.6 mi.N of Florette, 
along Cotaco Creek, 9 May 1986, Naczi 1030A (DOV*). Tuscaloosa Co.: ca. 8 mi. ESE of Windham 

prings, E side of Black Warrior River, 22 May 1996, Naczi 5454 (DOV). Wilcox Co.: ca. 4.5 mi. SW of 
Camden, of route 41,24 May 1993, Naczi 3095 (DOV). Winston Co.: ca. 10 mi.N of Double Springs, 
Sipsey River Recreation Area of Bankhead National Forest, 4 May 1991, Naczi 2870 (DOV*, MICH). 
ARKANSAS. Cross Co.: W of Birdeye, 20 May 1990, Naczi 2448 (DOV*). Howard Co.: ca. 8 mi. W of 
Umpire, 17 May 1988, Naczi 1924 (MICH, ctb). Lafayette Co.: Bodcow Bayou, 6 May 1995, Hyatt 6366 
& Hyatt (DOV*, MICH). Montgomery Co.: 1.2 mi. E of Pine Ridge, 11 May 1989, Naczi 2142 (MICH, 
UARK). Phillips Co.: 1.5 mi.N of West Helena, 15 May 1987, Naczi 1598 (MICH, UARK, ctb). Pope Co.: 
N of Dover, 20 May 1980, Davis 2231 (VDB). Scott Co.: ca. 2 mi.N of Y City, 10 May 1989, Naczi 2131 
(MICH, UARK), 20 May 1994, Naczi 3937 & ae '). Sevier Co.: 8 mi.E of DeQueen, 10 May 1989, 
Naczi 2126 (MICH, UARK, ctb). CONNECTICUT. [New Haven County]: nee 2/7 Jun 1901, Eames 
$.n. NY), [New London County]: ie oe 13 Jun 1896, Graves s.n. (GH). DELAWARE. New Castle 
Co.: ca. 3 mi. NW of Glasgow, 4 Jun 1986, Naczi 1313 (DOV); SW of McKays Corner, 7 Jun 1988, Naczi 
2008 (DOV*); 0.4 mi S of Townsend, 31 May 1999, Naczi 8089 (DOV*). Kem Co.: ca. 2.5 mi SSW of 


ee 5 May 2001, Naczi 8710 (DOV). Sussex Co.: between nd Millsboro, 21 Jun 
rue 200a are ISTRICT OF COLUMBIA. Washington, 9 Jun 1 901, Aare oat FLORIDA. 
ce Co.: 15 mi. N of Wewahitchka, 18 May 1965, McDaniel 6234 (IBE, MO, NY, VDB). Gadsden 


Co.: S side ie Apalachicola River, 18 Mar 1975, Kral 57343 (DOV, MICH, TENN, VDB); 
Chattahoochee, 27 May 1997, Naczi 6223 (DOV*). Jackson Co.: Floodplain of ea aoe River 
between U.S.90 and Jim Woodruff Dam, 14 Apr 1982, Manhart 425 & Gholson (DOV*); ¢ 
ae along W side of Chipola River,6 May 1989, Naczi 2109 (DOV). Santa Rosa Co.: an ay, 27 
Mar 1956, Godfrey 54609 (GH). GEORGIA. Burke Co.: 7 mi. NE of Shell Bluff, near Savannah River, 18 
May 1989, Naczi 2190 (DOV*). Chattahoochee Co.: ca.9 mi W of Cusseta, 23 May 1997, Naczi 6133 
(DOV*). Early Co.: ca. 1.5 mi. NW of Hilton, along Chattahoochee River, 18 May 1995, Naczi ees 
(DOV, MICH). acim Co.: ca. 2.5 MIN of Clya, 2 May 1991, Naczi 2832 (DOV*). Hancock Co.: ca. 1 
mi.E of Sparta, 20 May 1996, Naczi 5385 (DOV). Harris Co.: ca.6 mi.W of Mulberry Grove, along S een 
of Mulberry Creek, 24 May 1997, Naczi 6151 (DOV*). oe Co.: ca. 14 mi. SW of Monticello, 19 a 
1997, Naczi 6055 (DOV). Monroe Co.: just S of Dames Ferry, W side of ae River, 19 May 19 
Nacz! 6059 (DOV); ca. 2.5 mi SW of Russellville, 25 Apr 2000, Naczi 8400 & Ford (DOV). Screven 
Blue Springs, 2 May 1991, Naczi 2839 (DOV*). KENTUCKY. Callo euance 0.: os ong Dalton Road off KY 
280, 22 May 1990, McKinney 4198 & Hamilton (EKY, VDB). LOUISIANA. Grant Parish: 1.0 mi. S of 
Packton on US 167, 19 May 1995, Hyatt 6434 (MICH). Iberia Parish: Weeks Island, 15 Apr 1983, Wof 
ford 83-6 & Murrell (TENN). Washington Parish: ca. 2 mi E of Franklinton, 13 May 1972, Rogers 8067 
(TENN). West Feliciana Parish: ca. 4 mi. NW of Saint Francisville, 2 May 1988, Naczi 1820 (DOV*). 
MARYLAND. Anne Arundel Co.: Patuxent Research Refuge, | Jun 1945, Hermann 11437 (GH).Prince 
Georges Co.: 1.5 mi. SE of Suitland, 1 Jun 1939, Hermann 10331 (PH). Somerset Co.: 2 mi. N of 
ty eee May 1938, moe 3/98 (GH, PH). Talbot Co.: 4.25 mi. WNW of Easton, 5 Jul 1941, 
Earle 3414 (PH). Worcester Co.: 1.6 mi. SW of Public Landing, along Tanhouse Creek, 7 Jun 
Naczi oe (DOV*). ee Bristol Co.: Dighton, 24 Jun 1957, Seymour 17122 (GH). [Dukes 
County]: Martha's Vineyard, between Town Hall and Menemsha, Chilmark, 19 Jun 1917, a 
1882 (GH). Norfolk Co.: Dedham, 23 Jun 1 . Faxon s.n. (US). MISSISSIPPI. Bolivar Co.: ca iS 
of Choctaw, 23 Apr 1992, Bryson 11358 & Newton (DOV, ctb). Carroll Co.: N of Vaiden, ee a tb 
tary of Hurricane Creek, 5 Jun ap Bryson 15309 (DOV, MICH, ctb). Forrest Co.: ca. 10 m 
Hattiesburg, 30 Apr 1988, Naczi 1818 (DOV). Holmes Co.: 6.2 mi.E of Thornton, 14 Apr 1992, ae 


— 


NACZI ET AL 1011 


11224 & Newton (DOV, ctb). Lafayette Co.: ca.5 mi. ENE of Oxford, 8 Jun 1996, Bryson 15324 (DOV, 
ctb). Lee Co.: W of Tupelo, 17 Apr 1992, Bryson 11328 (DOV, ctb). Lowndes Co.: ca. 10 mi. of Colum- 
bus, 11 Apr 1987, Bryson 5336 (DOV, MICH, ctb). Oktibbeha Co.: ca. 15 mi. S of Starkville, Noxubee 
Wildlife Refuge, 26 May 1980, Bryson 3053 & Mathies (ctb). Pearl River Co.: just S of Carriere, 10 Apr 
1936, Harper 3477 (NY); ca. 3.2 mi S of Marion-Pearl River Co. line, 26 Apr 1991, Bryson 10750 et al. 
(DOV*, ctb). Tallahatchie Co.: ca. 8 mi. N of Charleston, 21 Apr 1988, Bryson 7345 & Morris (DOV, 
MICH, ctb). Tishomingo Co.: ca. 1.2 mi.S of Mingo, along Cedar Creek, 17 May 1992, Bryson 11707 & 
Warren (DOV, ctb). Union Co.: 6 mi. SE of New Albany, E branch of Okannatie Creek, 18 May 1990, 
sti 9954 (DOV*, MICH, ctb). Wilkinson Co.: 1.2 mi.W of Lessley, 11 May 1990, Naczi 2392 (DOV). 
nston Co.: ca.6.3 mi. NNW of Louisville, 29 May 1997, Naczi 6283 et al. (DOV). Yalobusha Co.: ca.6 
mi ie of Oakland, 9 Apr 1991, Bryson 10621 (DOV*, ctb). MISSOURI. Dunklin Co.: ca. 7 mi. NW of 
Campbell, along old channel of Saint Francis River, 22 May 1993, Reznicek 9431 et al. (DOV™, MICH). 
Ripley Co.: SW of Neelyville, 24 May 1989, Yatskievych 89-131 et al. (MO).NEW JERSEY. Bergen Co.: 
Paramus, 25 Jun 1935, Svenson 6655 (GH). Cumberland Co.: ca. 2 mi. N of Newport, 21 Jun 1994, 
Naczi eee Thieret (DOV*, MICH). Mercer Co.: Imlaystown, 30 Jun 1926, Long 32985 (GH). Salem 
Co.: 1 mi.S of Pittsgrove, 26 May 1923, ae isbach 1329 (CM).NEW YORK. Nassau Co.: Glen Cove, 15 
Jun ae Bicknell 2046 (PH). oer CAROLINA. Columbus Co.: ca. 2 mi. NE of Whiteville, 21 May 
1992, Reznicek 8967 & Reznicek (DOV, an Craven Co.: 1 mi. NW of Vanceboro, 13 May 1950, 
Blomquist 14961 & Batson (NY).Pitt Co.: 2 mi.E of Winterville, 14 Jun 1958, Radford 34985 (GH).OKLA- 
HOMA. McCurtain Co.: ca.9 mi. NE of Broken Bow, 12 May 1987, Naczi 1575 (DOV*). PENNSYLVA- 
NIA. Bucks Co.: Tullytown, 31 May 1899, Williamson s.n. (PH). Dauphin Co.: 1.75 mi.E of Lykens, 10 
Jun 1952, Berkheimer 13006 (PH). Lehigh Co.: 1.25 mi.W by slightly NW of Lehigh Furnace, 25 Jun 
1922, Pretz 1 ne ae Philadelphia Co.: near Cedar Grove, 8 Jul 1926, Benner s.n. (GH). Schuylkill 
C W of Summit Station, 5 Jul 1938, Wagner 7401 (PH). RHODE ISLAND. Bristol Co.: 
Bristol, 1 ] res se 19460 (MO). SOUTH CAROLINA. Cherokee Co.: 6.2 mi. NW of Gaffney, 
2 May 1990, Naczi 2355 (DOV*).Clarendon Co.: Manning, 23 May 1914, Stone 317 int PH). Edgefield 
Co.: Sumter National Forest, 3.2 air mi. SSE of Lick Fork Dam, 22 May 1997, Nelson 18240 (DOV, USCH). 
atl ae Co.: E of Caesars Head, 31 May 1995, Naczi 4761 & Naczi (DOV*, MICH). ame se ca. 
W of Newberry, 14 May 1996, Horn 10310 (DOV, NBYC). Richland Co.: Con 
a Monument, ca. 2 km N of Weston lake, 29 Apr 1998, eso 19096 (DOV*, ean aie 
Dyer Co.: ca.6 mi.S of Dyersburg, 27 Jun 1995, Naczi 4839 & Reznicek (DOV, MICH). Hamilton Co.: 
Behind Tennessee Welcome Center, I-75, 30 Apr 1996, McNeilus 96-197 (TENN). Lincoln Co.: just E of 
New Hope, 4 Apr 1983, Sommers s.n. (VDB).Marion Co.: ca. 2 mi.S of Jasper, 29 May 1990, Naczi 2509 
& Kral (DOV). Perry Co.: Mousetail Landing State Park, E side of Tennessee River, 17 May 1990, Kral 
77633 (VDB). Tipton Co.: ca. 2.5 mi. NW of Gilt Edge, 27 Jun 1995, Naczi 4827 & Reznicek (DOV, MICH). 
Weakley Co.: ca. 5 mi. NW of Greenfield, 28 Jun 1995, Naczi 4868 & Reznicek (DOV, MICH). TEXAS. 
Harris Co.: ca.3 mi.N of US 90 at Sheldon, 5 Apr 1989, Orzell & Bridges 9018 (MICH*). Jasper Co.: ca. 
14 mi.W of Kirbyville, 3 May 1988, Naczi 1830 (MICH, ctb). Marion Co.: ca. 3 mi.S of Jefferson, 12 May 
1988, Naczi 1871 (MICH, ctb). Smith Co.: ca. 15 mi.N of Tyler, 7 May 1987, Naczi 1539 (MICH, PH, TEX, 
ctb). VIRGINIA. Princess Anne Co.: Little Neck, 17 Jun 1935, Fernald et al. 4593 (GH, PH). Sussex Co.: 
3.8 mi. WNW of Sussex, 4 Jun 1986, Wieboldt 6019 (DOV*). Westmoreland Co.: 0.5 mi. S of Nomini 
Grove, 8 Jun 1940, Hermann 10504 (PH). 


— 


— 


APPENDIX B 


Specimens of Carex digitalis measured for scatter plot (Fig. 3). Equal numbers of specimens 
were measured for C. digitalis var. digitalis (d), C. digitalis var. asymmetrica (a), and C. digitalis var. 
macropoda (m).All vouchers are at DOV.U.S.A. ALABAMA. Butler Co.: ca. 0.5 mi N of Oaky Streak, 

7 May 1995, Naczi 4687 (m).Conecuh Co.: ca.5 mi SSE of center of Evergreen, 1 : May 1995, Naczi 
4696 (a). Coosa Co.: ca.6 mi SW of Unity, 29 Apr 2000, Naczi 8504 & Ford (a). Lawrence Co.: Borden 
Creek, Bankhead National Forest, 15 May 1992, Bryson 11612 (m). Madison Co.: Huntsville, Monte 


1012 BRIT.ORG/SIDA 19(4) 


Sano State Park,6 May 1991, Naczi 2881 (d).Marion Co.: S of Bear Creek, 15 May 1992, Bryson 11592 
(a). Monroe Co.: 1.8 mi N of route 47-83 junction in Midway, 28 Apr 1991, Naczi 2817 (m). ARKAN- 
SAS.Cross Co.: W of Birdeye, 20 May 1990, Naczi 2441 (m).Stone Co.: ca.3 mi ENE of Fifty Six, 20 May 
1986, Naczi 1204 (d). FLORIDA. Hamilton Co.: Holton Creek Wildlife Management Area, 3 Apr 2000, 
Abbott 13392 (a).Leon Co.: ca.1 mi W of Bloxham, 5 May 1989, Naczi 2099 (a). GEORGIA. Effingham 
Co.: ca. 2.5 mi N of Clyo, 2 May 1991, Naczi 2831 (a). pened Co.: Blue Springs, 2 a 991, Naczi 
pe Talbot Co.: ca.4 mi NE of Pleasant Hill, 25 Apr 200 7184.28 & Ford (m). Towns Co.: 0.3 mi 
W of Rabun Co. border, W of route 76, 3 May 1991, Naczi pie (d). INDIANA. peg ae 4.9 mi N of 
Bristow, 27 May 1990, Naczi 2504 (d). KENTUCKY. Adair Co.: ca.9 mi S of Columbia, 16 Jun 1998, 
Naczi 7/399 & Reznicek (d). Boone Co.: 3 mi S of Petersburg, 12 Jun 1994, Naczi 4106 (d). Campbell 
mi S of Alexandria, 5 Jun 1996, Naczi 5646 et al. (d). Hardin Co.: ca.8.5 mi ENE of center of 
Elizabethtown, 15 Jun 1997, Naczi 6507 & Heeg (d). Laurel Co.: ca. 5.5 mi WNW of Bernstadt, 19 Jun 
1997, Naczi 6540 & Heeg (m). Mclean Co.,: ca.0.6 mi SSW of Beech Grove, 14 Jun 1997, Naczi 6484 & 
Heeg (m). MICHIGAN. St. Clair Co.: ca. 2 mi WNW of Blaine, 17 Jun 1988, Naczi 2025 (d). MISSIS- 
SIPPI. Pearl River Co.: ca. 3.2 mi S of Marion-Pearl River Co. line, 26 Apr 1991, Bryson 10749 et al. (a). 
Winston Co.: ca.5 mi NW of Louisville, 11 Apr 2000, Naczi 8338 et al. (m). OKLAHOMA. McCurtain 
Co.: ca.9 mi NE of Broken Bow, 12 May 1987, Naczi 1574 (m). PENNSYLVANIA. Clinton Co.: ca.2.3 mi 
NE of Rosecrans,4 Jul 1995, Naczi 4894 & Naczi (d). SOUTH CAROLINA. Dorchester Co.: W of Givhans, 
19 Apr 1991, Naczi 2763 (a). York Co.: ca. 3 mi NW of Bethany, 3 May 1989, Naczi 2080 (m). TEXAS. 
Jasper Co.: ca. 14 mi W of Kirbyville, 3 May 1988, Naczi 1831 (a). VIRGINIA. Halifax Co.: 5 mi SW of 
bic Grove, 30 Apr 1986, Wieboldt 5944 (m). 


ACKNOWLEDGMENTS 


We thank Bruce A. Ford and Stanley D. Jones for helpful comments on the manu- 
script, Mary A. Naczi, Nancy B. Bryson, Anton A. Reznicek, Bruce A. Ford, Tho- 
mas F. Wieboldt, Lori A. Heeg, and James R. Manhart for assistance with field 
work; the Kentucky State Nature Preserves Commission for permission to col- 
lect on selected preserves; and Anton Reznicek for assistance with various de- 
tails of the manuscript. We also thank the curators of BEREA, CM, DOV, EKY, 
FLAS, GH, IBE, KNK, KY, MICH, MO, MSC, MUR, NA, NCU, NY, OS, PENN, PH, 
TENN, US, USAM, VDB, WKU, WVA and ctb (Charles T. Bryson personal her- 
barium) for loans of specimens or assistance during our visits. National Sci- 
ence Foundation Grant BSR-9001260, Block Grants from the Horace H. 
Rackham School of Graduate Studies of the University of Michigan, anda grant 
from the Kentucky State Nature Preserves Commission provided financial sup- 
port to RFCN for some of the field work. 
REFERENCES 
Bryson, C.T. 1980. A revision of the North American Carex section Laxiflorae (Cyperaceae). 
Ph.D. Dissertation, Mississippi State Univ., Mississippi State. 
Case, PR. and C.WertH. 1995. The Trillium pusillum Michaux (Liliaceae) complex in Virginia. 
Il. sozyme evidence. Castanea 60:15-29. 
CxHester, E.W., B.E. Worrorb, R. Krat. H.R. DeSeim, and A.M. Evans. 1993. Atlas of Tennessee vas- 
cular plants. Vol. 1. Pteridophytes, Gymnosperms, Angiosperms: Monocots. Misc. Publ. 
No. 9. The Center for Field Biology, Austin Peay State Univ., Clarksville, Tennessee. 


NACZI ET AL 1013 


Ciark, R.C. 1967. Andrachne phyllanthoides (Nuttall) Muell.on the Cumberland Plateau of 
Alabama. Castanea 32:73-74 

Cusick, A.W. 1988. Noteworthy collections. Castanea 53:311-314. 

Cusick, A.W. 1989. Seven indigenous taxa new to the vascular flora of Kentucky. Castanea 
54:268-270. 

Deam, C.C. 1940. Flora of Indiana. Indiana Dept. of Conservation, Indianapolis. 

Fernaco, M.L. 1950. Gray's Manual of Botany, 8th ed. American Book Co., New York. 

Funt, R.F.1971.Glacial and Quaternary Geology. John Wiley and Sons, New York. 

Geason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada, 2nd ed. New York Botanical Garden, Bronx. 

JOHNSON, G.P. 1988. Revision of Castanea sect. Balanocastanon (Fagaceae).J.Arnold Arbor. 
69:25-49, 

KUKENTHAL, G. 1909, Cyperaceae - Caricoideae. In: A. Engler, ed. Das Pflanzenreich, IV. 20, Heft 
38.Wilhelm Englemann, Leipzig. Pp. 1-824. 

Lirtte, E.L. 1977. Atlas of United States trees, vol. 4. United States Government Printing 
Office, Washington, DC. 

Lona, A.A. 1989. Disjunct populations of the rare shrub, Neviusia alabamensis Gray (Ro- 
saceae). Castanea 54:29-39. 

McKinney, L.E.,G. Ligay,and R. Mears. 2000.New and noteworthy records of Carex (Cyperaceae) 
from Kentucky. Castanea 65:221-224. 

Mackenzie, K.K. 1935. Cyperaceae - Cariceae. N. Amer. Fl. 18:169-478. 

Manuaart, J.R. 1986. Foliar flavonoids of the North American members of Carex section 
Laxiflorae (Cyperaceae). Biochemical Systematics and Ecology 14:85-90. 

McNeius, V.B. 1992. Carex ouachitana (Cyperaceae) new to Tennessee. Sida 15:154—-155. 

Mever, F.G. 1997a. Fothergilla. In: Flora of bog pals. eon Se eds. Flora of 
North America North of Mexico,Vol.3.Magnoliophyt nd Hamamelidae 
Oxford Univ. Press, New York. Pp. 365-366. 

Meyer, FG. 1997b. Magnolia. In: Flora of North America Editorial Committee, eds., Flora of 
North America North of Mexico, Vol. 3. Magnoliophyta: Magnoliidae and Hamamelidae. 
Oxford Univ. Press, New York. Pp. 4-10. 

Naczi, R.F.C. 1989. Circumscription of sections and phylogeny in a lineage within Carex 
(Cyperaceae). Suppl. to Amer. J. Bot. 76:261. 

Naczi, R.F.C. 1992. Systematics of Carex section Griseae (Cyperaceae). Ph.D. Dissertation, 
Univ. of Michigan, Ann Arbor. 

Naczi, R.F.C. 1997. Phylogeny reconstruction in Carex sections Careyanae and Granulares 
(Cyperaceae). Suppl. to Amer. J. Bot. 84:218-219. 

Naczi, REC. 1999. Chromosome numbers of some eastern North American species of 
Carex and Eleocharis (Cyperaceae). Contr. Univ. Michigan Herb. 22:105-119 

Naczi, RAC. and C.T. Bryson. 1990. Noteworthy records of Carex (Cyperaceae) from the 
southeastern United States. Bartonia 56:49-58. 

Naczi, R.F.C., A.A. Reznicek, and B.A. Foro. 1998. Morphological, geographical, and ecological 
differentiation in the Carex willdenowii complex (Cyperaceae). Amer. J. Bot. 85:434-447. 


1014 BRIT.ORG/SIDA 19(4) 


Nixon, K.C. 1997. Castanea. In: Flora of North America Editorial Committee, eds., Flora of 
North America North of Mexico, Vol.3. Magnoliophyta: Magnoliidae and Hamamelidae. 
Oxford Univ. Press, New York. Pp. 439-442. 

OHio Division Or NaTuRAL AREAS AND Preserves. 2000. Rare native Ohio plants: 2000-01 status 
list. Ohio Department of Natural Resources, Columbus. 

OvoxHam, M.J. and M. Zinck. 1997. New and noteworthy records from the vascular flora of 
Nova Scotia. Canad. Field-Naturalist 111:393-398. 

Rosison, H.W. and R.T. Auten. 1995. Only in Arkansas: A study of the Endemic Plants and 
Animals of the State. Univ. Arkansas Press, Fayetteville. 

Scora, R.W. 1967. Interspecific relationships in the genus Monarda (Labiatae). University of 
California Publications in Botany 41:1-59. 

Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, 2nd ed. 
Kinko's, Fayetteville, Arkansas. 

Starr, J.R., RJ. Bayer, and B.A. Foro. 1999. The phylogenetic position of Carex section 
Phyllostachys and its implications for phylogeny and subgeneric circumscription in 
Carex (Cyperaceae). Amer. J. Bot. 86:563-577. 

Tuckerman, E. 1843. Enumeratio methodica caricum quarundam. Schenectady, New York. 


ASYNOPSIS OF THEFERNS AND PERN ALLIES OP 
NEBRASKA, WITH MAPS OF THEIR DISTRIBUTION 


Steven B. Rolfsmeier Robert B. Kaul 
Charles E. Bessey Herbarium Charles E. Bessey Herbarium 
University of Nebraska State Museum University of Nebraska State Museum 
University of Nebraska-Lincoln University of Nebraska-Lincoln 
Lincoln, NE 68588-0514, U.S.A Lincoln, NE 68588-0514, U.S.A 


David M. Sutherland 


Omaha, NE 68182-0040, U.S.A. 


ABSTRACT 


An annotated list and county-by-county distribution an are presented for hs 2 2 oa of fer ns 
land 


and fern allies native to NED RASS: based upon betel a 
N la, Selaginella rupestris, five species of Equisetum, 


: literature 
and 25 species: in ie pena o ferns. tate ee species are here verified ve the pas time, based 
pensylvanica, Ophi 


P 
Pell labella ssp. glabella. Isoét | od li 2000 ete rear en 
Rejected are oublished reports of Ly ee innotinum, Selaginella densa, Equisetum xlitorale, E. 
palustre, E. pratense, E. variegatum, Asplenium tachentnes Azolla caroliniana, Botrychium 
dissectum, B. matricariifolium, B. multifidum, B. neglectum, Cryptogramma acrostichoides, 
Ophioglossum vulgatum, Osmunda cinnamomea, O. claytoniana, O. regalis, and Woodsia scopulina. 
e erroneous reports have been published repeatedly for more than a century, but others are 


recent. 
RESUMEN 


Se picseitan¢ un lista anolada y mae de eens oe condados, de las 32 especies de helechos y 
U,), basados en estudios de campo, examen de 


herbanids y evaluaclon critica de la literatura. Como nativas para este estado se cuentan Isoétes 
lanopoda upestris, cinco especies de aaa y 25 especies de helechos distribuidas 
e 


I 
en 18 géneros. Se citan tres especies como nativas por primera vez, basandose en recolec 
recientes: Matteuccia struthiopteris var pensylvanica, Soh an engelmannii, y Pellaea alabella 
Isoétes melanopoda fue redescubierta en el ano 2000, siendo el primer registro de esta especie de 
941. Se rechazan como erroneas las citas publicadas de la presencia de Lycopodium annotinum, 
Selaginalla densa, Equisetum Xlitorale, E. palustre, E. pratense, E ales Sania ate? 
Azolla caroliniana, Botrychium dissechuni, B. matricariifolium, B. multifidum, B. neglectum, 
Cryptogramma acrostichoides, Ophioglossum eeu Osmunda cinnamomea, O. claytoniana, O. 
Algunas de estas citas erroneas han sido publicadas en repetidas 


regalis, y Woodsia scopulin 
ocasiones desde hace mas un siglo tras que otras son recientes. 


SIDA 19(4): 1015 — 1026. 2001 


1016 BRIT.ORG/SIDA 19(4) 


INTRODUCTION 


While preparing our New Century Flora of Nebraska (Kaul, Sutherland, & 
Rolfsmeier, in prep.), we reinterpreted and corrected many dubious or errone- 
ous reports of the state’s flora, among which those of the ferns and their allies 
were especially confused and contradictory. 

The earliest credible report of Nebraska’s complete vascular flora was the 
list by HJ. Webber (1890), but the first definitive account of Nebraska’s ferns 
was that of Thomas J. Fitzpatrick (1920), which was based upon holdings in 
the Charles E. Bessey Herbarium (Nes) of the University of Nebraska-Lincoln. 
The only floristic manuals solely devoted to the state’s flora (Petersen 1923: 
Winter 1936) were also based upon that herbarium’s collection, which greatly 
expanded under Bessey’s direction from the 1880s until his death in 1915. Re- 
gional and more extensive books that include Nebraska were based upon that 
and many other collections in and outside Nebraska: Britton & Brown (1896, 
1913, but not Gleason (1952)); Rydberg (1932), Fernald (1950), Petrik-Ott (1979), 
Lellinger (1985), GPFA-Great Plains Flora Association (1977, 1986), FNA-Flora 
of North America Editorial Committee (1993), and various journal papers. With 
each succeeding publication, old distributional errors were perpetuated and new 
ones were introduced. 

The only county-by-county maps of all the state’s ferns and fern allies were 
those of Atlas ofthe Flora of the Great Plains (GPFA 1977), to which we contrib- 
uted, and of The Pteridophytes of Kansas, Nebraska, South Dakota and North 
Dakota (Petrik-Ott 1979). The state-outline maps of Flora of North America 
(FNA 1993) are more generalized. There are mapping errors in all those publi- 
cations that we seek to correct in this paper, and our extensive field and her- 
barium work has added many distributional records. 

A continuing source of errors is the E.M. Hussong collection at NEB, suppos- 
edly from Franklin County in the 1890s, according to the labels. The collection 
includes ferns and many angiosperms that are far out of range for Nebraska. 
For example, Osmunda regalis has been attributed to Nebraska for more thana 
century, based upon those specimens, but it is yet unknown anywhere in the 
state, and Franklin and nearby counties lack suitable habitat. The handwriting 
on the labels is not Hussong’s, and we reject the records as representing Nebraska 
plants. 

The massive cattle-ranching and agricultural development of the past 150 
years has greatly reduced the state’s natural vegetation, especially that of the 
prairies, but suitable habitats remain and most species can be found today, if 
only in remote or widely scattered sites. Most pteridophytes in Nebraska are at 
the edge of their range and, as such, many have always been scarce, and a few 
have not been seen for decades. Relatively pristine are the Sandhills, 23,000 
square miles of prairie-vegetated, uncultivated dunes and sandplains, the high 


ROLFSMEIER ET AL., FERNS AND FERN ALLIES OF NEBRASKA 1017 


water-table producing thousands of lakes, ponds, marshes, and fens; and the 
valley of the Niobrara River, its bluffs and sheltered tributaries lined for much 
of its length with forests of ponderosa pine and hardwoods. Some species in 
Niobrara Valley forests and Sandhills fens are apparently relicts from cooler, 
wetter, early post-Pleistocene times, such as the now-disjunct Ophioglossum 
pusillum, Dryopteris cristata, D.carthusiana, Athyrium filix-femina,and many 
angiosperms and animals (Kaul et al. 1988; Kaul & Rolfsmeier 1993). The pon- 
derosa pine forests of the Pine Ridge, Wildcat Hills, and Niobrara River valley 
still have much native flora, as do some of the oak-hickory forests of the Mis- 
souri River valley and its tributaries. With ranching and agrarian settlement 
came suppression of prairie fires, leading to major expansion of deciduous for- 
ests in the eastern, wettest third of the state, but their pteridophyte flora has 
not correspondingly expanded and is yet confined to the region of the original 
forests there. 


MATERIALS AND METHODS 


We have collected most species in the field during our combined 86 years of 
field work from 1964 through 2001. In addition, we examined all specimens at 
these herbaria in Nebraska: Charles E. Bessey Herbarium (nes) at the University 
of Nebraska-Lincoln; University of Nebraska at Omaha (oma); University of 
Nebraska at Kearney; Chadron (cscn), Peru, and Wayne state colleges; Cedar 
Point Biological Station; Doane College; and Nebraska Wesleyan University. We 
also examined all Nebraska-collected specimens at the University of Kansas 
(kANU), Kansas State University (ksc), Fort Hays State University, University of 
South Dakota (spu), South Dakota State University (spc), and University of 
Wyoming (rm), and we saw some Nebraska specimens at the Missouri Botani- 
cal Garden (mo) and the New York Botanical Garden (ny). 

Our nomenclature follows Flora of North America (FNA 1993), with syn- 
onyms as used in our cited references that specifically attributed the plants to 
Nebaska by text, map, or both. Species marked with a black diamond (@) are 
here verified for the first time as native to Nebraska and were discovered in the 
late 1990s and 2000. Year of first collection is given for the rarer species, some 
of which have not been found again. Counties and some physiographic features 
are named in Figure l, and all species are mapped by county in Figure 2. 


ACCEPTED SPECIES 


Division LYCOPODIOPHYTA 


ISOETACEAE 

Isoétes melanopoda Gay & Durieu ex Durieu.—Collected from the Rainwater 
Basins several times, 1880s-1941, but not again until 2000. Most habitat long 
since drained and plowed for agriculture. Mapped for the entire eastern half of 


1018 BRIT.ORG/SIDA 19(4) 


wt ie 
PR we wen - EVA PAHA BOYD \ 
wa ,} » \ \ ie fea ° # 
ve \\ he ® 
vine ay N e 
. 1X 
. KNOX KOT 
A 
s . iS 
ea ie 
el® ¢ CHERRY BROWN [ROCK [HOLT 1 Ld RSTO 
Xo IDAN s ANTE. ; 
pOPE | * a eles M 
Ni IGAR- Pas TAN 
iaett . RANT WooKer ITHOMAS frame lroue Jeep hwreeen | | ADISONITON _{cumi 
: ete ee . 
Ke . ~ yl -]oooce, Wash: 
BANNER e =a ATTE a is 
a INANCE 
PMERRIC! ~ ia: 
= - 
powssn! Andi r Ane 
KIMBALL HEYENNE _|DEVEL PLOK puis SAUNDERS 
af 
a fe / CASS 
HALL TON YORK _|SEWARD 
LAN— 
= J SJ Ww i CASTER |OTOE 
_ fh FALL 
ADAMS. |Chay £_qSALINE JOHN- 
ae rae SON MAHA 
RANK- BUC K- JEFFER- 
HARLAN [LIN WEBSTER JOLLS THAYER [SON IGAGE PAWNEE RICHARD: 
‘ y etal “ oe oe peer Sey aT frais 
Fic. 1. Map of Nebraska with counties, major rivers physiogray M: Missouri, N: Niobrara 
In. pl i +4 i rhiIimVHe| uth L | Pay J ee j , Nats Aas (0).C AL cil 
1d P: F R: Pine Ridge f | I 
prairie 1d 1 sandplains, with tt 1s of ponds, lakes, marshes, and fens; dashed line(Rw): Rai 
9 U ’ D 
tear Racine | In] f | | lud 


Ler DdSills Pv " y } y 


Scale bar = 100 mi 160 km. 


the state by FNA-Taylor et al. (1993), but known only from five south-central 
counties. 
SELAGINELLACEAE 
Selaginella rupestris (L.) Spring —Locally abundant in parts of the Niobrara and 
Little Blue river drainages, scattered elsewhere. 

Division EQUISETOPHYTA 


EQUISETACEAE 
Equisetum arvense L.—Common state-wide, except absent from the southwest. 

Equisetum xferrisii Clute [=-Equisetum intermedium (A. A. Eaton) Rydb.- 
Rydberg (1932)|—Occasional in the eastern half of the state; many reports in 
GPFA (1977) were based upon depauperate specimens of FE. hyemale. 

Equisetum fluviatile L. [=Equisetum limosum L.-Bessey (1892)]—Known 
only by collections from the eastern Sandhills in 1892, 1909, and 2000, and from 
one site near the Missouri River, 1941-1952, 1987, 2001. Mapped for the entire 
northern half of the state by FNA-Hauke (1993). 

Equisetum hyemale L. ssp. affine (Engelm.) Calder & Roy L. Taylor [=Equi- 
setum affine Engelmann-Rydberg (1932); Equisetum hyemale L.-Bessey (1892), 
Britton & Brown (1896, 1913), GPFA (1977), Equisetum hyemale L. var. affine 
(Engelm.) A.A. Eaton-GPFA (1986); Equisetum robustum A. Braun- Webber 
(1890), Fitzpatrick (1920), Rydberg (1932)]—Common over most of the state, 
except the southwest. 


ROLFSMEIER ET AL., FERNS AND FERN ALLIES OF NEBRASKA 1019 


Equisetum laevigatum A. Braun [=Equisetum kansanum J.H. Schaffn.- 
Rydberg (1932)]—Common statewide. 


Division POLYPODIOPHYTA 


ASPLENIACEAE 

Asplenium platyneuron (L.) Britton, Sterns, @ Poggenb.—First collected in 1986, 
by J. Locklear in Jefferson County (Rolfsmeier et al. 1988), where locally abun- 
dant; a single plant discovered and photographed in Washington County in 
1988 (Garabrandt 1988), the photograph at oma. It remains to be seen whether 
this plant will increase its range in Nebraska as it has elsewhere (Wagner & 
Johnson 1981). 


AZOLLACEAE 
Azolla mexicana C. Presl.—Sporadically abundant, sometimes massively so. 


DRYOPTERIDACEAE 

Athyrium filix-femina (L.) Mertens var. cyclosorum Rupr. [=Asplenium filix- 
foemina (L.) Bernh.-Botanical Survey of Nebraska (1893); Athyrium angustum 
(Willd.) C. Presl-Rydberg (1932)|—Known only as independent 1893 collections 
by J.M. Bates, FC. Clements, and PA. Rydberg, on different dates and at different 
sites in the Niobrara River valley, and perhaps yet present in pristine forests of 
Long Pine Canyon. The sites are disjunct from the main ranges of var. cyclosorum 
to our west and var. angustum (Willd.) G. Lawson to our east. FNA-Kato (1993) 
attributed only var. angustum to Nebraska, but the specimens from the wild, 
with their rounded sori and long-ciliate indusia, are var. cyclosorum. In 1979, 
R.E. Brooks annotated those specimens as ssp. cyclosorum (Rupr) C. Christens., 
but in GPFA (1986) he did not attribute the species to Nebraska; the key to vari- 
eties there is incorrect. Petrik-Ott (1979) did not specify infraspecific taxa, and 
an Adams County specimen she cited but did not map was from a garden; sev- 
eral other specimens in herbaria are of cultivated plants. 

Cystopteris bulbifera (L.) Bernh.-GPFA (1977, 1986).—First collected in 1972 
(Anderson 1974), in Richardson County in the extreme southeastern corner of 
the state, and still abundant at that site, but unknown elsewhere in the state. 
Not attributed to Nebraska by FNA-Haufler et al. (1993). 

Cystopteris fragilis (L.) Bernh. [=Filix fragilis (L.) Gilib-Petersen (1923), 
Rydberg (1932)|—Common in the northern and eastern halves of the state. 

Cystopteris protrusa (Weath.) Blasdell—Common in the eastern one-fifth 
of . state, especially southeastward. 

stopteris tenuis (Michx.) Desv. |=Cystopteris fragilis (L.) Bernh., in part- 
GPFA Vee 1986)|—Locally abundant in the eastern one-fifth of the state. Moran 
(1983) and FNA-Haufler et al. (1993) mapped it for one station in the extreme 
west, but we cannot verify its presence there. 

Dryopteris carthusiana (Vill.) H.P Fuchs[=Aspidium spinulosum Sw Web- 


1020 BRIT.ORG/SIDA 19(4) 


ber 1890); Dryopteris spinulosa (O.F Mull.) Kuntze-Fitzpatrick (1920), Petersen 
(1923), Rydberg (1932), Dryopteris spinulosa (O.F Mull.) Watt-GPFA (1986)]— 
Collected infrequently between 1892 and 2001; locally common today at scat- 
tered sites in the paper-birch forests of the Niobrara River valley in Cherry 
County, and in oak-wooded ravines between bluffs of the Missouri River in 
Dakota and Thurston counties. 

Dryopteris cristata (L.) A. Gray.—Collected by Rydberg in 1893 from shrub- 
filled fens of Hooker County, in the Sandhills, disjunct 150 mi/240 km west of 
the main range; not found since then, but potentially persisting in the many 
remaining fens. Erroneously attributed also to adjacent Thomas County by 
Petersen (1923) and GPFA (1977), based upon incorrect interpretation of the lo- 
cation as written on the labels. 

¢Matteuccia struthiopteris (L.) Tod. var. pensylvanica (Willd.) CV. Morton.— 
First collected from the wild in 2000, in native habitat in Thurston County, 
remote from past or present habitations. Commonly cultivated but not known 
to have naturalized in the state, although seemingly capable of doing so. 

Onoclea sensibilis | —Abundant today in Sandhills marshes and in the 
Elkhorn, Loup, and lower Little Blue river valleys, but rare and mostly extir- 
pated elsewhere. 

Woodsia obtusa (Spreng.) Torr—Occasional to locally common in rocky 
woods in the eastern one-fifth of the state. 

Woodsia oregana D.C. Eaton ssp. catheartiana (B.L. Robins.) Windham.— 
Occasional in the western two-thirds of the state. Mapped for all but the ex- 
treme southeastern corner of the state by FNA-Windham (1993), but absent 
from a much greater area (Fig. 2). 


MARSILEACEAE 
Marsilea vestita Hook. & Grev.—Locally abundant, sometimes massively so, in 
the Platte River valley and Rainwater Basins, scarce elsewhere. 

Pilularia americana A. Braun.—First and last collected in 1966, froma large, 
shallow Sandhills lake in Cherry County (McGregor 1967). 


OPHIOGLOSSACEAE 
Botrychium campestre W.H. Wagner & Farrar.—First collected in 1982, from a 
bur-oak and red-cedar forest on the floodplain of the Niobrara River, Brown 
County (Freeman & Churchill 1983, as B. matricariifolium A. Braun, but later 
formally described as B. campestre in Wagner & Wagner (1986)), and still 
thriving there; two nearby populations discovered in 2000. 
Botrychium virginianum (L.) Sw—Common in eastern and northern counties, 
less so westward. Mapped for the entire state by FNA-Wagner and Wagner (1993), 
but unknown to us in the entire southwestern half, which lacks suitable habitat. 
¢Ophioglossum engelmannii Prantl.—Known only by collections in 1999 
from Pawnee County, the northern edge of its range. 


ROLFSMEIER ET AL., FERNS AND FERN ALLIES OF NEBRASKA 1021 


Ophioglossum pusillum Raf.[=Ophioglossum vulgatum L. var. pseudopodum 
(S.E Blake) Farw.-Fernald (1950), Brooks (1980), GPFA (1986)|—Locally abun- 
dant on peat mounds in northwestern Sandhills fens, and on shallow peat near 
springs in the Niobrara River valley. First collected in 1912, but not again until 
1982 and subsequently at seven other sites. The habitats are not especially threat- 
ened, and a few are preserved. 


PTERIDACEAE 

Adiantum pedatum L.—Common in oak-hickory forests on bluffs near the Mis- 
souri River and nearby tributaries, and known from a single collection from 
Rock ie in the Niobrara River valley. 

Argyrochosma dealbata (Pursh) Windham [=Notholaena dealbata Kunze- 
Webber (1890), Britton & Brown (1913), Fitzpatrick (1920), Petersen (1923), 
Rydberg (1932), Fernald (1950), GPFA (1977, 1986)]—First and last collected in 
1888, from Weeping Water Valley, Cass County. The site, now severely disturbed 
by quarrying of limestone, is northernmost for the species. 

Cheilanthes feei T. Moore [=Cheilanthes lanuginosa Nutt.-Webber (1890)|— 
Uncommon in western counties, absent elsewhere; mapped for the entire state 
by FNA-Windham & Rabe (1993). 

Pellaea atropurpurea (L.) Link —Mapped for the entire state by FNA- 
Windham (1993), but known to us only from one western county, where it is 
locally abundant in narrow, shady, cool canyons, and from seven eastern coun- 
ties, where it is uncommon. 

¢Pellaea glabella Mett.ex Kuhn ssp. glabella—The only specimen we have 
seen is our collection from Gage County, 1998, although the plant was earlier 
mapped for Nebraska by Gastony (1988), but without citation of specimens. It 
was then attributed to the state and mapped for Richardson County by FNA- 
Windham (1993), perhaps based upon Gastony’s map. 


THELYPTERIDACEAE 

Thelypteris palustris Schott var. pubescens (G. Lawson) Fernald [=Aspidium 
thelypteris (L.) Sw-Webber (1890); Dryopteris thelypteris (L.) A. Gray- 
Fitzpatrick (1920), Petersen (1923), Rydberg (1932), Thelypteris palustris Schott- 
GPFA (1977, 1986)|—Common now in Sandhills marshes and fens, sometimes 
massively so, and locally abundant in the Elkhorn and Loup river drainages 
and in Jefferson County; essentially extirpated elsewhere. 


EXCLUDED SPECIES 
Division LYCOPODIOPHYTA 


Seana 

inum L. was reported for Nebraska by Petrik-Ott (1975, 1979) and GPFA (1977), based 
ipen an 1892 specimen supposedly from Cherry County and attributed to J.M. Bates. Bates included 
no label with the specimen, and all label data are in the handwriting of Thomas J. Fitzpatrick. Al- 


1022 


BRIT.ORG/SIDA 19( 


mail 


4 in = 
Adiantum pedatum 


| | | Le ee IDs 

ian {A 

tee y i" 
EH 


Coan EES | 1 1 
Athyrium filix-femina 
cyclosorum 


a \— Le F a ae 
eet HAE oie 


Azolla mexicana Botrychi 


L 


Ebi 


Botrychium virginianum 


[ p 
Cystopteris protrisa 


a! 


ch : 
Cystopteris tenuis 


[Tt im 74, 
a= 1 | | ee 
Equisetum x ferrissii Equisetum fluviatile 
Fic. 2. Distributi ps of all Nebraska species of f 1 f llies, as collected from 1873 through 2001.The records 
£, kK kK d Pl + ime 1 L = Al Poe & 2 L L hd inf a is 
é rer i 


ri 


though correctly identified, the specimen was rejected by Brooks (1978) because the mosses with the 
specimen are not native to Nebraska, and Petrik-Ott (1979), in the addendum to her book, agreed. 
Neither this nor any other species of Lycopodium, sensu lato, is known in Nebraska. 
SELAGINELLACEAE 
Selaginella densa Rydb. was reported from Holt County by Petrik-Ott (1975, 1979) and GPFA (1977), 
ut based upon a misidentified specimen of S. rupestris. It was attributed to Nebraska and mapped 
for the extreme northwestern corner of the state by FNA-Valdespino (1993), but we have not found it 
there, although it is nearby in Wyoming and South Dakota. 


ROLFSMEIER ET AL., FERNS AND FERN ALLIES OF NEBRASKA 1023 


Y Uy [iL ae 
ie cL 
ae ine Cy! 
MUG Y oe ry 

Yes TT 

Reisen laevigatum ; Isoetes melanopoda 


— 


“— 


t ia q 
aetna stiopterts 
NSU 


Thelypteris palustris a er 


YY, 
OW 


Wecduncban obtusa 7 2 


Fic. 2. (cont.) 
Division EQUISETOPHYTA 


EQUISETACEAE 

Equisetum Xlitorale mnees ex Rupe was bapoee for the entire northern half of Nebraska by FNA- 
Hauke (1993), but her collected it in the state nor have we seen voucher sf 

herbarium 


Equisetum palustre L. was reported for Nebraska by Taylor (1970), but we have not seen specimens. 
Equisetum pratense Ehrh. Bessey (1892) reported a specimen from Sowbelly Canyon, Sioux 
County, supposedly collected in August, 1890, but we have not found such a epeomen: 


Tea: 


Equisetum variegatum Schleich. ex F Weber & D. Mohr 
the state by Webber (1890), Fitzpatrick (1920) and P 1923 bas those reports were based upon 
misidentified, depauperate specimens of Equisetum laevigatum. The records were rejected by Petrik- 
Ott (1975, 1979) and Brooks (1986). 


Division POLYPODIOPHYTA 


ASPLENIACEAE 
Asplenium trichomanes L. ssp. trichomanes was not reported for Nebraska by Moran (1982), but it was 
both mapped and cited for the state by FNA-Wagner et al. (1993). Moran (1995, in litt.) could not 


1024 BRIT.ORG/SIDA 19(4) 


account for the discrepancy. The nearest vouchered stations we know are in the Black Hills of South 
Dakota. 

ZOLLACEAE 
Azolla caroliniana Willd. Reports before 1950 were made by authors who did not distinguish this 


from Azolla mexicana, porn species were reported for Nebraska by FNA-Lumpkin (1993), but the 
, which are lacking in most of our specimens. 


distinction 
We reject reports of A. nim. as did Pecule Out (1979), until further evidence is available 
DRYOPTERIDACEAE 

Woodsia scopulina D.C. Eaton was reported by Rydberg (1932), but we have not found vouchering 
specimens. It occurs in the Black Hills of South Dakota and perhaps is in the nearby Pine Ridge of 
Nebraska's northwestern corner 

OPHIOGLOSSACEAE 

Botrychium dissectum Spreng. was reported (as Botrychium obliquum Muhl.) by auiagh oe _ wh 
suggested that the latter name might be correct for the Hussong collection of B. multi 


see, below). 

Botrychium matricariifolium A. Braun was reported as new to Nebraska by Freeman & Churchill 
(1983) and Brooks (1986), based upon specimens of Botrychium campestre (q.v. above), which was not 
described until 1986 

B hi Itifidurn (S.G.Gmel.) Rupr. A (H ) at NEB, supposed|y from Fran- 
klin County, was so identified by Petrik-Ott 1975, 1979). The enecinen is from a large collection of 

very doubtful provenance, and the record was excluded by GPFA (1977, 1986) and is rejected by u 

Botrychium neglectum A. Wood was reported by Britton and Brown (1913), Fitzpatrick (1920), 
and Petersen (1923), the latter two reports being based on the specimen later reported as B. multifidum 
by Petrik-Ott (1979). The placement of this name is in doubt, and Brooks (1986) included it as a syn- 


(or 


onym of B. matricariifolium. 

Ophioglossum vulgatum |. was first reported for Nebraska by Bates (1912). All reports before 
1980 were by authors who did not distinguish this species from O. pusillum Raf., of which there are 
old and recent records for the state. 
OSMUNDACEAE 
Osmunda cinnamomea L. was reported for Nebraska by Lellinger (1985) but not - GPFA (1977, 1986) 
or FNA-Whetstone Sadho 993); we nave not found vouchering specimen 
al s reported by Fitzpatrick (1920), based NEB “with- 


Osmunda cl Be 
out definite rere far “loubtfully referred to oNebuade? The report was slater accepted by Rydberg 
(1932) and rejected ae Petrik-Ott (1975, 19 
unda regalis |. var. spectabilis (Will d.) A. Gray was first reported for Nebraska in 1897 (Fern 
Bull., p. 66), based upon specimens at nrp that were collected ees E. M. Hussong, supposedly from 
ei County but of doubtful provenance. The report was reiterated by Britton & Brown (1913), 
ee (1920), Petersen (1923), Petrik-Ott 1975, 1979), and iene (1985), but not by Fernald 
), Gleason (1952), GPFA (1977, 1986), or FNA-Whetstone & Atkinson (1993). Franklin County 
ne suitable habitat, and the plant has not been found anywhere in the state. 


PTERIDACEAE 

ryptogramma acrostichoides RK. Br. was reported by Fitzpatrick (1920) and Petersen (1923) on the ba- 
sis of a single frond that was sent to C_E. Bessey for determination. The specimen is from a collection 
by E.M. Hussong and supposedly from Franklin County, but with doubtful locality data that is not 
written in oo hand. Petrik-Ott (1975, 1979) considered the specimen not to be from Nebraska, 


and so do 


ROLFSMEIER ET AL., FERNS AND FERN ALLIES OF NEBRASKA 1025 


REFERENCES 


ANDERSON, G.J. 1974. Cystopteris bulbifera new to Nebraska. Amer. Fern J.64:30. 

Anonymous. 1894. Additions to the reported flora of Nebraska made during 1893. Botani- 
cal Survey of Nebraska 3, report for 1893:4-19. 

Bates, JM. 1912. Ophioglossum vulgatum in Nebraska. Fern Bull. 20:67. 

Bessey, C.E. 1892. A second edition of Webber's “Appendix to the catalogue of the flora of 
Nebraska," with a supplementary list of recently reported species. Contr. to Botany, 
Univ. Nebraska, n.s. 3:1-55. 

Britton, N.L.and A. Brown. 1896 (15 ed.), 1913 (24 ed). An illustrated flora of the northern 
United States, Canada, and the British Possessions, vol. 1. New York Botanical Garden, 
New York. 

Brooks, R.E. 1978. A deletion [Lycopodium annotinum] from the pteridophyte flora of Ne- 
braska. Amer. Fern J. 68:94-95. 

Brooks, R.E. 1980. Additions to the pteridophyte flora of the Great Plains. Amer. Fern J. 
70:91-92. 

Brooks, R.E. 1986. Equisetum, Botrychium. |n: Flora of the Great Plains (GPFA-Great Plains 
Flora Association, 1986, q.v.). Pp.42-48. 

FerNALD, M.L. 1950. Gray's Manual of Botany, 8th ed. American Book Co., New York. 

Fitzpatrick, T.J.1920.The fern flora of Nebraska. Amer. Fern J. 10:5-15, 33-44. 

FNA—Fiora of NortH America EpitoriAL Committee. 1993. Flora of North America, North of 
Mexico, vol. 2.Oxford University Press, New York.C.H. Haufler, R.C. Moran, M.D.Windham- 
Cystopteris; R.L. Hauke-Equisetum; T.A. Lumpkin—Azolla; W.C. Taylor, N.T. Luebke, D.M. 
Britton, RJ. Hickey, D.F. Brunton-/soétes;|.A.Valdespino—Selaginella; W.H.Wagner Jr.,R.C. 
Moran, C.R. Werth—Asplenium; W.H. Wagner Jr. and F.S.Wagner—Botrychium; R.D.Whet- 
stone and T.A.Atkinson—Osmunda;M.D.Windham-—Pellaea, Woodsia; M.D.Windham and 
EW. Rabe—Cheilanthes. 

Freeman, C.C.and S.P.CHurcHilt. 1983. Noteworthy pteridophyte records for Nebraska. Amer. 
Fern J. 73:29-30. 

Garasranot, M.M. 1988. An annotated list of the vascular plants of Fontenelle Forest and 
Neale Woods in eastern Nebraska. Trans. Nebraska Acad. Sci. 16:31-49. 

GastTony, G.J. 1988. The Pellaea glabella complex: electrophoretic evidence for the deriva- 
tions of the agamosporous taxa and a revised taxonomy. Amer. Fern J. 78:44-67. 

GPFA—Grear PLains Flora Association. 1977. Atlas of the flora of the Great Plains. lowa State 
University Press, Ames. 

GPFA-Grear PLAINS Flora Association. 1986. Flora of the Great Plains. University Press of 
Kansas, Lawrence. 

GLeason, H.A.1952.The new Britton and Brown illustrated flora of the northeastern United 
States and adjacent Canada, vol. 1. New York Botanical Garden, NY. 

KAUL, R.B., G.E. Kantak, and S.P. CHURCHILL. 1988. The Niobrara River valley,a postglacial migra- 
tion corridor and refugium of forest plants and animals in the grasslands of central 
North America. Bot. Rev. 54:44-81. 


1026 BRIT.ORG/SIDA 19(4) 


Kaut, R.B. and S.B. Rotrsmeier. 1993. Native vegetation of Nebraska. Map 1:1,000,000, 24 x 
36" 16 colors + text. Conservation & Survey Division, University of Nebraska-Lincoln. 

Levuincer, D.B. 1985. A field manual of the ferns and fern-allies of the United States and 
Canada. Smithsonian Institution Press, Washington, D.C 

McGrecor, R.L. 1967. Pilularia americana A. Braun new to Nebraska. Amer. Fern J.57:136. 

Moran, R.C. 1982. The Asplenium trichomanes complex in the United States and adjacent 
Canada. Amer. Fern J. 72:5-11. 

Moran, R.C. 1983. Cystopteris tenuis (Michx.) Desv.:a poorly understood species. Castanea 
48:218-223. 

Petersen, N.F. 1923. Flora of Nebraska, 3rd edition. Plainview, Nebraska, published by the 
author. 

Perrik-Ort, AJ. 1975. A county checklist of the ferns and fern allies of Kansas, Nebraska, 
South Dakota, and North Dakota. Rnodora 77:478-511. 

Petrik-Ort, AJ. 1979. The pteridophytes of Kansas, Nebraska, South Dakota, and North 
Dakota. Beih. Nova Hedwigia 61:1-332. 

Rotrsmeier, $.B., R.B. KAuL, M.M. GaraBRaANot, and D.M. SUTHERLAND. 1988. New and corrected 
floristic records for Nebraska. Trans. Nebraska Acad. of Sci. 16:115-121. 

Ryoser, PA. 1932. Flora of the prairies and plains of central North America. New York Bo- 
tanical Garden, NY. 

Taytor, T.M.C. 1970. Pacific Northwest ferns and their allies. University of Toronto Press, 
Toronto. 

Waener, W.H.,Jk.and D.M. JoHNsoN. 1981. Natural history of the ebony spleenwort, Asplenium 
platyneuron (Aspleniaceae), in the Great Lakes area. Canad. Field-Naturalist 95:1 56-166. 

Wacner, W.H., Jr.and F.S.Wacner. 1986.Three new species of moonworts (Botrychium subg. 
Botrychium) endemic in western North America. Amer. Fern J. 76:33-47. 

Winter, J.M. 1936. An analysis of the flowering plants of Nebraska. Bull. 13:1-203, n.s.10, 
Conservation and Survey Division, University of Nebraska-Lincoln. 

Weeser, HJ. 1890. A catalogue of the flora of Nebraska. Nebraska State Board Agric. Rep. 
1889:177-302. 


— 


EXPANSION OF THE EXOTIC AQUATIC PLANT 
CRYPTOCORYNE BECKETTII (ARACEAE) IN THE 
SAN MARCOS RIVER, TEXAS 


Robert D. a 
niversity of North Te 
a ely pe Institute ee Sciences 
PO Box 310559, Denton, TX 76203-0559, U.S.A. 


ABSTRACT 


Cryptocoryne beckettii Thw. ex R. Trim. is an exotic aquatic plant recently found in the San Marcos 
River, Texas. The species is currently expanding rapidly within the lower portions of the upper San 
Marcos River. The distribution and areal extent of the species was quantified on = occasions be- 
tween April 1998 and coas 2000. ee this 28- month period, the number of 
increased pos nee 63, 1 f 171 to 646 m*. The average rate of 
areal expa ring this mele was 80% per ae Most colonies of C. beckettii were found to be 
small e 5 m?), 2), although i in August 2000 three colonies were greater than 50 m’ in size. All colonies 
were found at water eepus pees? oe ane 120 cm and appeared to favor more rapidly flowing 


} 


f the river makes C. beckettii a potentially 

nt endemic to the San Marcos River, that 
occupies a similar river zone. All known colonies of G ne are currently downstream from the 
remaining stands of Z. texan 


Lf 
oF; . . 1 


a 
serious threat to t Hitche 


RESUMEN 
] sti be oenoontradarecient t 
Cryptocoryne beckettii Thw.ex R. ue I 
ee rio San saab Texas. | | pandiénd Apid por las 5 baj 
1S Maeve La distribucion y area d idn de | ie f tificad 


tres ocasiones entre abril de 1998 y agosto de 2000 Durante este ponds de 28 meses, en numero de 
colonias individuales se incremento de 11 a 63, y el area total 171 


a 646 m?. La tasa media de expansion del area durante este periodo fue del 80% por ano. La mayoria 
de las es de C. beckettii son ‘Pequefias s 5 m?), aundae en naga de 2000 tres colonias eran 


mayores de 50 m’. Todas las col en p g e 30 y L2Ocmy 
parecian ae un flujo de agua mas rapido. Esta preferencia por las areas del rio poco ana 
yon sae hace de C. peceert I I tchc., una 

d io San Marcos, q fluvial similar. Todas colonias 


C 
Greens de C. beckettii estan actualmente aguas abajo de las posiciones restantes de Z. texana 


INTRODUCTION 


The San Marcos River, Hays County, Texas, originates from a series of springs 
along the San Marcos Springs fault within the city of San Marcos, Texas. The 
river flows 130 km to its confluence with the Guadalupe River near the city of 
Gonzales. The upper river, defined as the eight km between its origin and its 


'Current Address: Baylor University, Department of Biology, PO Box 97388, Waco, TX 76798, U.S.A. 
SIDA 19(4): 1027 — 1038. 2001 


1028 BRIT.ORG/SIDA 19(4) 


confluence with the Blanco River, is the subject of substantial interest due to 
the presence of several endangered species (USFWS 1996). The river maintains 
relatively constant temperature and water chemistry year around, and this con- 
stancy may be responsible for the biological uniqueness of the system (Hannan 
S Dorris 1970; Lemke 1989; Groeger et al. 1997). This river supports the greatest 
known diversity of aquatic organisms in Texas, and several of the more rare 
species are limited to the upper portion of the river (Staton 1992; USFWS 1996). 
Considered an ecologically sensitive and critical habitat by the US Fish and 
Wildlife Service (USFWS 1996), the upper river harbors several endangered 
aquatic species, including Zizania texana Hitche. (Texas wild rice), Iyphlomolge 
rathbuni Stejner. (San Marcos blind salamander), and Etheostoma fonticola Jordan 
& Gilbert. (fountain darter), and the threatened Eurycea nana Bishop. (San 
Marcos salamander). Gam busia georgei Hubbs & Penden. (San Marcos gambusia) 
isa fish species thought to have recently gone extinct from the San Marcos River. 

The river's spring-fed waters provide excellent habitat for the luxuriant 
growth of numerous common aquatic plant species as well. Lemke (1989) 
reported the presence of 27 species of aquatic flowering plants as well as two 
species of aquatic ferns and two aquatic bryophytes. Unfortunately, eight of 
these species are exotic to the United States including some that are known to 
spread aggressively. The most abundant plant found in the upper river is now 
the notorious Hydrilla verticillata (L£.) Royle,a non-native plant know to cause 
substantial management and ecological problems (see Langeland 1996). An- 
other introduced aquatic plant species here is Hygrophila polysperma (Roxb) T. 
Anderson, native of India and Malaysia. Angerstein and Lemke (1994) hypoth- 
esized that the high growth potential of H. polyspermaalso posesa serious threat 
to the native flora and the biotic integrity of the San Marcos ecosystem. Al- 
though there is little historic information on the distribution of H. polysperma 
during the 25 or so years it has been in the system, it appears to have expanded 
dramatically during the last 15 years. In his survey of the upper river, Lemke 
(1989) categorized H. polysperma (although misidentified as Hygrophila 
lacustris (Schlecht. & Cham.) Nees as “uncommon” which he defined as “re- 
stricted to one or a few locations in the study area.” Today, this species is wide- 
spread in the upper river and second only to H. verticillata in abundance (Doyle, 
unpublished data). Hygrophyla polysperma appears able to compete with H. 
verticillata (Les and Wunderling 1981) and apparently can easily outcompete 
Ludwigia repens Forst.,a native aquatic macrophyte which hasa similar growth 
form (Francis 2000). One other exotic aquatic plant species know to inflict dam- 
age on native flora is Myriophyllum spicatum L. (see Madsen et al. 1991) a spe- 
cies found abundantly in Spring Lake at the headwaters of the San Marcos River, 
although only rarely in the river itself. 

Recently, a new exotic aquatic plant species identified as Cryptocoryne 
beckettii Thw.ex R.Trim. was found in the San Marcos River (Rosen 2000). The 


DOYLE, TEXAS 1029 


identification provided by Rosen (2000) was based only on vegetative charac- 
teristics, but more recent observations of floral characteristics from a sample 
collected from the river and grown in quarantine culture appear to confirm 
this species identification (Ken Saunders, Texas Parks and Wildlife Department 
(TPWD), personal communication). In this paper, | report information on the 
distribution and recent expansion of this species within the San Marcos River 
based on three vegetation assessments conducted between April 1998 and 
August 2000. 


METHODS 


The distribution and abundance of C. beckettii in the San Marcos were assessed 
in 1998, 1999, and 2000 as part of a larger vegetation assessment of the flora of 
the entire river. The specific survey dates (with river flow rates for those dates 
in parentheses) were: April 02, 1998 (6.20 cms); August 27, 1999 (3.88 cms); and 
August 25, 2000 (3.42 cms). 

For each colony seen, its geographic location and the colony size were re- 
corded using a high-resolution GPS unit (Irimble Pro XR), which was set to 
record data only where spatial resolution was considered “excellent” (+70- to 
100-cm point resolution). Colonies larger than approximately 16 m2 were 
mapped by circumnavigating each colony with the GPS unit. Water depth was 
also measured, using a standard depth pole depth gauge. For the larger colo- 
nies, the depth recorded was the depth considered to be most representative of 
the overall colony. Smaller colonies were mapped by recording a GPS point in 
the geographic center of a colony and recording North/South and East/West 
dimensions. Water depth was recorded where the GPS point was taken. In sec- 
tions of the river where water was deeper and without a clear view of the bot- 
tom (> 17 m),1 searched for C. beckettii by dragging a heavily weighted plant 
rake attached toa rope. Although not quantitative, this sampling technique is 
effective in determining presence/absence of submersed aquatic plants. 
Cryptocoryne beckettii was not found in any of the deeper portions of the river. 

In 1998 and again in 2000 vegetative voucher specimens were collected 
from the river. | have not observed the species in flower in the river. 

Flow velocities were taken in March 200] at two of the larger colonies lo- 
cated near the upper end of Reach 12, using a Marsh-McBirney (Flo-Mate Model 
2000). The measurements were taken just above the plant canopy (ca. 20 cm off 
the bottom). 


RESULES 


Location of colonies, August 2000 

Cryptocoryne beckettii has only been observed in the lower portions of the 
upper San Marcos River (Fig. 1). In August 2000, the most upstream colony was 
located just below the “power line.” That location, named for the high-voltage 


1030 BRIT.ORG/SIDA 19(4 


San Marcos, River, TX 
USFWS Reach 12 


~ Spring Lake | 100.0 100 200 Meters 
J S| 
N pr San Marcos wastewater 
0 i aiieiers treatment plant outfall 
“Power Line” | 7% 


gQINN d 


f = i 
| Interstate 35 | 
| for Zizania texana 


/ ca | Initial colony? - , 

Lf yf ae 

! va USFWS Reach 12 a | 
a we Confluence with the 

; | Blanco River | 
8/00 upstream boundary 


USFWS Reach 13 


Fic. 1. Distributi f Crypt beckettiiin the San M River, Hays County, Texas, U.S.A. on August 25, 2000 


power line that crosses the river, is the beginning of the USFWS monitoring 
Reach 12. This colony was located about 20 m downstream of the southern- 
most stand of Z. texana, which is just upstream of the power line crossing. 
Cryptocoryne beckettii was widely distributed throughout Reach 12 and was 
the dominant aquatic macrophyte in that Reach (Table 1). Unlike most of the 
river, Reaches 12 and [are largely shaded by riparian tree canopy and are very 
sparsely populated by aquatic macrophytes. Total coverage by aquatic macro- 
phytes in these sections is only about 3.6% of the total Reach area, and C. 
beckettii accounted for 73% of all aquatic vegetation coverage in that area. 
Only four tiny colonies of C. beckettii totaling 1.0 m2 were found in Reach 13. 
These were located just downstream of the division between Reaches 12 and 13. 


Water depth and size of colonies, August 2000 

In August 2000, most C. beckettii colonies were located at depths of 30 to 90 
cm (Fig. 2). No colonies were located in water shallower than 30 cm nor in wa- 
ter deeper than 120 cm. Although 27% of all colonies were located in water 
deeper than 90 cm, these colonies tended to be small, so the areal coverage in 
deeper water was only about 13% of the total areal coverage. Because the Au- 
gust 2000 survey was conducted at relatively low river flow (3.42 cms), I be- 
lieve all colonies surveyed remain completely submersed throughout the year. 


DOYLE, CRYF TEXAS 1031 


Taste 1. Abundance of all aquatic plant species in USFWS Reaches 12 and 13 of the San Marcos 
River, TX, in Oct 2000. Species are identified as exotic (E) or native (N). 


Species Total Area (m2) Number of colonies 
REACH 12 
ee PeCe (E) 644.5 59 
a* (N) 79 
Hydrilla verticillata (E) 11.3 8 
Hygrophila polysperma (E) 197.9 29 
lusticia americana (N) 43 3 
Nuphar luteum (N) 5.5 4 
ag! ttaria platyphylla a (N) <0.1 ] 
ria americana N 9.5 2 
Zosterella dubia* (N) 79 9 
REACH 13 
Cryptocoryne beckettii (E) 1.0 
Hygrophila polysperma (E) 7.25 12 
Nuphar luteum (N) 424 7 
*Zosterella dubia = Heteranthera dubia = Het thera lieb 


Most C. beckettii colonies in the San Marcos River were small in August 
2000. Of the 63 discreet colonies identified, 42 were <5 m2. At the same time 
48% of the total population occurred in only three colonies, which ranged in 
size from 53 to 160 m¢*. 


Expansion of colonies over three years, 1998-2000 

In all three surveys, the majority of all colonies were less than 5 m? in size, and 
relatively few colonies were greater than 10 m? (Fig. 3A). However, between 1998- 
2000 the absolute number of colonies in each size class increased (Fig. 3B) indi- 
cating that the population continued to expand and that individual colonies 
were continuing to grow in size. 

The number of colonies and the area covered by C. beckettii increased sig- 
nificantly during the survey period (Fig. 4). In April 1998 there were only ll 
colonies of C. beckettii, and 68% of the total areal coverage was found in one 
large colony located in the shallows on the inside of a bend in the river just 
above the wastewater treatment plant (Fig. 1). The total number of colonies be- 
tween April 1998 and August 2000 increased from 1] to 63 (Fig. 4A), while total 
areal coverage of the species increased from 171 m? to 646 m? (Fig. 4B). The rate 
of increase in areal coverage averaged about 80% per year. 


Flow velocity over Cryptocoryne beckettii colonies 

In March 2001, the flow over two large C. beckettii colonies was 0.56 and 0.75 m 
s!. These colonies were located near the upstream limits of the population, but 
appeared visually similar to most of those observed within the river. 


1032 BRIT.ORG/SIDA 19(4) 


wv”) 
e |A 

Cc 

2 

© 

e. oo 

.@) 

Soe 

oO 

O 

E 

= 20 + 

© 

~— 

ie) 

et 

ee 

OC 10- 

~~ 

Cc 

ce) 

O 

o no colonies 
O. at these depths 
. |B 

0) 

40) 

— 

g 30 

(@) 

O 

O 

<0) 

w 

mew 20 

w 

et 

oO 

~~ 

we 

eo) 

—_— 

Cc 104 

oO 

O 

Seen 

oO 

Oo. colonies 

0 at these depths 
<30 30-50 51-70 71-90 >90 
Colony Depth (cm) 

Fig y) Nisesth at f Cry Lb L he iT 4 I L {AVC 1 y £ | . iheda ed lh (B) 
Frequency of percent of | | t different depths. M I August 25, 2000 in the San 


Marcos River, Hays County, Texas, U.S.A. 


DOYLE, CRYE TEXAS 1033 


100 


80 


Pe Bg Apr 1998 
le 7 zzzza Aug 1999 
mmm AUG 2000 


Percent of total number of colonies 


“) 
oO 
c 3074 
© 
(e) 
oO 
Ke) 
$e 20 4 
O 
_ 
Oo 
ke 

10 + 

sam |” 
0-5 5-10 10-25 25-50 >50 
Colony area (m*) 
1c. 3. Distributi fC beckettii colonies in the San M Ri Texas by size classes (A) P t of col 


ff ra 
nies in each size class. (B) Number of colonies in each size class. 


1034 BRIT.ORG/SIDA 19(4) 


50 + 


Total Number of Colonies 


500 - 


Total areal coverage (m*) 


Apr 2, 1998 Aug 27, 1999 Aug 25, 2000 


Survey Date 


Fic.4 NI n f{-n] * (A) 4 n 1 (B) £ ; 4 n 1 vee Lac aa n:. Texas, 1998— 


DOYLE, TEXAS 1035 


DISCUSSION 


Many introduced aquatic plant species have spread beyond their native ranges, 
and some problematic weeds h { in spectacular fashion in their new 
habitat (Cook 1990). These alien species may interact with the native flora in 
various ways (Falinski 1998), such as filling a long-empty ecological niche in 
the community (supplementary interaction), filling a recently vacated niche 
(compensatory), or displacing one or more native species with similar biologi- 
cal and ecological requirements from a filled niche (substitutive). Although 
reliable criteria to determine accurately the ultimate weed potential of new in- 
vaders have not been developed (Mack 1996; Zamora et al. 1989), species that 
are likely to interact with native flora in negative ways should be considered to 
have high noxious weed potential (Bazzaz 1986). 


Expansion of Cryptocoryne beckettii 

During the survey period, C. beckettii increased at an alarmingly rapid rate. 
Between April 1998 and August 1999, C. beckettii areal coverage expanded at 
an annual rate of 80.2%. Between August 1999 and August 2000, the areal cov- 
erage of the population increased by 82.5%. Although the rate of expansion is 
likely to slow as the most suitable habitats become colonized, at the current 
average rate of expansion (80% per year) C. beckettii could cover 100% of Reach 
12 in less than five years. 


Potential threat to Zizania texana 
The morphology of Z.texanaand C. bekettii differ substantially. Zizaniatexana 
is characterized by long, ribbon-like leaves often observed to be 1-2 m in length 
(Terrell et al. 1978) while the ovate leaves of C. beckettii are typically only 15- 
30 cm in length, including both petiole and blade. Even so, | believe the intro- 
duction of C. beckettii into the San Marcos ecosystem may pose a substantial 
threat to Z. texana, because the two species appear to have similar depth and 
flow preferences. Poole and Bowles (1999) surveyed 44 individual stands of Z. 
texana in August 1994 and found that the water depth for these stands aver- 
aged 0.75+0.16 m (95% c.i.). In August 2000, the 63 surveyed colonies of C. 
beckettii were in an average water depth of 0.72+0.07 m (95% ci), indicating 
that the two species occupy virtually identical depth zones within the river. 
Furthermore, both species appear to favor locations with relatively high cur- 
rent velocity. Poole and Bowles (1999) further reported that the 44 stands of Z. 
texana had an average current velocity of 0.56 (+0.20) ms! (95% c.i.). Although 
current velocity near C. beckettii was measured only once and at only two colo- 
nies, the observed velocity range of 0.56 to 0.75 m s™ indicates that this exotic 
species occupies a flow environment very similar to that of Z. texana. 

One major habitat difference between the two species may indicate a dif- 
ference in the degree of shading that can be tolerated. Zizania texana is most 
frequently found in full sun, while, at present, C. beckettii is found predomi- 


1036 BRIT.ORG/SIDA 19(4) 


nantly in more heavily shaded regions of the river. However, historical, unpub- 
lished data from TPWD indicate that within the past five years there have been 
stands of Z. texana extending farther downstream than the current distribu- 
tion into sections of the river now dominated by C. beckettii. 

The fact that C. beckettii may have similar habitat preferences to Z. texana 
does not indicate that it can or will displace remaining stands of the native plant. 
In fact, it is possible that established Z. texana stands, like other established 
native species such as Vallisneria americana Michx. will be a very effective 
competitor against an invading alien species (see Smart et al. 1994). However, 
given that the distribution of Z. texana is currently much reduced from historic 
levels (USFWS 1996), an aggressively expanding alien species suchas C. beckettii 
may quickly occupy habitat that might otherwise be re-colonized by Z. texana. 


Management recommendation 

Eradication is ultimately the most desirable response to a new plant invasion, 
especially when it appears likely to interfere with an important native species. 
However, this outcome is most likely when plant populations are relatively small 
and contained; large or widely distributed populations require considerably 
more knowledge, money, and effort to achieve eradication (Coblentz 1990; 
Zamora & Thill 1999). The current abundance and distributional pattern of C. 
beckettii appears to lend itself to implementing an eradication effort. In Au- 
gust 2000, the population of C. beckettii was relatively small (646 m2) and lim- 
ited toa 1.7 km stretch of the upper San Marcos River. All existing stands of Z. 
texana are located upriver of this area, and very few other native species are 
present within the affected reaches. However, the explosive increase in C. 
beckettii observed to date indicates that the situation is likely to quickly be- 


come much more difficult to control. 
ea | | 


Should control actions be 


,a holistic, ecosystem-based approach 
should be utilized. Control plans that focus on a single alien species without 
addressing the underlying disturbance phenomena that permitted the success- 
ful invasion, deal only with the effect of environmental degradation and not 
the causes (Edwards 1998). Single species approaches may simply delay an in- 
evitable permanent establishment of the alien in the region or simply trade one 
invasive species for another equally damaging one (Hobbs and Humphries 
1995). In the San Marcos River, C. beckettii has invaded a portion of the river 
that was virtually empty of aquatic vegetation. Given the apparent suitability 
of this species for this particular area and the apparent popularity of the spe- 
cies within the aquarium trade (illustrated by the large number of sites on the 
World Wide Web devoted to this genus), the likelihood of re-introduction is 
high. At a minimum, eradication efforts should be followed by an aggressive 
restoration effort to fill the “empty niche” with more desirable, native vegeta- 
tion such as Zizania texana and Vallisneria americana. 


DOYLE, CRYE TEXAS 1037 


ACKNOWLEDGMENTS 


I wish to thank Matt Francis and Joel Johnson (US Army Corps of Engineers, 
Lewisville Aquatic Ecosystem Research Facility) as well as Tyson Galusky (Uni- 
versity of North Texas) for assistance in the field. Ken Saunders (TPWD) pro- 
vided valuable information on his observations related to Cryptocoryne 
beckettii. This research was supported by a grant from the US Fish and Wild- 
life Service. 


REFERENCES 


Ancerstein, M.B.and D.E. Lemke. 1994. First records of the aquatic weed Hygrophila polysperma 
(Acanthaceae) from Texas. Sida 16:365-371. 

Bazzaz, F.A.1986.Life histories of colonizing plants:some d hic, genetic,and physi- 
ological features. In: H.A. Mooney and J.A. Drake, eds. Sadly afbiclogicsl invasions of 
North America and Hawaii. Springer-Verlag, New York. Pp. 96-110. 

Copstentz, B.E. 1990. Exotic organisms:a dilemma for conservation biologists. Biol. Conserv. 
4:261-265 

Cook, C.D.K. 1990. Origin, autecology, and spread of some of the world’s most troublesome 
aquatic weeds. In: A.H. Pieterse and K.J. Murphy, eds. Aquatic weeds: the ecology and 

management of nuisance aquatic vegetation. Oxford University Press, New York. Pp. 

31-38 


Eowaros, K.R. 1998. A critique of the general approach to invasive plant species. |n:U. Starfinger, 
K. Edwards, |. Kowarik, and M. Williamson, eds. Plant invasions: ecological mechanisms 
and human responses, Backhuys Publishers, Leiden, The Netherlands. Pp. 85-94. 

FauiNski, J.B. 1998. Invasive alien plants and vegetation dynamics. In: U. Starfinger, K Ed- 
wards, |. Kowarik, and M. Williamson, eds. Plant invasions: ecological mechanisms and 
human responses. Backhuys Publishers, Leiden, The Netherlands. Pp. 3-21. 

Francis, M.D. 2000. Interspecific competition between Hygrophila polysperma and Ludwigia 
repens, two species of importance in the Comal River, Texas. M.S. thesis, University of 
North Texas, Denton. 

Groecer, A.W., PF. Brown, T.E. Tieuen, and T.C. Ketsey. 1997. Water quality of the San Marcos 
River. Texas J. Sci..49:279-294 

HANNAN, H.H. and T.C. Dorris. 1970. Succession of a macrophyte community in a constant 
temperature river. Limnol. Oceanogr. 15:442-453 

Hoses, R.J. and S.E. Humenries. 1995. An integrated approach to the ecology and manage- 
ment of plant invasions. Conserv. Biol. 9:761—770 

LANGELAND, K.A. 1996. Hydrilla verticillata (Lf) Royle (Hydrocharitaceae), the perfect aquatic 
weed. Castanea 61:293-304. 

Lemke, D.E. 1989. Aquatic macrophytes of the upper San Marcos River, Hays Co., Texas. 
Southw. Naturalist 34:289-291 

Les, DH. and R.P. Wunbertin. 1981. Hygrophila polysperma (Acanthaceae) in Florida. Florida 
Sci. 44:189-192. 


1038 BRIT.ORG/SIDA 19(4) 


Mack, R.N. 1996, Predicting the identity and fate of plant invaders; emergent and emerg- 
ing approaches. Biol. Conserv. 78:51-58. 

Maosen, J.D., J.W. SUTHERLAND, J.A. Bloomrieto, L.W. EicHter, and C.W. Boyten. 1991. The decline of 
native vegetation under dense Eurasian watermilfoil canopies. J. Aquat. Plant Manage. 


Poote, J.and D.E. Bowtes. 1999. Habitat characterization of Texas wild-rice (Zizania texana 
Hitchcock), an endangered aquatic macrophyte from the San Marcos River, TX, USA. 
Aquat. Conserv. Mar. Freshw. Ecosyst. 9:291—302. 

Rosen, D.J. 2000. Cryptocoryne beckettii (Araceae), a new aquatic plant in Texas. Sida 19: 
399-401. 

Smart, R.M., J.W. Barko, and D.G. McFarLaNb. 1994. Competition between Hydrilla verticillata 
and Vallisneria americana under different environmental conditions. Aquatic Plant 
Control Research Program, Technical Report A-94-1. U.S. Army Engineer Waterways 
Experiment Station, Vicksburg, MS. 

Staton L.L. 1992. Assessment of changes in the aquatic macrophyte community in the 
upper San Marcos River. M.S. thesis. Southwest Texas State University, San Marcos. 
Tere et, E.E.,W.H.P. Emery, and H.E. Beary. 1978. Observations on Zizania texana (Texas wildrice) 

an endangered species. Bull. Torrey Bot. Club 105:50-57. 

USFWS 1996. San Marcos/Comal species recovery plan (revised). United States Fish and 
Wildlife Service, Albuquerque, NM. 

Zamora, D.L.and D.C. THitt. 1999. Early detection and eradiction of new weed infestations. 
In:R.L.Sheley and J.K.Petroff eds. Biology and management of noxious rangeland weeds. 
Oregon State University Press, Corvallis. Pp. 73-84. 

Zamora, D.L., D.C. THitt, and R.E. Eptee. 1989. An eradication plan for plant invasions. Weed 
Technol. 3:2-12. 


VASCULAR PLANT TYPES IN THE 
ARIZONA STATE UNIVERSITY HERBARIUM 


Stefanie M.lckert-Bond and Donald J. Pinkava 


Department of Plant Biology 
Arizona State University, Box 871601 
Tempe, AZ 85287-1601, U.S.A. 


ABSTRACT 


Founded in 1896 by Fredrick M. Irish, the vascular plant herbarium at Arizona oe ome (ASU) 
acts asa EEpOSHOry for plants particularly from the desert Southwest and Mexico. It now contains 
of which 735 have been designated as types incling paratypes. Here we 
present a list that abouts 384 major vascular plant type specimens and includes verified holo 
types, lestovypes, syntypes and neetypes and their oh aida Regionally ie majority of the speci- 
1 Mexico (178), w 13 from South America and 7 from the 


Old World. anaes the majority of type specimens res o the Asteraceae (84), followed by 
Polygonaceae (56), Cactaceae (34), Scr tophusanacese (27), Habart: (16), the Agavaceae (13), and 


Acanthaceae (10). Historically ¢ i | by CG. Engl a Blom: 
nent early plant collector in Mexico from 1885 to 1909.N L] b 

James R. ee Noel H. Holmgren, Howard S y and George B. nice For each basionym typi- 
fied by an ASU specimen, whic ch is accompanied by a copy of the original publication, we here list 


by 
collector(s) a collection number, locality information (restricted to country and state or equiva- 
lent demographic unit), and date of collection. 


RESUMEN 


Fundado en 1896 por Frederick M. Irish, el herbariod l le la Universidad del Estado 
de Arizona (ASU) actta como un Sia - plantas eigen del desierto del Suroeste y 
México. Actualmente el h 1 jemplares, de los cuales 735 corresponden a 
ejemplares tipo ae ae psi Se presenta una lista que ee 384 de los mas 
importantes ejemplares tipo de plantas vasculares, incluyendo holotipos, lectotipos, sintipos, y 
eon pes, ye sus Sau teades: Reponaien la one a nee ejemplares son de los Estados Unidos 
Vi ie Mundo. Taxonémicamente, 
oe raya ade s familias A 84), seguida de Polygonaceae 
S 6), sCactacene GH), perpinlaaceae Co Fabaceae te) Agavaceae (13), y Acanthaceae (10). 

I p oc ean prominente Sones 
en México entre 1885 y | 1909 S ta adema j ] i James R eal, 
Noel H. Holmgren, Howard S. Gentry y George B. incon eur Be cae tipificado por un eee 
en ASU, incluye una copia de la publicaci6n original. Se listan Sua . os y numero de 


colecta, informacion de la localidad (restringida al pais y estado o mografica equival 


y a la fecha de la colect 
INTRODUCTION 


Since the herbarium was founded by Fredrick M. Irish in 1896, the vascular plant 
herbarium has grown to over 230,000 specimens, centering on floristics of the 


SIDA 19(4): 1039 — 1059. 2001 


1040 BRIT.ORG/SIDA 19(4) 


desert Southwest and Mexico, but many worldwide collections are represented 
Frederick Mortimer Irish received his Bachelor of Science degree at the Univer- 
sity of lowa in 1895 and briefly taught high school in his hometown of Dubuque, 
lowa, before accepting the position at Tempe Normal School in 1896 (Thomas 
1960). “Cap” Irish served not only as the biology science teacher, but also as foot- 
ball coach, captain of the Military Company, and leader of student activities 
such as the Cactus Walking Club and the Girl’s Hiking Club (Hopkins & Tho- 
mas 1960; Trauth-Nare & Pinkava 2000). 

“He knew the flowers and could call them by name. Asa botanist he was 
an authority” (Thomas 1960, p. 549). Irish was instrumental in establishing the 
herbarium. When Irish came to Tempe he brought with him his personal col- 
lections. Over the years more than 700 specimens bearing his name have been 
processed; some duplicates have been eventually deposited at the Desert Bo- 
tanical Garden (DES). His early collections in the Salt River Valley are a valu- 
able historical record of the diversity of plants in the Phoenix area of the time. 
Irish was appointed Registrar at the newly named Arizona State Teachers Col- 
lege in 1925 and served in this position till his death in 1941 (Thomas 1960). 

James A. McCleary followed Irish as curator and botany professor. His in- 
terests were in bryophytes and desert plants and he and his students added 
approximately 3,000 specimens from 1947-1959 with only minimal financial 
assistance (McCleary, pers. comm.). By 1960, according to the new curator, Nor- 
man H. Russell, professor and departmental chair, more than 15,000 specimens 
were shelved in the herbarium. It was housed on the third floor of the now Life 
Science Center A Wing, Arizona State University (Rubinoff 1973). Russell's re- 
search interests included study of Viola and numerical taxonomy. 

Donald J. Pinkava became director of the herbarium and Assistant Profes- 
sor of Botany in 1964, after receiving a Ph.D. in Botany from Ohio State Univer- 
sity, studying the genus Berlandiera DC. (Asteraceae). His research has focused 
on the Asteraceae, floristics of Cuatro Ciénegas, Coahuila, Mexico (resulting in 
52 type collections and 5 original descriptions) and most recently on cacti, par- 
ticularly the Opuntioideae of the Southwest. The ASU Herbarium houses one 
of the nation’s largest collections of Cactaceae, including the best documented 
vouchered collection for chromosome counts. The Arizona State University 
Herbarium received official recognition when assigned its acronym, ASU, in 
1966 (Anonymous 1966). It was then recognized as one of only 105 resource 
herbaria in the United States (Anonymous 1974). In 1971 the herbarium was 
moved to its present site, the third floor of the new C Wing of the Life Science 
Center (LS C392). During Pinkava’s tenure as director, several members of the 
faculty, curators and especially his graduate students were responsible for the 
growth of the herbarium. 

Throughout the 1960s and 1970s Pinkava received much assistance from 


TLACATL HEADARIUM 1041 


the very knowledgeable curator, Ms. Elinor Lehto; her contributions were im- 
measurable. Upon her retirement in 1980, Bruce Parfitt served as acting curator, 
until the position was upgraded to half-time curation and half-time research 
position, which was filled by Thomas Daniel, who received his Ph.D. from the 
University of Michigan. As an expert on Neotropical Acanthaceae, his studies 
resulted in several types. Upon the appointment of Tom Daniel as curator of 
the herbarium at the California Academy of Sciences, Leslie R. Landrum, who 
also received his Ph.D. from the University of Michigan, accepted in 1986 the 
position as curator and research scientist in the Department of Botany. He is an 
expert on New World Myrtaceae. An herbarium grant from the National Sci- 
ence Foundation created a permanent half-time position for a collections’ man- 
ager, which rotates from student to student and provides an important learn- 
ing experience. As curator, Landrum’s objectives have been to enlarge the 
holdings of Southwestern plants, and acquire material from Mexico, South 
America and the Caribbean, where he and many of his students carry out cur- 
rent studies. Increased interest has also been placed upon the Phoenix Flora 
project (URL: http://wwwasu.edu/ces/CAPLTER.htm), part of the Central Ari- 
zona- Phoenix Long-Term Ecological Research Project (CAP LTER) awarded to 
the Center for Environmental Studies, funded by the National Science Founda- 
tion. Present staff of the ASU herbarium includes Pinkava, retired Director 
(Biosytematics of Opuntioideae, Southwest floristics), and Landrum, Curator 
(Systematics of South American Myrtinae, Southwest floristics). 

The ASU herbarium serves as repository of specimens worldwide and par- 
ticularly from the southwestern United States. Emphasis is on producing a 
Manual of the Vascular Plants of Arizona, portions of which are being pub- 
lished as they become available in the Journal of the Arizona-Nevada Acad- 
emy of Science. From 1992 to 2001, treatments of 68 families or portions thereof, 
were prepared by experts, including ASU staff and students (Mason et al. 1992). 
These publications make the treatments available for researchers, pending the 
publication of the completed Flora. 


TYPE COLLECTIONS 


The ASU vascular plant herbarium houses 735 type specimens including 26 
holotypes, 338 isotypes, 3 isolectotypes, 13 isosyntypes, 4+ isoneotypes, 233 
paratypes, and 118 isoparatypes. The following list documents 384 vascular 
plant type specimens in the Arizona State University Herbarium (ASU) and 
includes verified holotypes, lectotypes, syntypes and neotypes and their dupli- 
cates. Regionally, the majority of the specimens are from the United States (186) 
and Mexico (178), while 13 are from South America and 7 from the Old World. 
Taxonomically the majority of type specimens belong to the Asteraceae (84), 
followed by Polygonaceae (56), Cactaceae (34 of which 15 are holotypes), 


1042 BRIT.ORG/SIDA 19(4) 


Scrophulariaceae (27), Fabaceae (16), Agavaceae (13), and Acanthaceae (10). 
Historically prominent are 60 isotypes collected by CG. Pringle, a prominent 
early plant collector in Mexico from 1885 to 1909 (Davis 1936). Numerous type 
collections were also made by James R. Reveal, Noel H. Holmgren, Howard S. 
Gentry, and George B. Hinton (ckert-Bond and Pinkava 2000). The list of type 
specimens is arranged phylogenetically by division and then alphabetically 
by family, genus, species, and infraspecific categories. For each basionym typi- 
fied by an ASU specimen accompanied by a copy of the original publication, 
we here list collector(s) and collection number, locality information (restricted 
to country and state or equivalent demographic unit), and date of collection. 

In the 1960s the irreplaceable type specimens were filed among the general 
collection and were thus subject to much unnecessary handling. The specimens 
are now housed in two separate herbarium cabinets. The use of these specimens 
is limited to only trained taxonomists, who appreciate the value of these speci- 
mens in accordance with the recommendations set forth by the International 
Code of Botanical Nomenclature (Greuter et al. 2000), “Recommendation 7A . 
It is strongly recommended that the material on which the name of a taxon is 
based, especially the holotype, be deposited ina public herbarium or other pub- 
lic collection with a policy of giving bona fide botanists open access to depos- 
ited material, and that it be scrupulously conserved “. The types are specially 
curated in acid-free genus folders (e.g., palm folders) and additionally each speci- 
men is protected by a species folder, all from Herbarium Supply Company, Cali- 
fornia. Copies of original descriptions are accompanying the type specimens. 
They are fastened on a herbarium sheet with archival clear-hold press-on 
mounting corners from Light Impressions, New York. Specimens checked 
against the original publication and the type status was verified. Cross-refer- 
encing dummy sheets are placed in the main herbarium. 

Some specimens that are cited in the protologue are missing from the ASU 
type collection; they are out on loan or were never received. This type list is the 
first account of the history of the ASU herbarium and the first list of its type 
specimens. We plan to update this list in the future with a supplement of all 
new types since this publication. We feel that the printed version of the type 
list allows for convenient checking of type specimens. A searchable list of all 
type specimens including paratypes, isotypes and clonotypes as well as some 
images of types is planned for the future, and will be made available at a later 
point through the web page of the ASU herbarium (URL: http://Isvl.la.asu.edu/ 
herbarium/). The construction of sucha web page will take some time and this 
paper version will serve in the interim. 


LAER 


1043 


HACADARIVM 


VASCULAR PLANT TYPES IN THE 
ARIZONA STATE UNIVERSITY HERBARIUM 


POLYPODIOPHYTA 


foc ant ae 
lenium montanum CL. Willdenow forma 
shawangunkense T. Reeves, Amer. Fern J. 
64:105. ee USA. New York: 18 Oct 1973, T. 
TYPE), ASU 67981. 


Pp 
MNECVES AIYY (NUL 


DRYOPTERIDACEAE 

Cystopteris utahensis M.D. Windham & C.H 
Haufler, Amer. Ly 81:13.1991.USA. Utah: 
2 Jul 1990, M.D. Windham 90-282 with 
Windham ie ASU not found. 

Woodsia cochisensis M.D. Windham, Contr. 
Univ. Michigan Herb. 19:54. 1993. USA. Ari- 
zona: 31 Aug 1985, M.D. Windham 781 with 
G. Yatskievych tele ta not found. 

Woodsia neom Windham, Contr. 
aera ca oe 1993. USA. New 
Mexico: 23 Aug 1990, M.D. Windham 90-365 
with Rabe (ISOTYPE). ASU not found. 


POLYPODIACEAE 

Goniophlebium pringlei W.R. Maxon, Proc. U.S. 
Natl. Mus. 27:953. 1904. Mexico. Veracruz: 30 

11855 (iSOTYPE). ASU 


Nov 1903, C.G. Pringle 
60335. 


PTERIDACEAE 
Astrolepis cochisensis (L.N. Goodding) D.M. 
m subsp. arizonica D.M. Benham, 
Amer. Fern J.82:60. 1992. USA. Arizona: 5 Sep 
1989, D.M. Benham 1312 (IsoTYPE). ASU 
158448. 

Cheilanthes yavapensis |. Reeves ex M.D. 
Windham, Contr. Univ. Michigan Herb. 19:33. 
1993, USA. Arizona: 18 Nov 1980, M.D. 
Windham 202 (isotype). ASU not found. 

Pellaea lyngholmii M.D.Windham, Contr. Univ. 
Michigan Herb. 19:40. 1993. USA. ea 23 
Sep 1990, M.D. Windham 90-420 w 

Lyngholm (isotype). ASU not fou 

Pellaea ternifolia (A.J. Saeed J H.F. Link 
subsp.arizonica M.D.Windham, Contr. Univ. 
Michigan Herb. 19:47. 1993. USA. Arizona: 18 
Mar 1981,M.D.Windham 246-C (ISOTYPE). ASU 
115789. 


MAGNOLIOPHYTINA-DICOTS 


ACANTHACEAE 

Dyschoriste novogaliciana T.F. Daniel, 
Polibotanica 2:1. 1996. Mexico. Nayarit: 16 
Mar 1982, TF. Daniel 2057 (isoTyPE). ASU 
208318. 

Justicia masiaca T.F. Daniel, Brittonia 47:408. 
1995.Mexico ee 1983, TF Daniel 
2456 (iSOTYPE). ASU 220 

Mirandea andradenia IF ae Southw. Natu- 
ralist Se ioe Mexico. Taumalipas: 12 
Aug 1984, TF. pe 3693 with M. Baker 
oe ou 

Mirandea Fenech TF. Daniel, Syst. Bot. 
3:428.1979.Mexico. Nuevo eee 1978 
TF. Daniel 252 (ISOTYPE). ASU 11 

sie eos: S.Acosta, ee 57:249. 

o.Tabasco: oe 1 1983,F Ventura 
ie , Heat 

Tetramerium eae T.F. Daniel, 
Syst. Bot. Monogr. 12:89. 1986. Mexico. 
Michoacan: 27 Nov 1983, I-F Daniel 3267 with 
M. Butterwick (isoTYPE). ASU 1415 

So emilyanum T.F. Daniel, Syst. Bot 

12:74. 1986. Mexico. Michoacan: 26 
eee a TF, Dani o cl with M ae 
(ISOTYPE). ASU 1417 

Tetramerium ae aah TF. Daniel, Syst. Bot. 

nogr. 12:92. 1986. Mexico. Michoacan: 26 
a 1983, LF. re — with M. Butterwick 
(ISOTYPE). ASU 1415 

ae ela redowsi T.F. Daniel, Syst. Bot. 

Mono 


— 


1986. Mexico. Michoacan: 24 
Mar ee A Danie! 2145 (iSOTYPE). ASU 
141586 


Tetramerium sagasteguianum IF. Daniel, Syst. 
nogr. - 10. 1986. Peru. Cajamarca: 
_ LF. Daniel 4319 with M. 

Butterwick an ASU 142681. 


APIACEAE 

Angelica callii M.L. Mathias & L.C. Constance, 
Madrono 24:78. 1977.USA. California: 18 a 
1965, . &V. Call 2459 (isotype). ASU 10872 

Lomatium junceum R. Barneby & ee 
Holmgren, Brittonia 31:96. 1979. USA. Utah: 


1044 


27 May 1978, N.H. Holmgren et al. 8778 
(isoTYPE). ASU 113743. 

Prionosciadum megacarpum J.M. Coulter & 
J.N. Rose, Contr. U.S. Natl. Herb. 3:308. 1895. 
Mexico, pie ~ _ 1894, C.G. Pringle 4688 
pe TYPE). ASU 5 


& B.G. 


Lc 
Baldwin, Brittonia 50:122. 1998. Mexico. Baja 
California: 19 May 1973, R. Moran 20929 
(isoTyPe). ASU 170073. 

AQUIFOLIACEAE 

Ilex oe - Standley, Contr. U.S. Natl. Herb. 

3. Mexico. Puebla: 25 Apr 1904, 
a ee 10003 (isotype). ASU 50602. 


ASCLEPIADACEAE 

Asclepias welshii N.H. Holmgren & PK. 
Holmgren, Brittonia 31:110.1979. USA. Utah: 
20 Jun 1978, N.H. Holmgren 9009 with P 
Holmgren (\soTyPE). ASU 113750 

Marsdenia tressensii S.A. excess iG N.Morillo, 
Bonplandia pea 7:34. 1993. Argen- 


tina.: 21 Oct 1988, S.A. Caceres 426 (ISOTYPE). 
ASU 191309 

ASTERACEAE 

Achaetogeron versicolor J.M.Greenman, Proc. 
Amer. Acad. Arts 41:255. 1905. Mexico. 


Hidalgo: 7 Aug 1904, C.G. Pringle 8849 
(ISOTYPE). ASU 50584. 
Acourtia fragrans J. Rzedowski, Bol. Soc. Bot. 


exico 45:102. 1983. Mexico. Puebla: 6 Jan 
1981, J. Rzedowski 37174 (ISOTYPE). ASU 
184636. 


Acourtia peeees B.L. Turner, Phytologia 
5:404. 3. Mexico. Nuevo Leon: 
1993, oo ca 23934 et al. (ISOTYPE). ASU 

202296 

Acourtia huajuapana BL. Turner var. 
actinomorpha J. Rzedowski, Bol. Soc. Bot. 
Mex. 45:104. 1983. Mexico. Guerrero: 23 Dec 
. Rzedowski 37160 (ISOTYPE). ASU 
1845 

Alomia enol S.F. Blake, J. Wash. Acad. Sci. 
27:375. 7.Mexico. Sonora: 7 Mar 1935,H.S. 
Gentry oe (ISOTYPE). ASU 89075. 

Barroetea subuligera A. Gray var. latisquama 


— 
QJ 


J.M. Greenman, Proc. Amer. Acad. Arts 40:35. 
1904. Mexico. Jalisco: 27 Oct 1903, CG. Prin- 


gle 8773 (\SOTYPE). ASU 17584. 


BRIT.ORG/SIDA 19(4) 


nald, Proc. Amer. Acad. Arts 43:68. 
acan: 1 Nov 1905, C.G. 


stint rosea Schz. Bip. var. aequisquamosa 
M.L 


oo rears Michoa 
Pringle 10109 (ISOTYPE). ASU 50605. 

Carminatia alvarezii J.Rzedowski & G.Calderon, 
Anales Esc. Nac. 7 Biol. 31:9. 1987. Mexico. 
Oaxaca: 25 Oct 1980, J. Rzedowski 37075 
(ISOTYPE). ASU 1 ao 

Carminatia anomala B.L. Turner, Pl. Syst. Evol. 
160:173. 1988. Mexico. Puebla: 6 Oct 1984, 
Sundberg 3032 with M. Levin (isotyPe). ASU 

518. 

Chrysactinia lehtoae D. Keil, Madrono 23:374. 
1976. Mexico. Sinaloa: 25 Nov 1975, £. Lehto 
£19557 with TH. a and J.J. Landye (HOLO- 
TYPE), ASU 8000 

ae pee — B.L. Turner, sie 

4:437. 1988. Mexico. Micho 
i : n 15222 with MLL. one ine TYPE oe 
ASU 12362 52 

a nauseosus (P.S. Pallas) Britt. 
subsp. iridis L.C. Anderson, Great Basin Natu 
ralist: 41:311. 1981. USA. Utah: 29 Sep 1979 
S.L. Welsh 19258 (ISOTYPE). ASU 108397, 

Cirsium clokeyi S.F. Blake, Proc. Biol. Soc. Wash. 
51:8. 1938. USA. Nevada: 6 Aug 1937, LW. 
Clokey 7456 (ISOTYPE). ASU 66713 

Cymophora pringlei B.L. Robinson, Proc. Amer. 
fae Arts 43:39. 1907. Mexico. Gue 22 

905, C.G. Pringle 10068 (ISOT ae 


— 


se 


~w 


Erigeron acomanus R. Spellenberg & P. Knight, 
tenes 36:115.1989.USA.New Mexico: 14 
Jul 1983, PKnight 2689 (isotype). ASU 176290. 
erigeron ieee G.L. Nesom, Phytologia 
243.1990. USA. Arizona: 28 Aug 1981, GL. 

ee 4521 (ISOTYPE). ASU 174639. 


Erigeron ba saseachensl: .L. Nesom, 
Phytologia . 1989. Mexico. Chihuahua: 
18 Aug 1984, 1 aa 5089 with P. Lewis 
(ISOTYPE). ASU 160286. 


Erigeron ne Nesom & B. Hevron, 
Madrono 42:12. 1995. USA. New Mexico: 3 
Jun re : a in with J. Merz (ISOTYPE). 
ASU 1 

ig tae. ee . Sundberg & GL. Nesom, 
Phytologia 69:278. 1990. Mexico. Chihuahua: 
19 May 1985,R. Scott 471 with T. Ayers, M.Lavin 
& A. Whittemore (ISOTYPE). ASU 202297, 


TAREE 


Erigeron chiangii G.L.Nesom var.lamprocaulis 
G.L.Nesom, Phytologia 73:119.1992.Mexico 
Coahuila: 3 Jun 1992, G. Nesom 7408 with M. 
Mayfield (isotype). ASU 190098. 

Erigeron coroniglandifer G_L. aaa 
Phytologia 66:432. 1989.Mexico.Chihuah 
28 oe 1983, G.L. Nesom 4964 (ISOTYPE). ee 
16028 


ae cuatrocienegensis Gl. Soa 
Madrono 28:143. 1981. Mexico. Coahuila: 1 
Jun 1968, DJ. oan see 1 with E. Lehto, os 
Keil (HOLOTYPE). ASU 8 

Erigeron eruptens aL ce om, Phytologia 
66:438. 1989.Mexico.cChihuahua:27 Apr 1985, 
R. Spellenberg 8070 et al. ina tee 143251. 

aes Ae G.L. Nesom, Syst. ce ae 

A. New ce 30 Aug 1981, G.L. 

ae ree (isoTYPE). ASU oe 

Erigeron sceptrifer G.L. Nesom, Phytologia 
69:248. 1990. Mexico. Chihuahua: 23 Aug 
1981, G.L. Nesom 4477 (ISOTYPE). ASU 174638. 

Erigeron scopulinus G.L.Nesom &V.D. Roth, J. 
Arizona-Nevada Acad. Sci. 16:39. 1981. USA. 
Arizona:9 Jun 1976, VD. Roth s.n. (SOTYPE).ASU 


140787. 

Eupatorium saltillense B.L. belay Proc. 
Amer. Acad. Arts 43:31. 1907. Mexic 
Coahuila: 5 Oct — - Pringle 0% 
(ISOTYPE). ASU 1877 

Flaveria mcdougallii : E. Theroux, DJ. Pinkava 
& D. Keil, Madrono 24:13.1977.USA. Arizona: 
27 Jan 1976, M.E. Theroux 1675 (isoTyPe). ASU 
84121. 
illardia | Wii B.L.T Southw. Natural- 
ist 17:183. 1972. Mexico. Coahuila: 11 Apr 

70, B.L. Turner 6023 (I ORVEE): ASU 130037. 

Grindelia fraxino-pratensis J.L. Reveal . : 
Beatley, Bull. Torrey Bot. Club 98:334. 1 
USA vada: 21 Aug 1968, JL. Reveal oe 

H. Holmgren (isotype). ASU 50079. 

Grae hintoniorum G.L.Nesom, Phytologia: 
68:328. 1990. Mexico. Nuevo Leon: 15 May 
1984, G.B. Hinton 18666 et al. (isoTyPE). ASU 
147845. 

Gutierrezia ionensis C.L. Lunell, Amer. Mid. 
Naturalist 2:194. 1912. Neotypified by M.A. 
Lane, Syst. Bot. 10:25. 1985. USA. Oregon: 27 
Aug 1983, M.A. Lane 3054 (ISONEOTYPE). ASU 
130892 


1045 


Halimium exaltatum J.N. Rose & PC. Standley, 
r. U.S. Natl. Herb. 23:833. 1923. Mexico. 
Michoacan: 27 Oct 1907, C.G. Pringle 10409 
(isotype). ASU 16235. 
Haplopappus enormidens R. Moran, Phytologia 
371. 1976. Mexico. Baja California: 19 Jul 
1975, R. Moran 22602 pate ASU 85778. 
Baa a eee R. Moran, Trans. San 
= oc. Naturalist Hist. i 154, 1969. 
i Baja eu 4 1966,R. Moran 
i eee ASU 2 
dea a rosaricus ey om a ee 
ie c. Naturalist Hist. 15:159. 
Mexico. i Ca a ie a. 967, R. ie 
14020 (ISOTYPE). ASU 2 
Haplopappus ign i ek & E. 
Ezcurra, Ci. Interamer. 26:16. 1986. Mexico. 
Sonora: 17 De oo E. Ezcurra 84001 
(isoTYPE). ASU to 
Heterotheca fulcrata peed Shinners vat. 
arizonica J.S. Semple, eee 39: 380.1987. 
U.S.A. Arizona: 13 Sep ee 
Heard 7923 (ISOTYPE). ene on loa 
Heterotheca ellie S.Semple, i Waterloo 
iol. Ser. 38: 83. 6. U.S.A. Arizona: 16 Oct 
1993,/.5.Semple i (isoTyPe). ASU on loan. 
Heterotheca mucronata V.L. Harms 
Turner var. harmsiana J.S.Sem 
loo Biol. Ser. 38:60. 1996. Mexico. Zacatecas: 
18 Jun 1976, D.J.Pinkava P13486 et al. (HOLO- 
Type). ASU on loan. 
Heterotheca zionensis J.C. Semple, shea 
39:384. 1987. USA. Utah: 15 Sep 1 C. 
Pillai Aoasss hae cs, ASU ek 
gh, Contr. Univ 
” Michigan Herb. 9:416. He fe Jalisco: 
1] oe) 1958,R.McVaugh 17078 (ISOTYPE). ASU 
12763 
Hymeno0 turneri K. Parker, Phytologia 20:192. 
970.USA. Texas: 21 _ 1965,B.L. Turner 5154 
ene 17464 
socoma menziesii on Hooker & G.A.W. 
Arnott) G.L.Nesom var.diabolica G.L.Nesom, 
Phytologia 70:96. 1991.USA. California: 5 Oct 
1985, D. Keil 19042 (ISOTYPE). ASU 146650. 
Isocoma tomentosa GL. Nesom, Phytologia 
70:109. 1991. Mexico. Chihuahua: 14 Oct 
1986, G.L. Nesom 5478 with W.L. Vorbik 
(isoTYPE). ASU 184811. 


1046 


Laphamia cochisensis W.E. Niles, Mem. New 
York Bot. Gard. 21:47. 1970. USA. Arizona: 10 
May 1964, WE. Niles 377 (ISOTYPE). ASU 75480. 

Machaeranthera amm J.L. Reveal, Bull. 
ue Bot. Club 97:172. 1970. USA. Nevada: 

1968, J.L. Reveal 1882 with N.H. 
aes (ISOTYPE). ASU 96211, 

Machaeranthera asteroides (J). [orrey) ELL. 
Greene var. glandulosa B.L. Turner 
oe 60:77.1986. USA. Arizona: 19 Sep 

Lehto L18904 et al. (isotype). ASU 


: — 

Machaeranthera canescens (F.1.Pursh) A.Gray 
SH aa B.L. Turner, Phytologia 60:77. 
1986. USA. Nevada: 27 aay 968, S. Stephens 
2830/7 a ASU 837 

ae cue iat debile G. ae var.arsenei 
J.Rzedowski, Bol. Soc. Bot.Mexico 31:81.1970. 

Michoacan: 18 aa 1967, J). Rzedowski 
so ine ASU 4493 
Parthenium lozanianum i H. Bartlett, Proc. 
mer. Acad. Arts 44:636. 1909. Mexico. Nuevo 
Leon:1 May 1906, C.G. Pringle 10247 (ISOTYPE). 
ASU 50587. 

Pectis exilis D. Keil, Phytologia 67:349. 198 
Mexico. Jalisco: 6 Oct 1903, CG. Pringle 8760 
(ISOTYPE). ASU 26006 

Pectis holochaeta (SF Blake) . Keil var. cana 
D. Keil, Phytologia 67:354 . Mex 
Michoacan: 7 Sep 1981, ee fe pene 

Luckow ee ASU 1 17. 

Pectis papposa W.H. ces & A. Gray var. 
grandis D. ai ee 26:32. 1974, USA. 
Texas: 15 Aug 1 - K/768 with LA. 
McGill (ISOTYPE). pet 5] 

Pectis purpurea IS. bate var. sonorae D. 
Keil, os 26:35.1974. Mexico. Sonora:07 
Sep 1977, D. Keil K8644 & J. ae Canne (ISOTYPE). 
ASU ae 

Perezia hooveri R.McVaugh, Contr. Univ. Michi- 
gan Herb. 9:466, 1972. Mexico. Jalisco: 3 Apr 

1951, R. McVaugh 11750 (isotype). ASU 
1274 

Perezia fritivees La Lave & aM se re xaldal 
Nov.Veg. Descr. 1:24.1824. 
genus Acourtia by B.L. ieee Prtoloi 


74:390. 1993. Mexico. Michoa 
1907, C.G. Pringle 10410 (ISONEOTYPE . a 
25734 


BRIT.ORG/SIDA 19(4) 


Perezia lozanii J.M.Greenman, Proc. Amer. Acad. 
68. 1905. Mexico. Hidalgo: 19 Sep 
1904, C.G. Pringle 8871 (ISOTYPE). ASU 50588. 

Perityle turneri M.A. Powell, Madrono 21:456. 
1972. Mexico. Chihuahaua: 2 Apr 1970, M.A. 
Powell 1858 (isotype). ASU 108726. 

Perymenium uxor R. McVaugh, Contr. Univ 
Michigan Herb. 9:439. 1972. Mexico. Nayarit: 
20 Dec 1970,R. McVaugh 25583 (ISOTYPE). ASU 

127508. 

ee ree B.L. eset Proc. 

. rA ts 39:116. 1903. Mexico 
ae 8 | 1902, ae as 8720 
ee 26126. 

Porophyllum pygmaeum 5 & J.B. 
Morefield, Syst. Bot. 14:583. 1989. USA. N 
vada: 4 Jun 1987, J.B. pare IE with : 
Ehrendorfer (ISOTYPE). ASU 160750. 

Psilostrophe mexicana R.C. Brown, Brittonia 
26:115.1974.Mexico. Chihuahua: 23 Jul 1973, 
RC. ith DJ. Pinkava & L.A. McGill 
(HOLOTYPE). ASU 50454, 

Sabazia anomola J.M. Greenman, Proc. Amer. 
Acad. Arts 41:262. 1905. 1970. Mexico. 
Hidalgo: 7 Aug 1904, C.G. Pringle 8853 
(ISOTYPE). ASU 26332 

Senecio cyclophyllus J.M. Greenman, Publ. 
Field Columbian Mus. Bot. Ser. 2:276. 1907. 
Mexico. Nuevo Leon: 31 on 906, C.G. Prin- 
gle 10230 (isotype). ASU 230 

Senecio filaris R. McVaugh, so Univ. Michi- 
gan Herb. 9:470. 1972. Mexico. Colima: 7-8 
Dec 1959, R. beech 1550 with WN. Koelz 
(isOTYPE). ASU 1 

Senecio ne coe R.M.Beauchamp, 
Brittonia 26:106.1974. USA. California: 23 May 
1973, 8. Moran & M. Douglas 21038 (ISOTYPE). 


SU 54676, 

Senecio platypus J.M.Greenman, Publ.Colum- 
bian Field Mus. Bot. Ser. 2:278. 1907. Mexico. 
Nuevo Leon: 21 Dec 1906, CG. Pringle 10352 
(IsoTYPE). ASU 23014. 

enecio rzedowskii J. Garcia Pérez, Phytologia 
57:275. 1985. Mexico. Mexico: 18 Jul 1982, / 
Rzedowski 37876 (ISOTYPE). ASU 184568. 
eolldago-'s ericamerioides G.L. Nesom, 
Phytol 9/:143.1989.Mexico. Nuevo Leon: 
26 Aug 1 1984, G.B. Hinton et al. 18763 (ISOTYPE). 
ASU 148389 


TACKY 


peta julia G.L. Nesom, ek 67:445. 
USA. Texas: 23 Sep 1989, G.L. Nesom 
with J. Nesom ors ASU 174657. 
Sphaeromeria ruthiae A.H. Holmgren, L.M. 
Shultz & T.K. Lowry, ae 28:257. 1976. 
USA. Utah: 20 Sep 1974,A.H. Holmgren 16003 
with L. Shultz & TK. Lowry (ISOTYPE). ASU 91050 
Stevia purpusii B.L. Robinson, Contr. Gray Herb. 
:18. 1930. Mexico. Morelos: 19 Nov 1902 
C.G. Pringle 11294 Se ASU 26690. 
a Haas J.D. Morefield, Madrono 
2.USA. ee 7 May 1991, 1.8. 
ree . (isoTYPE). ASU 188414. 
Tagetes stenophylla B.L. Robinson, Proc. Amer. 
Acad. Arts 43:44. 1907. Mexico. Michoa 
Jan 1907, CG. Pringle 10361 (ISOTYPE). eh 
646. 


— 
\O 
tae Oo) 


Tanacetum eaten es R. Moran, Trans. 

o Soc. Naturalist Hist.: 15:292. 1969. 

ee a See : se 1968,R. Moran 
15613 (ISOTYPE). ASU 2 

Townsendia jonesii (J.H. sete J.L.Reveal var. 
tumulosa J.L. Reveal, Great Basin Naturalist 
30:37. 1970. USA. Nevada: 17 Jun 1968, JL. 
Reveal 1326 (ISOTYPE). ASU 29721. 

Townsendia smithii L.M. Shultz & A.H. 
Holmgren, Brittonia 32:144. 1980. USA. Ari- 
zona: 4 Jun 1977, A.H. Holmgren 16380 with 
L.M. Shultz, J. Shultz & F. Smith (isotype). ASU 

217. 

Tridax yecorana B.L. Turner, Phytologia 79:286. 
1995. Mexico. Sonora: 7 Sep 1995, T.R. Van 
Devender 95-836 with A.L. Reina G., DA. 

an & M.E. Fishbein (ISOTYPE). ASU 206994. 

Vernonia joyaliae B.L. Turner, Phytologia 73:16. 


992. Mexico. Sonora: 17 May 1990, E. Joyal 


1465 with A. Alvarez, C. Smith. & J. Rascon (HO- 
LOTYPE). ASU 184020. 
I phalumd g M.L.Lane, Syst 
Bot. 8:305. 1983. Mexico. Durango: 24 Auc 
1979, M. Lane 2738 with D.W. Longstreth 
(isoTYPE). ASU 111180. 

Xanthocephalum gymnospermoides (A. 
Gray) G. Bentham & W.J. Hooker var. 
intermedium M.L. Lane, Syst. Bot. 8:305. 
1983. Mexico. Chihuahua: 23 Aug 1978, M. 
Lane 2479 with D.W. Pe (isoTYPE). ASU 
111179 


Xanthocephalum petrodoria S.L. Welsh & S. 


1047 


Goodrich, Brittonia 33:301. 1981. USA. Utah: 
24 Sep 1980, S.Goodrich 15240 (ISOTYPE). ASU 
120136. 


BETULACEAE 

Alnus arguta (Schlechtendal) E. Spach var. 
cuprea HH. Bartlett, Proc. Amer. Acad. Arts 
44:610. 1909. Lectotypified by JJ. Furlow, 
Rhodora 81:107.1979. Mexico. Oaxaca: 27 
May 1906, CG. Pringle 10251 (ISOLECTOTYPE). 
ASU 6251. 

Alnus arguta (Schlechtendal) E. Spach var. 
subsericea H.H. Bartlett, Proc. Amer. Acad. 
Arts 44:610. 1909. Mexico. Oaxaca: 27 May 
1906, C.G. Pringle 10252 (isoTyPE). ASU 6250. 

Alnus firmifolia M.L. Fernald, Proc. Amer. Acad. 
Arts 43:61. 1907. Mexico. Mexico: 30 Aug 
1905, C.G. Pringle 10040 (isoTyPe). ASU 50603. 


BORAGINACEAE 
Heliotropium calcicola M.L. Fernald, Proc. Amer. 
Acad. Arts 43:62. 1907. Mexico. Gue 28 
ae 905, C.G. Pringle 10262 on el 
576. 


gui turneri A. Richardson, Sida 6:237.1976. 
oahuila: 29 Jun 1971, AT. Richard- 
son 1595 (sorve, a 66700. 
Standley, Contr. U.S 
Natl. Hie 23:1230. 1924. Mexico. San Luis 
Potosi: 2 Jul 1890, C.G. Pringle 3518 (ISOTYPE). 
ASU 50580 


BRASSICACEAE 

Draba oreibata J.F. Macbride & E.B. Payson var. 
serpentina A. Tiehm & P.K. Holmgren, 
Brittonia 43:20. 1991. USA. Nevada: 29 Jun 

1966, N fees 2761 with J.L. Reveal 

(corved. ASU 177 

Lesquerella vicina - a J.L. Reveal & 
R. Rollins, Novon 7:9 1997. USA. Colorado: 9 
May 1988, J.L. Anderson 88-156 (isoTYPE). ASU 
206/733 


CACTACEAE 
Cochiseia bbinsorum W.H. Earle, 
Saguaroland Bull. 30:65. 1976. USA. Arizona: 
1 Oct 1975, J. eae & WH. Earle s.n. (HOLO- 
pues sae: See? 
is L.Bremer, Cact. Succ 
LW. S.) 56:165. 1984. Mexico. Queretaro: 14 
Oct 1976, L. Bremer 1076-8 (HOLOTYPE). ASU 
132677 


1048 


Coryphantha cuencamensis L. Bremer, Cact. 
Succ. J. (U.S.) 52:183. 1980. Mexico. Durango: 
26 Mar 1979,L. Bremer 379-4 (HOLOTYPE). ASU 
108730. 


ee delicata L. Bremer, Cact. Succ. J. 

.1979,Mexico. Tamaulipas: 11 Apr 

1976, L. rks A4/6-7 (HOLOTYPE). ASU 
poe 


ta L. Bremer, Cact.Succ.J.(U 
53: 276. 1981 Mexico. Tamaulipas: 1 de a 
A.B. Lau s.n. (HOLOTYPE). ASU 1 

Coryphantha laredoi C. Glass os Cact. 
Succ. J.(U.S.) 50:235. 1978. Mexico. Coahuila: 
1 Feb 1972,C. Glass & R. Foster 3761 (ISOTYPE). 
ASU 10485] 

Coryphantha laui L. Bremer, Cact. Succ. J.(U.S.) 
51:278.1979.Mexico. Coahuila:n.d., lL. Bremer 
476-3 (HOLOTYPE). ASU 108 

Coryphantha maliterrarum L. Bremer, Cact. 
Succ.J.(U.S.) 56:71. 1984. Mexico. Queretaro: 
13 Oct 1976,L. Bremer 1076-7 (HOLOTYPE). ASU 
128514. 

Coryphantha pusilliflora |. Bremer, Cact. Succ. 
J.(U.S.) 54:133. 1982. Mexico. Coahuila: 1] Jan 
1977, AB. Lau s.n. [= lL. Bremer 477-2] (HOLO- 
TYPE). ASU 121235, 

Coryphantha vivipara (T. Nuttall) N.L. Britton 

var. buoflama PC. Fischer, Cact. 

Succ. J.(U.S.) 52:28.1980.USA. Arizona: 22 May 
1979, PC. Fischer 6582 (isorTyPe). ASU 110901. 

sr delgadilloana J.P. Rebman & 

a, J. Arizona-Nevada Acad. $ 
ae os si . Mexico. Baja California: 9 = 
1994, J.P Rebman 2566 (HOLOTYPE), ASU 
838 


Echinocereus mapimensis E. Anderson, W.C. 
son & P Quirk, Cact. Succ. J.(U.S.) 70:285. 
1998. USA. Arizona: 22 Aug 1971, D. Keil 8136 
with L.A. McGill (isotype). ASU 216692, 
Echinocereus santaritensis W.Blum & J. Rutow, 
lum, Der Echinocereenfreund 12:93. 
USA. Arizona: 16 Apr 1996, J. Rutow JR 
52-56 (HOLOTYPE). ASU 211396. 
Mammillaria miegiana W.H. Earle, Saguaroland 
Bull. 26:77. 1972. Mexico. Sonora: 15 May 
oo WH. Earle s.n. (ISOTYPE). ASU 50081. 
Mammillaria tobuschii W.T. Marshall, 
Saguaroland Bull. 6:79. 1952. USA. Texas: 24 
Jun 1952, WT. Marshall & E.T. Blakely 1501 
(ISOTYPE). ASU 44125. 


BRIT.ORG/SIDA 19(4) 


Opuntia amarilla D. Griffiths, Bull. Torrey Bot. 
C 919. Mexico. Tabasco: 4 Aug 
1905, D. Griffiths 8032 (ISOLECTOTYPE). ASU 


129843. 

Opuntia anteojoensis D.J. Pinkava, Madrono 
23:75, 1976. Mexico. Coahuila: 7 May 1973, 
M.C. Johnston 10911 with T.L. Wendt & F. 
Chiang (isotype). ASU 59583 

Opuntia , D. Griffiths, Bull. Torrey Bot. 
Club 4 1916. Mexico. Tabasco: 1 Auc 

Seen ene ae yy 33265, 
140760 


sey ieee eee D. Griffiths, Bull. Torrey 
Club: 46:197. 1919. ae la D. 

S oe 2836 snes ASU ] 

ee ae V. Grant & oh aan Bot. 
Gaz 205.1979.USA. psi sel 1979, V. 
ae I (ISOTYPE). ASU 210 

Opuntia eee D. Griffiths, aa Biol. Soc. 
Wash. 29:11.1916. USA. Arizona: 1 Sep 1911, 
D. Griffiths ae es ASU 140753. 

Opuntia go iths, Monatsschr. 
ca me 34. os 3. Mexico. Aguas 
Calientes: Aug 1905, D. Griffiths 8092 
(ISOTYPES). ASU 129842, 133264. 

Opuntia intricata D. Griffiths, Proc. Biol. ie 
Wash. 29:1 6.USA. California: 1 May 19 
D. ae 7 (ISOTYPE). ASU 140754. 

Opuntia ithypetala D. Griffiths, Bull. Torrey Bot. 
Club 43:529. 1916, USA. Texas: San Antonio 
Garden SAG506 pian ASU 141 

Opuntia lindsayi J.P R n, Cact. & Succ. J. 
(U.S.) 69:67. 1997, vee Baja California: 24 
Jun 1 ee oe Rebman 2782 with CA. Cano & 


M sendiz (HOLOTYPE). ee 195943 

Opuntia linguiformis D. Griffiths, Rep. (Annual) 
see Bot. Gard. 19:270. 1908. USA. Texas: 

Aug. 1906, D. Griffiths 8377 (ISOTYPE). ASU 
140761. 

Opuntia pachona D.Griffiths, Rep. (Annual) Mis 
souri Bot.Gard.21:168, plate 22 (in part). 1910. 
Mexico, Zacatecas:19 Sep 1905, : Griffiths 
8141 (iSOTYPES). ASU 129844, 13325 

ie pinkavae B.D. Parfitt, ae 99:223. 

USA. Arizona: 30 oo 980, B.D. Parfitt 
a (HOLOTYPE). ASU 111 

Opuntia sanfelipensis J. ane Hasseltonia 
6:17.1999. Mexico. Baja California:8 Apr 1994, 

JP Rebiian 2540 (HOLOTYPE). ASU 195789. 


N 
any 


TLRERI 


Opuntia valida D.Griffiths, Proc. Biol. Soc.Wash 
24. 1914. USA. New Mexico: 1908, D. 
Griffiths 9194 (ISOTYPE). ASU 140780. 
Opuntia x campii M.A. Baker & DJ. Pinkava [= 
O. acanthocarpa G. Engelmann & J.M. Big- 
elow x O. bigelovii Engelmann], Cact. Succ. 
J.(U.S.) 71:320.1999.USA. Arizona:9 Mar 1995, 
M.A. Baker 11661 (HOLOTYPE of 3 sheets). 
ASU 203945, 203946, 203947. 
Sclerocactus parviflora D. Woodruff &L.D.Benson 
blessingiana W.H. Earle, Saguaroland 
Bull. 34:29. 1980. USA. Arizona: 10 Mar 1978, 
W.H. Earle s.n. (HOLOTYPE). ASU 108791. 
Turbinicarpus booleanus G.S. Hinton, 
Phytologia 80:62. 1996.Mexico. Nuevo Leon: 
1 Mar 1992, G.B8. Hinton et al. 21805 (ISOTYPE). 
ASU 186715 


CAPRIFOLIACEAE 
shir occidentalis J.A.Villarreal, Brittonia 49:84. 
7. Mexico. Durango: 22 Aug 1995, JA. 

ries 8882 (ISOTYPE). ASU 235854. 

Viburnum cuneifolius H.H. Bartlett, Proc. Amer. 
Acad. Arts 45:635. 1909.Mexico. Nuevo Leon: 
27 Mar 1906, C.G. Pringle 10234 oe 
7545 


CELASTRACEAE 
Canotia wendtii M.C. Johnston, Brittonia 27:121. 
Mexico. Chihuahua: 25 Aug 1973,M.C. 

Johnston et al. 12345 (isoTyPe). ASU 137501. 

Celastrus tetramerus PC. Standley, Contr. U.S. 
Natl. Herb. 23:679. 1923. Mexico. Guerrero: 8 
Oct 1906, C.G. Pringle 10319 (IsoTyYPe). ASU 
69983. 


CHENOPODIACEAE 

Atriplex frankenioides R. Moran, Bees 
30:1.1975.Mexico. Baja California Sur: 
1974, R. Moran 21184 (isoTYPE). ASU 84120. 

Nitrophila mohavesis PA. Munz & J.C. Roos, 
Aliso 3:112.1955.USA. California: 15 Jun 1954, 
J.C. Roos & A.R. Roos 6140 (iSOTYPE). ASU 
133206 


CISTACEAE 
Halimium exaltatum J.N. Rose & PC. Standley, 
U.S. Natl. Herb. 23:833. 1923. Mexico. 
Michoacan: C.G. Pringle 10409 (ISOTYPE). ASU 
16235, not found. 


CONVOLVULACEAE 
Ipomoea igualensis C.A.Weatherby, Proc. Amer. 


1049 


Acad. Arts 45:427.1910.Mexico. Guerrero: 21 
Sep 1905, CG. Pringle 10054 (isotype). ASU 
50589. 


CROSSOSOMATACEAE 

Glossopetalon spinescens A. Gray var. 
microphyllum N.H. Holmgren, Brittonia 40: 
272.1988. USA. Utah: 23 May 1987, N.H. 
Holmgren 11292 with RK. Holmgren (\sOTYPE). 
ASU 215438. 


EUPHORBIACEAE 

Acalypha salicioides H.H.Rusby, Descr.S. Amer. 
P|.:-46-47. 1920.Colombia. Santa Marta: 1898 
1901,H.H. Smith 1428 (isoTYPE). ASU 127470. 

Euphorbia aaron-rossii A.H. Holmgren & N. 
Holmgren, Brittonia 40:357. 1988. USA. Ari- 
zona: 5 May 1971,A.H. Holmgren et al. 15558 
(ISOTYPE). ASU 177194 

Euphorbia ariensis F.\W.H.A Humboldt, A.J.A. 
Bonpland & K.S. Kunth var. villicauli MLL. 
Fernald, Proc. Amer. Acad. Arts 43:62. 1907. 
Mexico. Michoacan: 29 Oct 1905, C.G. Pringle 

10116 (ISOTYPE). ASU 50606. 
sap chalicophila C.A. Weatherby, Proc. 
cad. Arts 45:426.1910.Mexico. Jalisco: 

12 ae 1903, C.G. Pringle 11846 (\soTyPE). ASU 
50590. 

Euphorbia henricksonii M.C. Johnston, 
Madrofto 22:374. 1974. Mexico. Chihuahua: 
29 Sep 1972, M.C. Johnston 9585 with F 
Chiang & T.L. Wendt (isoTyPe). ASU 137473. 

Manihot intermedia C.A. Weatherby, Proc. 
Amer. Acad. Arts 45:427. 1910. Mexico. 
Guerrero: 27 Jul 1907, C.G. Pringle 13938 
(IsoTYPE). ASU 50591. 

FABACEAE 

Astragalus ackermanii R. Barneby, ae 
32:26. 1980. USA. Nevada: 27 May 1979, N.H. 
Holmgren et al. 9200 (ISOTYPE). ASU 11 Be 


1942. USA. Nevada: 14 Jun 1937, .W. Clokey 
7572 (ISOTYPE). ASU 66709 

Astragalus atwoodii S.L. Welsh & K.H. Thorne, 
Great Basin Naturalist 37:103. 1977. USA. Ari- 
zona: 15 Jun 1976,N.D. Atwood 6794 (ISOTYPE). 
ASU 108002. 

Astragalus chuskanus R. Barneby & R. 
Spellenberg, Brittonia 39:188. 1987.USA.New 
Mexico: 28 Jun 1986, R. Spellenberg et al. 8521 
(isoTYPE). ASU 148180. 


1050 


Astragalus holmgreniorum R. Barneby, 
Brittonia 32:24. 1980. USA. Arizona: 25 May 
1979, N.H. Holmgren et al. 9175 (ISOTYPE). ASU 

13742. 

Astragalus tortipes J.L. Anderson & J.M. Porter, 
Syst. Bot. 19:116.1994,USA. Colorado: 25 Apr 
1989, J.L. Anderson 89-16 (ISOTYPE). ASU 


200009. 

Ateleia herbert-smithii A. Pittier, Contr. U.S. 
Naturalist Herb, 20:112. 1918. Colombia. 
Santa Marta: 1898-1901, H.H. Smith 817 
(ISOTYPE). ASU 127480. 

Coursetia paniculata M.Sousa, Syst. Bot. 12:1 

987.Mexico.Oaxaca:9 Mar 1985,M. ee et 

al. 5320 (Isotype). ASU 150423, 

Dalea ananassa R. Barneby, Mem. New York Bot. 
Gard. 27:175. 1977. Mexico. Sonora: 26 Oct 
1961,H.S. Gentry, A.S. Barclay & Arguelles 19384 
(ISOTYPE). ASU 79047, 

a oe R.Barneby, ea Flora 

989. USA. Nevada: 24 J 

ea 77193. 


ae et al. 9902 


je') 


~~ 


1970, LL. Reveal 2202 (iSoTYPE). ASU 41443. 
Lotus mearnsii (N.L. Britton) ELL. ae var. 
eee ee L.Anderson,Madron 61. 
USA. Arizona: 3 Apr 1987, J/.L. hes 
- ~ (HOLOTYPE). ASU 166482. 
Lupinus bicolor J. Lindley subsp. marginatus 
D.B. Dunn, Aliso 3:159. 1955. USA. California: 
19 Apr 1946, ai au & C. Epling 2129 
(ISOTYPE). pe 67 
D.Do glas subsp medius L_E. 
Dating, Aree ve ren 45:488. 1951. 
on: : ie 949, L.E. Detling 6546 
“or. ASU 7 
osa ape “ D. Parfitt & DJ. Pinkava, 
pices 30:172. 1978. Mexico. Coahuila: 14 
Aug 1975,D.J. Pinkava P13073 (HOLOTYPE). ASU 
89319 


Ovyvutroanic dafl 


y (P.M. Pallas) A.P de Candolle 
var. pulcherrima S.L.Welsh & A.Huber, Great 
Basin Naturalist 55:277, 1995. USA. Utah: 18 


Jul 1994, A. Huber 1673 (iSoTYPE). ASU 209524. 


FAGACEAE 
Quercus acherdophylla W. Trelease, Mem. Natl. 
cad. Sci. 20:283, plate 367. 1924. Mexico. 
ce 1 Sep 1904, C.G. Pringle 10008 
(ISOTYPE). ASU 50601 


BRIT.ORG/SIDA 19(4) 


Quercus hypoxantha W. Trelease, Mem. Natl. 
Acad. Sci. 20:170, plate 339. 1924. Mexico. 
Coahuila: 12 a oe C.G. Pringle 10227 
(isoTyPE). ASU 313 

seis Heese : es Amer.J.Bot. 
79:1 1992. Mexico. Chihuahua: 22 Jun 
ee . Spellenbera et al. 9281 (ISOTYPE). ASU 
181983 

Quercus nwsophyl C.A.Weatherby, Proc. Amer. 

Acad. Sci 3.1910.Mexico, Nuevo Leon 


10025 ISOSYNTYPE). 


—= 


2/7 Mar 1 ae Pringle 
ASU 50581 
Quercus rysophylla CA. Seah Proc. Amer. 
Acad. Sci.45:423. 1 Mex 
12 Mar 1906,C.G. Pringle oe Foe ene 


ASU 50582. 
Quercus rysophylla C.A. Weatherby, Proc. Amer. 
Acad. Sci.45:423. 1910. Mexico. Nuevo Leon: 
10 Jul 1907, C.G. Pringle 10379 (ISOSYNTYPE). 
ASU 50583. 
Quercus subtriloba W. Meroe Mem. Natl. 
cad. Sci. 20:81, plate 118. 1924. Mexico. 
ae algo: 2 Sep 1906, ee Pringle 10303 
(ISOTYPE). ASU 50600 


GENTIANACEAE 

Centaurium namophilum J.L. Reveal, CR. 
Bloome & J.Beatley, Bull. Torrey Bot. Club 100: 
353.1973.USA. Nevada: 25 Jul 1972, J. Beatley 
13447 with JL. Reveal (ISOTYPE). ASU 50077. 

Centaurium pusillum A. Eastwood, Proc. Amer. 
Acad. Arts 44:605. 190 ico. Michoacan 
29 Nov 1907,C.G. Pringle 10408 (\soTYPE). ASU 


— 


Gentiana durangensis J.A. Villarreal, Acta Bot. 
Mex. 34:49. 1996. Mexico. Durango: 6 Oct 
1985, S. Gonzdlez 3464 (ISOTYPE). ASU 235855. 
apes Jone ulenta J.S. Pringle, Rhodora 
68:2 966. USA. poe 7 Oct 1906, Chase 
oe ee ASU 3210 
Halenia hintonii A.A. fee Hooker's Icon. Pl. 
34:tab. 3399, 1939. Mexico. Mexico: 9 Aug 
1935, G.B. Hinton 8273 (ISOTYPE). ASU 152531. 
GERANIACEAE 
Geranium toquimense N.H. Holmgren & AH. 
olmaren, Brittonia 29:34. 1974.USA. Nevada: 
1518 with LL. Re- 


3 Aug 1964, NH. a 

veal es ASU 6 
HYDROPHYLLACEAE 
Phacelia beatleyae J.L. Reveal & L.C.Constance, 


ILACAT 


Brittonia 24:199. 1972. USA. Nevada: 29 Apr 
1971,/. Beatley 12358 (ISOTYPE). ASU 50080. 

Phacelia crenulata J. Torrey in S. Watson var. 
angustifolia N.D. Atwood, Great Basin Natu- 
ralist: 35:158.USA. Arizona: 18 ihe 1970,N.D. 
Atwood 2597 (ISOTYPE). ASU 203 

Phacelia ae S.L. ieee radars 
95:402 3. USA. Utah: 13 Jun 1983, N.D. 
Aanecd ne (IsoTYPE). ASU 130043. 


Phytologia 26:437. 1973. USA. Utah: 6 May 
1970,N.D. Atwood 9809 (isotypes). ASU 71165, 
203837. 

Phacelia marshall-johnstonii N.D. Atwood & 
DJ. Pinkava, Madrono 24:212. 1977. Mexico. 
Coahuila: 15 Aug 1975, DJ. Pinkava 13100 
with T. Reeves (HOLOTYPE). ASU 92596. 


HYPERICACEAE 

Hypericum hintonii A.A. Bullock, Kew Bull. 
1936:390. 1936.Mexico.Mexico:21 Sep 1932, 
G.B. Hinton 1796 (ISOTYPE). ASU 50465. 


LAMIACEAE 

Salvia flaccidiflora M.L. Fernald, Proc. Amer. 
Acad. Arts 43:66. 1907.Mexico. Hidalgo:6 Sep 
1906, C.G. Pringle 10298 (ISOTYPE). ASU 32872. 

Salvia hispanica C. Linnaeus var. chionocalyx 
M.L. Fernald, Amer. Acad. Arts 43:63. 1907. 
Mexico. Michoacan: 16 Oct ° C.G. Pringle 
8837-1/2 (ISOTYPE). ASU 3360 

Salvia lilacina M.L. Fernald, aa Amer. Acad. 
Arts 45:418.1910.Mexico.Michoacan: 15 Oct 
1904, C.G. Pringle 13279 a 33625. 

Salvia muralis M.L. Fernald, Amer. Acad. Arts 

65. 1907. Mexico. Guerrero: 28 Sep 190 

C.G. Pringle 10072 (ISOTYPE). ASU 50599. 

Salvia subpatens E. Epling, Repert. Spec. Nov. 
Regni Veg. Beih. 110:97.1938.Mexico, Mexico: 
2/ 1933, G.B. Hinton 4376 (Isotype). ASU 
5449 


Poca 


Salvia uruapana MLL. Fernald, Proc. Amer. Acad. 
Arts 45:418.1910.Mexico.Michoacan: 16 Oct 
1904, C.G. ised a (isoTyPE). ASU 33675. 

Satureja maderensis J. Henrickson, Brittonia 
33:211.198] es Coahuila: 5 Aug 1973, 
J. Henrickson 11888 with T.L. Wendt (ISOTYPE). 


ASU 74254. 
Scutellaria lutilabia M. Lane & GL. Nesom, 
adrono 35:112. 1988. Mexico: G.L. Nesom 
4273 (ISOTYPE). ASU not found. 


1051 


LOASACEAE 

Mentzelia collomiae C.M. Christy, Novon 7:25. 
1997. USA. Arizona: 25 ee . 94,C.M. Christy 
2385 (HOLOTYPE). ASU 2 

Mentzelia goodrichii K. " ae & S.L.Welsh, 
Rhodora 95:407.1993.USA.Utah: 15 Jul 1992, 
S. Goodrich 23806 (\soTYPE). ASU 203839. 


LORANTHACEAE 
santa alni : H. Bartlett, Proc. se Acad. 
Arts 44:9 Oo. Oax 2 May 
1906, C.G. : ne on any ee 37488. 


LYTHRACEAE 

Cuphea imberbis R.N. Rose, In E. Koehne, Bot. 
Jahrb. Syst. 41:94. 1907. Mexico. Hidalgo: 29 
Jul 1904, C.G. Pringle 8979 (isoTyPE). ASU 
50593. 


Cuphea lanceolata W. Aiton, Hortus Kewensis. 
Il (ed. 2):150.1911.Neotypified by $.Graham 
in Syst. Bot. Monogr. 20:84. 1988. Mexico. 

0 Sep 1904, C.G. Pringle 11983 
(ISONEOTYPE). ASU 38289. 


ACEAE 

Abutilon holosericeum G.H.A. Scheele, Linnaea 
21:471.1848.USA. Texas: 1846, Lindheimer 354 
(ISOTYPE). ASU 37985 

Abutilon pinkavae P.A. Fryxell, Phytologia: 
37:285.1977.Mexico. Coahuila: 14 Aug 1975, 
DJ. Pinkava 13044 with T. Reeves (HOLOTYPE). 

SU 79373. 

Decaschistia brynesii P.A. Fryxell, Austral. J.Bot. 
22:188. 1974. Australia. Northern Territory: 30 
Jan 1969, N. Byrnes 1320 (isoTyPE). ASU 
199035. 

Decaschistia brynesii P.A. Fryxell subsp. 
landulacea PA. Fryxell, Austral. J. Bot. 22:189 
1974. Australia. Northern Territory: 28 May 

1971,N. Byrnes 2284 (ISOTYPE). ASU 206992. 
arcs se pee i FD. Wilson, Austral. J. Bot. 
— ae Nile: Northern Territory: 20 
= ie 9, N. Byrnes 1390 (ISOTYPES). ASU 
oe 99038, 199039. 
goons ae T. Howell & G.H. True, Four 
S 4:20. 1972. USA. California: 30 Jul 
at ee ee ell 7630 (ISOTYPE). ASU 
15515 

MELIACEAE 

Trichilia oblanceolata H.H. Rusby, Descr. New 

Sp. South Amer. PI.:36-37. 1920. Colombia. 


1052 


Santa Marta: ol 901,H.H. Smith 447 
(ISOTYPE). ASU 127617. 


MENISPERMACEAE 

Menispermum mexicanum J.N. Rose, Contr. 
U.S. Natl. Herb. 13:302. 1911. Mexico, Nuevo 
Leon: 8 Jul 1907, C.G. Pringle 10378 (ISOTYPE). 
ASU 50594 


MORACEAE 
Ficus subrotundifolia J.M. Greenman, Proc. 
Acad. Sci.41:237.1905.Mexico: Jalisco: 
28 Sep 1903, C.G. Pringle 11850 (ISOSYNTYPE). 
ASU 50596. 


MYRTACEAE 

Calycolpus bolivarensis L.R. Landrum, Ann. 
Missouri Bot. Gard. 76:930. 1989. Venezuela. 
Bolivar: 5 Mar 1977, A. Fernandez 3068 
(ISOTYPE). ASU 160293. 

manesia espiritosantensis LR. 
Landrum, Brittonia 39:245. 1987. Brazil. 
Espirito Santo: 27 Jan 1984, D.A. Folli 494 
(isoTyPe). ASU 145539, 

Eugenia anglohonduransis C.L. Lundell, 
Wrightia 2:123.1961. British Honduras. Stann 
Creek District: . ii 1954, PH. Gentle 8354 
(ISOTYPE). ASU 15370 

Eugenia blanda 7 Sobral, Bradea 26:234. 1993. 

Brazil. Minas Gerais: 12 Jun 1990,G.&M 
Hatschbach 54169 with V. Nicolack (ISOTYPE). 
ASU 209876. 

Eugenia gentlei CL. Lundell, Publ.Carnegie Inst. 
Wash. 47 eg. Peten):216. 1937. British 
Honduras. eee 7 1935,PH.Gentle 1684 
(ISOTYPE). ASU 1 

Myrcia eer are H.F.C. Kiaerskou, 
Enum.Myrt. Bras. 90. 1893. Brazil:n.d.,Glaziou 

11986 (ISOTYPE). ASU nhs 
ee microph Ila O.K. Berg var. australis 
iels, Bot. Jahrb. ens 37:593. 1906. Bo- 
vee 1892, M. Bang 1573 (ISOSYNTYPE). 
ASU 152743. 
sieges ee O.K. Berg var. australis 
s, Bot. Jahrb. Syst. 37:593. 1906. Bo- 
orate 892, M. Bang 1911 (ISOSYNTYPE). 
ASU 152744 


NYCTAGINACEAE 
Abronia argillacea S.L. Welsh & S. Goodrich, 
Great Basin Naturalist 40:78. 1980.USA. Utah: 


BRIT.ORG/SIDA 19(4) 


30 May 1979, S.L. Wels ’ . ea Welsh 
16689 (ISOTYPE). ASU 1 

Mirabilis pringlei CA. ey Proc. Amer. 
Acad. Arts 45:424. 1910.Mexico. Guerrero: 23 
Jul 1907, C.G. Pringle 10384 (isoTYPE). ASU 


50597. 

ONAGRACEAE 

Gayophytum decipiens H. Lewis & J. 
Szweykowski, Brittonia Bee 964. USA 
California:8 Aug 1962,H. Lew 1311 (\SOTYPE) 


ASU 127955. 


POLEMONIACEAE 

Gilia nyensis J... Reveal, Bull. Torrey Bot. Club 
96:480. 1969. USA. Nevada: 29 May 1968, /.L. 
Reveal 1048 (\SOTYPE). ASU 30054 

Ipomopsis sancti-spiritus D. Wilken & R. 
Fletcher, Brittonia 40:48. 1988. USA. New 
Mexico: 15 Jul 1986, D. Wilken & R. Fletcher 
14645 entices 154587, 

Linanthus jamauensis R. Moran, Madrono 
24:147.1977. ene Baja California: i a 
1973, R. Moran 20930 (IsoTYPE). ASU 1 

POLYGONACEAE 

seated rosulenta JL. Reveal, Phytologia 

66:95, . Mexico. Baja California: 23 Mar 
1988, ie Reveal et al. 6729 (iSOTYPE). ASU 


= 


153148. 
Dedeckera eurekensis J.L. Reveal & J.T. Howell, 
Brittonia 28:246. 1976. USA. California: 29 Jul 
19 .L. Reveal et al. 3909 (isotype). ASU 
| 19050. 
Eriogonum aliquantum J.L. Reveal, Phytologia 
34:460. 1976. USA. New Mexico: 14 Jul 1972, 
JL. Reveal 2770 with C. Reveal (ISOTYPE). ASU 
119046, 
Eriogonum apachense J.L. Reveal, J. Arizona- 
Nevada Acad. Sci. 5:222. 1969. USA. Arizona: 
7 Sep 1968, £. Lehto 13400 with DJ. Pinkava & 
D. Keil (SoTYPES). ASU 12590, 12589, 116493, 


Eriogonum argophyllum J.L. Reveal, 
Phytologia 23:168. 1972. USA. Nevada: 7 Jul 
1969, N.H. Holmgren 3667 with PM. Kern 
(isoTYPE). ASU 40397, 

Eriogonum atrorubens G. Engelmann var. 
nemorosum W.). Hess & J.L. dines Great 
sr Naturalist 36:302. 1976. 

ango: 11 Aug 1971, JL. ees én 
ie Hess & R.W. Kiger (ISOTYPE). ASU 5007 


ILACHI 


gai cane beatleyae J.L. Reveal, Aliso 7:415. 
2.USA.Nevada: cee 1971, 1.L. Reveal et 
io (ISOTYPE). ASU 50070. 
Eriogonum bifurcatum ‘in sei 7:357. 
971. USA. Nevada: 13 Jun 1970, J.L. Reveal 
2283 (ISOTYPE). ASU 50069. 
sae capistratum J.L.Reveal,Phytologia 
:254. 1989. USA. Idaho: 11 Jul 1975, JL. Re- 
pe with B.J.Ertter (ISOTYPE). ASU 119199. 
Eriogonum capistratum J.L. Reveal var. welshii 
J.L. Reveal, 66:257. 1989. USA. 
Idaho: 16 Jun 1976, AL. 4501 with SLL. 
Welsh eas ASU 11 
Eriogonum concinnum fa oa Bull. Torrey 
Bot. Club 96:476. 1969. USA. Nevada: 5 Jul 
1968, J.L. Reveal 1501 (ISoTyPE). ASU 30850. 
set vo es correllii J.L.Reveal, Sida 3:198. 1968. 
5:23 Aug 1967, J.L. Reveal 878 with 
: a de (isoTyPE). ASU 13753. 
Eriogonum corymbosum G. Bentham var. 
albogilvum J.L. Reveal, Great Basin Natural- 
ist 27:218. 1968. USA. Utah: 26 Aug 1967, J.L. 
Reveal 726 with C.G. Reveal (ISOTYPE). ASU 


4. 

Eriogonum corymbosum G. Bentham var. 
davidsei J.L. Reveal, Great Basin Naturalist 
27:216. 1968. USA. Utah: 9 Sep 1967, J.L. Re- 
veal 956 with G. Davidse (isOTYPE). ASU 13756. 

ee corymbosum G. Bentham var. 
erectum J.L. Reveal & J.D. Brotherson, a. 
a Naturalist 27:213. 1968. USA. Utah: 1 
Aug 1966,N.H.Holmgren 3022 with J.L. a 
(iSOTYPE). ASU 13659. 

Eriogonum corymbosum G. Bentham var. 
velutinum J.L.Reveal, Great Basin Natrualist: 
27:224, 1968. USA. New Mexico: 5 Sep 1967, 
JL. Reveal 919 with G. Davidse (IisoTYPE). ASU 
13745, 

Eriogonum cronquistii J.L. Reveal, Madrono 
19:289. 1969. USA. Utah: 14 Aug 1966, N.H. 
Holmgren 3010 with J.L. Reveal (ISOTYPE). ASU 

661. 

Eriogonum deflexum J. Torrey var.nevadense 
J.L. Reveal, Phytologia 25:206. 1973.USA.Ne- 
vada: 18 Jul 1972, JL. Reveal 2785 with C.G, 
Reveal (ISOTYPE). ASU 50075. 

Eriogonum deflexum J. Torrey var. rectum J.L 
Reveal, ec 66:263.1989.USA. Califor 

87, JL. Reveal 6385 with CR. 
Broome ene ASU 153150. 


1053 


ee duchesense J. Reveal, aes 
5:183. 1973. USA. Utah: 2 Sep 1964, JL. Re- 
veal 678 (ISOTYPE). ASU 13664 
Eriogonum ericifolium J. Torrey & A. Gray var. 
thornei J... Reveal & J. Henrickson, Madrono 
23:205. 1975. USA. California: 26 Aug 1973, J. 
Henrickson 12676 (ISOTYPE). ASU 114198. 
Eriogonum fimbriatum W.J. Hess & J.L. Reveal, 
Great Basin Naturalist 36:308. 1976. Mexico. 
Nuevo Leon: 19 Sep 1972, J.L. Reveal 3144 
with WJ. Hess (isoTYPE). ASU 50072 
Eriogonum gracile G. Bentham var. incultum 
J.L.Reveal, Lory 66:265. aan USA. Cali- 
fornia: 28 Jun 1987, /.L. veal 6620 with CR. 
Broome fa ASU 153 
sain ee Le he Phytologia 
9:291, 1968. USA. Utah: 13 Aug 1966, N.H. 
i hae es with JL. Reveal (ISOTYPE). ASU 
13667 


sal 


Eriogonum hylophilum J.L. Reveal & J.D. 
Brotherson, Great Basin Naturalist 27:190. 
1968. USA. Utah: 15 Aug 1966,N.H. ti 
3017 with JL. Reveal (isoTYPE). ASU 1 

Eriogonum inflatum J.Jorrey & J.C. svar var. 
contiguum J.L. Reveal, Aliso 7:221.1970.USA. 
Nevada: 3 Jun 1969, LL. Reveal 2157 with ME. 
Matthews (ISOTYPE). ASU 50076. 

Eriogonum lancifolium J.L. Reveal & J.D. 
Brotherson, Great Basin Naturalist 27:187. 

1968. USA. Utah: 9 Sep 1967, a ea 957 
with G. Davidse (ISOTYPE). ASU 13 
ia leptocladon J. Torrey : -— var. 
apiliunculum J.L. Reveal, Brittonia 26:92. 
a USA. Arizona: 7 i 1972, J.L. Reveal 
2906 (ISOTYPE). ASU 5 

Eriogonum lewisii ee Basin Natu- 
ralist 45:277. 1985. USA. Utah: 30 Jul 1976, J.L. 
Reveal 4596 with C.G. Reveal (iSOTYPE). ASU 
119203. 

Eriogonum meledonum J.L. Reveal, Phytologia 
66:254. 1989. USA. Idaho: 5 Jul 1974, J.L. Re- 
veal 3674 (ISOTYPE). ASU 116438. 

Eriogonum microthecum T. Nuttall var. 
corymbosoides J... Reveal, Brignam Young 
Univ. Sci. Bull. Biol. Ser. 13:25. 1968. USA. Cali- 
fornia: 10 Sep 1968, J/.L. Reveal 2090 (ISOTYPE). 
ASU 50071. 

Eriogonum microthecum T. Nuttall var. 
lapidicola J.L. Reveal, Brigham Young Univ. 
Sci. Bull. Biol. Ser. 13:28. 1968. USA. Nevada: 


1054 


25 Aug 1968, JL. tee i with N.H. 
Holmgren USOT VRE) ASU 30 

Eriogonum mort J. a Brittonia: 
26:90. 1974. USA. Arizona: 16 Aug 1972, /.L. 
Reveal 2904 (isoTyPe). ASU 50068 

Eriogonum natum J.L. Reveal, Great pee Natu- 
ralist 35:363. 1975, USA. Utah: 13 Aug 1975, 
JL. Reveal 3924 with M.L. Reveal (ISOTYPE). ASU 
119044. 


Eriogonum ochrocephalum S. Watson var. 
alexanderae J... Reveal, Great Basin Natu- 
ralist 45:276.1985.USA. Nevada: 21 Jun 1978, 
JL. Reveal et al.4737 (isotype). ASU 119197. 

Eriogonum ochrocephalum S. Watson var. 
sceptrum J.L. Reveal, Phytologia 66:252. 
1989. USA. Idaho: 7 Jul 1974, .L. Reveal 3687 
(ISOTYPE). ASU 116445. 

Eriogonum ordii S. Watson, Proc. Amer. Acad. 
Arts 21:468. 1886. USA. Arizona: 1 a 1884, 
CJ. Lemmon ices pele ASU 4504 

Eriogon TN iceeih 
J.L. Reveal, Phytologia 66:259. 1989. USA. 
Idaho: 12 Jul 1975, LL. Reveal 3883 with BJ. 
Ertter (isoTYPE). ASU 119201. 

Eriogonum da tee dia J.L. Reveal, Great 
Basin Naturalist 33:120.1973.USA. Colorado: 
16 Jun 1972, = fra 2/80 with C.G. Reveal 
(isOTYPE). ASU 5 

Eriogonum Si ed J. Torrey var. 
cervinum J.L. Reveal, Great Basin Naturalist 
34:246, 1974. USA. Utah: 18 Aug 1973, N.D 
sig 5895 with L.C. Higgins (ISOTYPE). 


ASU 


Bo 
= 


ios ncn preclarum J.L. Reveal, Brittonia 
8:338. 1976. Mexico. Baja California: 9 Feb 
1973,R. Moran 19964 with JL. Reveal (ISOTYPE). 
ASU 116456. 
et acie pro J.L. Reveal, Aliso 7:417, 
2. USA. Oregon: 22 Jun 1971, JL. Reveal 
a (ISOTYPE). ASU 50066 
Eriogonum rupinum J.L. Reveal, Aliso 7:226. 
1970.USA. Nee : Aug 1969,/. Beatley 9458 
(ISOTYPE). ASU 500 
Eriogonum saurinum on Reveal, Great Basin 
Naturalist 27:196. 1968. USA. Utah: 15 Aug 
1966, N.H. Holmgren 3019 with JL. Revea 
(ISOTYPE). ASU 13675 
Eriogonum scabrellum J.L. Reveal, Ann. Mis- 
sourl Bot. Gard. 55:74. 1968. USA. Utah: 8 Sep 


rad 


BRIT.ORG/SIDA 19(4) 


1967, JL. Re veal )4 with UJ. Davidse \IOULTPRC). 
ASU ] 

Eriogonum shockleyi S. Watson var.packardae 
J.L. Reveal, Ac 66:257. 1989. USA. 
Idaho: 7 Jul 1974, JL. Reveal et al. 3686 
PORES | 16433 

Eriog ithii J.L.Reveal, Great Basin Natu- 
ralist 27:202. 1968. USA. Utah: 14 Aug 1966, 
NH. foci 3012 with JL. Reveal (\SOTYPE). 
ASU 136 

Eriogonum sor edium J.L. Reveal, Great Basin 

Naturalist 41:229. 1981. USA. Utah: 29 Aug 
1980, S.L. Welsh et al. 20192 (\soTyPE). ASU 
120398. 

Eriogonum thompsonae S. Watson var. 
albiflorum J.L. Reveal, Madrono 19:299. 

1969.USA.Utah: 11 Aug 1966, N.H. fel 
2991 with LL. Reveal ae: ASU 1368 

Eriogonum veal, Sida 5:265. 1 ie 
Mexico. Nuevo ee i: Sep 1972, /.L. Reveal 
3174 with WJ. Hess (\soTYPE). ASU 50074. 

Eriogonum umbellatum J. Torrey var. 
chlorothamnus J.L. Reveal, Calif. Fl. Suppl.: 
44.1968. USA. California: 23 Jul 1966, N.H. 
Holmgren 2938 with J.L. Reveal (isoTyPE). ASU 


Eriogonum umbellatum J. Torrey var. 
furcosum J.L. Reveal, Great Basin Naturalist 
45:278.1985.USA. California: 23 1975, A.L. 
Reveal 3971 (isotype). ASU 119 

Eriogonum umbellatum J. Torrey var. vernum 
J.L.Reveal, Great Basin Naturalist 28:157, 1968. 
USA. Nevada: 4 Jun 1968, JL. Reveal 1139 
(ISOTYPE). ASU 30107. 

pies sal ied L.Reveal, Phytologia 

253. 1989. USA. Idaho: 6 - 1974, JL. Re- 
ce (isoTyPe). ASU 11644 
eoniues ACEAE 
— ees F.T.Pursh var.idahoensis 
vis, een: Leads 1966. USA. Idaho: 
ate 2,RID 29 (ISOTYPE).ASU 93460. 
pare pare JR. Swanson & W.A. Kelly, 
Madrono 34:155. 1987. USA. California: 4 Jul 
1959, 1.R. Swa 490 (ISOTYPE).ASU 152432. 


PRIMULACEAE 
Primula capillaris NH. Holmgren & AH. 
Holmgren, Brittonia 29:313. 1974. USA. Ne- 
vada: 15 Jul 1965, N.H. Holmgren 2154 with 
JL. Reveal (ISOTYPE). ASU 65092 


WLACHI 


Samolus ebracteatus F.W.H.A. Humboldt, AJA. 
Bonpland, K.S. Kunth var. coahuilensis J. 
Henrickson, Southw. Naturalist 28:312.1983. 
Mexico. Coahuila: 23 Sep 1972, J. Henrickson 
7950 (ISOTYPE). ASU 82091. 


RANUNCULACEAE 

Aquilegia scopulorum |. Tidestrom subsp. 
perplexans |.W.Clokey, Bull. S. Calif. Acad. Sci, 
37:3. 1938. USA. Nevada: 16 1936, LW. 
Clokey 7094 (ISOTYPE). ASU 6671 

Clematis ee D. Keil, sal Totes Bot. 
Club 104:10. 1977. Mexico. Coahuila: 4 Aug 
1973, J. eta 1880 (isotype). ASU 
es 


S.Watson f.insigne J.A. 
Ewan, Univ. Colorado Stud. Ser. D, Phys. Sci. 
2:165. 1945. Mexico. Nuevo Leon: 26 Sep 
1934, C.G. Pringle 10178 Siar oe 50614. 
Delphinium subsca , Univ. 
Colorado Stud. Ser. a Phys. a _ ia suis 
Mexico. Mexico: 25 Sep 1934,G 
(isoTyYPE). ASU 42227. 
Ranunculus pentadontus PC. a 2 
Naturalist Hist. Bot. Ser. 1 
ae era te Sep soon ee 
2787 (ISOTYPE). ASU 79075. 


ROSACEAE 

Cercocarpus ledifolius 17. Nuttall var. 
intermontanus N.H. Holmgren, oe 
39:424, 1987, i Utah: 24 May 1 


elas 11298 with PK. Holmgren (ISOT YPE): 
ASU 215437. 


Ivesia hypo B.J.Ertter & J.L. Reveal, Syst. Bot 
14:239. 1989. USA. Oregon: 13 Jul 1975, JL. 
ee 3894 with BJ. Ertter (isorype). ASU 
119052. 


pos oi hl N.H. Holmgren, me 
39:34 7. USA. Utah: 10 Jul 1984, N.H. 
err Holmgren 10623 (isot a ASU 
177195. 

Potentilla basaltica A. Tiehm & BJ. Ertter, 
Brittonia 36:228. 1984. USA. Nevada: 31 Ma 
1983, - re 7770 with M. Lavin (ISOTYPE). 
ASU 1 ; 

Purshia inka C.Schaack, ees 63:301. 

7. USA. Arizona: 11 Apr 7,C. Schaack 

2046 with B. Schaack (\SOTYPES). a 152078, 
154249, 

Rosa stellata R. Weston subsp.abyssa A.M. Phil- 


FAEADARIUM, 1055 


rH Madrono 39:31. 1992. USA. Arizona: 15 
1980, A.M. Phillips 80-103 (IsoTYPE). ASU 
ae 


RUBIACEAE 

Galium collomae J.T. Howell, Leafl.W. Bot. 5:150 
1949. USA. Arizona: 1937, Collom 596 (ISOTYPE). 
ASU 27801 

Hedyotis nigricans (J.B.A.P. de Monnet de 
Lamarck.) FR. Fosberg var. A ean BL. 
Turner, ee 79:1 . Mexico. 
Nuevo L May 58 a eae et al. 
eee rei 1310 

Paederia pringlei J.M. eee Proc. Amer. 
Acad. Arts 39:92. 1903. Mexico. Morelos: Oct 
1902, C.G. Pringle 8682 (ISOTYPE). ASU 23846. 


SALICACEAE 
alix melanopsis T. Nuttall var.gracilis C.R. Ball, 

Proc. Natl. Acad. USA 21:182. 1935. USA. Cali- 
fornia: 10 May nie Ae cae 101 
(ISOSYNTYPE). ASU 7 

Salix melanopsis T. el var. set CR. Ball, 
Proc. Natl. Acad. USA 21:18 35. USA. Cali- 
fornia: 30 May 1931,H.E. si ae Tracy 935 
(ISOSYNTYPE). ASU 73807 

Salix parksiana CR. Ball, Univ. Calif. Publ. Bot. 
17-400. 1934.USA. California: 21 May 1931,1.P 
Parks 983 (ISOSYNTYPE). ASU 72617 

Salix parksiana CR. Ball, Univ. Calif. Publ. Bot. 
17:400. 1934. USA. California: 14 May 1931,/.P 


1} 


FU rks (ISU TINI TPE). ASU 72 
7 ae C.R. Ball, Univ. Calif. Publ. Bot. 17:403. 
4. USA. California: 15 May 1932, CR. Ball, 
: ae & J.B. Tracy 1288 (ISOSYNTYPE). ASU 
72619 


SAPINDACEAE 

Urvillea biternata C.A. Weatherby, Proc. Amer. 
Acad. Arts 45:425.1910.Mexico. Guerrero: 24 
Jul 1907, C.G. Pringle 10380 (ISOTYPE). ci 
50598. 


SCROPHULARIACEAE 

Castilleja aquariensis N.H. Holmgren, Bull. 
Torrey Bot. Club 100:87. 1973. USA. Utah: 11 
Aug 1970, N.H. Holmgren 4726 with PK. 
Holmgren (\sOTYPE). ASU 65095 

Castilleja christii NH. Holmgren, Bull. Torrey Bot. 
Club 100:91. 1973. USA. Idaho: 12 Jul 1966, 

H. Holmgren 2866 with J.L. Reveal (ISOTYPE). 

ASU 65096. 


1056 


este sepa N.H. Holmgren, Mem.New 
ork B ard. 21:46. 1971. USA. Nevada: 3 

if ie N.H. Holmgren 2772 with J.L. Reveal 
(ISOTYPE). ASU 21567. 

Castilleja kaibabensis N.H. Holmgren, Bull. 
We Bot. Club 100:89. 1973. USA. Arizona: 
7 1970, N.H. Holmgren i with PK, 
Holmgren sates ASU 6509 

Castilleja linifolia N.H. ure Brittonia 
28:200. 1 eae Durango: 26 Sep 1973, 
J.L. Reveal 3482 with N.D. Atwood (ISOTYPE). 
ASU 130198 


~~ 


~~ 


Castilleja pringlei M.L.Fernald, Proc. Amer. Acad. 

04. Mexico. Hidalgo: 26 Aug 

1902, CG. Pringle 8666 (ISOSYNTYPE). ASU 
98 


oe) . 

Castilleja revealii NH. Holmgren, Bull. Torrey 
Bot. Club 100:87. 1973. USA. Utah: 24 Jun 
1965, N.H. Holmgren 2017 with LL. Revea 
(ISOTYPE). ASU 65100. 

Castilleja salsuginosa N.H. Holmaren, Bull. 
Torrey Bot. Club 100:83. 1973. USA. Nevada: 
4 Jul 1966, N.H. Holmgren 2764 (ISOTYPE). ASU 
65101 


Mimulus gemmiparus W.A. Weber, Madrono 
21:423.1972.USA. Colorado: 8 Jul 1970, WA. 
Weber & T. Grove 10462 (ISOTYPE). ASU 59439. 

Penstemon acuminatus D. Douglas ex J. 
Lindley var. latebracteatus N.H. Holmgren, 
oe 31:232.1979. USA. Oregon: 21 Ma 

N.H. Holmgren 8741 with PK. Holmgren 
ie VPE), ASU 113745. 

Penstemon angustifolius FT. Pursh var. 
vernalensis N.H.Holmgren, Brittonia 31:229. 
1979.USA. Utah: 25 May 1 - Holmgren 

et al. 8748 (ISOTYPE). ASU 1 

ills barnebyi N.H. Teen Brittonia 

31:226.1979, USA. Nevada: 23 Jun 1977, N.H. 
haingren we Lent : ASU 113746. 

e & J.L.Anderson, 

eae npr toe 11 Jun 
S.L. O'Kane, Jr. & JL. Anderson 2424 
tite ASU 152920. 

Penstemon distans N.H. Holmgren, Brittonia 
32:326. 1980. USA. Arizona: 25 May 1979,N.H. 
Holmgren et al. 9173 (isotype). ASU 113754. 

Penstemon dolius M.£.Jones ex FW.Pennell var. 

uchensnensis N.H. Holmgren, Brittonia 
31:219. 1979. USA. Utah: 25 May 1978, N.H. 
Holmgren et al. 8762 (Isotype). ASU 113748, 


n Cc] 


BRIT.ORG/SIDA 19(4) 


Penstemon goodrichii N.H. Holmgren, 
Brittonia 30:416. 1978. USA. Utah: 25 May 
1978, N.H. Holmgren et al. 8760 (isoTyPe). ASU 
113752 

gre henricksonii R.M. Straw, Madrono 
23:263. 1976. Mexico. Coahuila: 5 Aug 1973, 
J. he 1903 with TL. ve (ISOTYPE). 
ASU 6 

Penstemon S nanifesie N.H. Holmgren, 
sal 30:334. 1978. USA. Nevada: 7 a 

7,N.H. glee 8493 with PK. Holmgren 
rea ASU 113749. 

Penstemon moriahensi N.H. Holmgren, 

a 30:422. 8. USA. Nevada: 17 Jul 
ra rineniee 8609 with PK. Holmgren 
fees ae 3753 

Penstemon mucronatus N.H. Holmgren, 
Brittonia 31:234. 1979. USA. Utah: 25 May 
1978, H. Holmgren et al. 8747 (isotype). ASU 
11375 

Penstemon navajoa N.H. Holmgren, ae 
30:4 978. USA. Utah: 13 Jul 1 H. 
nae 8587 with PK. Holmgren eee 
ASU 113751 


alle 


Penstemon pudicus J.L. Reveal & J.Beatley, Bull. 
Torrey Bot. Club 98:332. 1971. USA. Nevada: 
5 Jul 1971, J. Beatley 12950 (isotype). ASU 


Penstemon rhizomatosus N.H. Holmgren, 
ay 50:162. 1998. USA. Nevada: 4 Aug 
1985 see 10967 with KA. Vincent 
eae ASU 215 

Penstemon ca i var. anestius J.L. 

eveal & J. Beatley, Great Basin Naturalist 
34:230. 1974. USA. Nevada: 20 Jun 1973, J. 
Beatley 13460 with Ackermann (ISOTYPE). ASU 


Penstemon tiehmii N.H. Holmgren, Brittonia 
50:159. 1998, USA. Nevada: 26 Jun 1986, A. 
Tiehm 10658 with S. Crisafulli (ISOTYPE). ASU 


Seymeria deflexa A. Eastwood, Proc. Amer. 
Acad. Arts 44:607. 1909. Mexico. Nuevo Leo 
19 Sep 1907, C.G. Pringle 10398 (\SOTYPE). ii 


50579, 

Stemodia macrantha B.L. Robinson, Proc. Amer. 
Acad. Arts 43:27. 1907. Mexico. Michoacan: 
28 Jan 1907, C.G. Pringle 10356 (isoTYPE). ASU 
50622. 


IWAACRKI 


THEACEAE 

Ternstroemia huasteca B. Bartholomew, 
Phytologia 64:458. 1988. Mexico. Hidalgo: 5 
May 1987,8.Barthol 4035 (Isotype). ASU 
155155. 


THYMELAEACEAE 

Dirca mexicana G.L. Nesom & M.H. Mayfield, 
Sida 16:460. 1995. Mexico. Tamaulipas: 3 Mar 
1995, G. Nesom et al. 7863 (ISOTYPE). ASU 
202295 


TILIACEAE 
aaa se aneanNaee A.A. Bullock, Kew 
:292.1937.Mexico. Mexico: 10 Dec 
— a Hinton 4917 (ISOTYPE). ASU 59079. 
Triumfetta quercetorum A.A. Bullock, Kew Bull. 
1937:295.1937.Mexico. Mexico: 13 Dec 1934, 
G.B. Hinton 6746 (isOTYPE). ASU 59077. 


VERBENACEAE 

Stachytarpheta hintonii H.N. oo 
Phytologia 1:431. 1940. Mexico. Mexico 
Aug 1935, G.B. Hinton 8236 (ISOTYPE). noe 
152532. 


VIOLACEAE 

Viola hemsleyana G. Calderon, Phytologia 
52:101. 1982. Mexico. Mexico: 1 Jul ha ue 
Rzedowski 36144 (ISOTYPE). ASU 3614 

Viola litheon N.H. Holmgren & PK. fn mgren, 
Brittonia 44: 300. 1992. USA. Nevada: 3 Aug 
1980, N.H. Hol oe 9727 with PK. Holmgren 
(ISOTYPE). ASU 215440. 

Viola oxydontis H.E. Ballard, Phytologia 74:359. 
1993. Mexico. Mexico: 15 Jun 1934, G.B 
Hinton 6167 (ISOTYPE). ASU 39975, 


MAGNOLIOPHYTA-MONOCOTS 


AGAVACEAE 
Agave arizonica H.S. Gentry & J. Weber, Cact. 
Succ. J.(U.S.) 42:223,1970.USA. cris : Jun 
1968, J. Weber s.n. (ISOTYPE). ASU 1 
Agave delamateri W.C. Hodgson & . Gace 
See 3:133. 1995. USA. Arizona: 5 Jul 
WC. hora 5478 (\SOTYPE of 2 sheets). 
oo 15984 849. 
ss pet : aie ssp. simplex H.S. 
Gentry, Occas. sa Calif. Acad. Sci. 130:22. 
1978. USA. Ariz 12 Jun 1974, H.S. Gentry 
23404 (ISOTYPE). a 76871. 


1057 


Agave gigantensis H.S. Gentry, Occas. Pap. Ca- 
lif. Acad. Sci 130:63. 1978. Mexico. Baja Cali- 
fornia: 20 Jun 1973,H.S.Gentry 23320 with L.A. 
McGill (isotype). ASU 50795. 

Agave impressa H.S. Gentry, Agaves Cont. N. 
America:146. 1982. Mexico. Sinaloa: 16 Feb 

/H.S. Gentry 23366 (ISOTYPE). ASU 131488. 

Agave jaiboli H.S. Gentry, USDA Agric. Hand- 

book 399:89. 1972. Mexico. Sonora: 1 May 
1965,H.S.Gentry 21177 (isoTYPE). ASU 109644. 

eae abe eee Sida 17:191.1996. 

y 1995, /A. Villarreal 
eta iene (isoTYPE). ASU 235856 ; 

Agave moranii H.S. Gentry, Occas. Pap. Calif. 

cad. Sci. ee 1978. Mexico. Baja Califor- 
nia: 13 Jun 1973, oe need (ISOTYPE of 
2 sheets). ASU 50 

Agave oroensis a paca — Cont. N 

America:294. 1982.Mexico. Zacatecas: 1 Aug 
23592 (ISOTYPE). ASU 128162. 

Agave shrevei H.S. Gentry subsp. magna H.S. 
Gentry, Agaves Cont.N. America 4:451.1982. 
Mexico. Chihuahua: 10 Oct 1973, H.S. Gentry 
233060 with R. Bye (ISOTYPE of 2 sheets). ASU 
131486, 131487. 

Agave titanota HS. Gentry, oie en N. 
America:176. 1982. Mexico. Oax e) 

1967,H.S. ae 22474 with on ane 
ASU 1539 

Agave tiongularis J.A. Jacobi, Zweiter 
Nachtr zu dem _ Versuch” einer 
pee eee Ordnung der Agaveen: 149. 
1869. Neotypified by H.S. Gentry, Agaves 
Cont. N. America:180. 1982. Mexico. Puebla: 
11 Mar 1974, H.S. Gentry 23399 (ISONEOTYPE). 
ASU 131533 

Yucca harrimaniae W. Trelease var. sterilis E. 
Neese & S.L. Welsh, Great Basin Naturalist 
45:789. 1986. USA. Utah: 31 M 
Welsh 18461 (isoTYPE). ASU 108176. 


CYPERACEAE 

Carex ciliaris M.L. Fernald, Proc. Amer.Acad. Arts 
43:61. 1907. Mexico. Hidalgo: 26 Aug 1905, 
C.G. Pringle 10039 (ISOTYPE). ASU 11 

Carex seatoniana LH. Bailey, Bot. Gaz. 25:270. 
1898. Mexico. Hidalgo: 24 Jun 1897,C.G. Prin- 
gle 7452 (\soTYPE). ASU 1396. 


FLAGELLARIACEAE 
Joinvillea ascendens A.1.de Brongniart & JAA. 


1058 


Gris subsp. glabra T.K. Newell, J. Arnold 
Arbor. 50:550. 1969. New Caledonia. Plateau 
Dogny: 1 a — TK. Newell 196 (ISOTYPE, 
oa) ASU 6 

Joinvillea ae A.I.de Brongniart &J.AA. 
ris subsp. samoensis [.K. Newell, J. Arnold 
Arbor. 50:548. 1969.Western Samoa. Upolu: 
26 Oct 1966, IK. Newell 239 (ISOTYPE, spirit 

ASU 67967 


oS 


IRIDACEAE 

cia are pape C. s Weatherby, Proc. Amer. 
Acad. Arts 45:423. 1910. Mexico. Guerrero: 
29 Jul Pee at 10391 (isotype). ASU 
50592 


Tigridia morelosana B.L. Robinson, Proc. Amer. 


BRIT.ORG/SIDA 19(4) 


Brittonia 29:309. 1974. USA. Utah: 13 Jun 1960, 
N.H. Holmgren et al. 13125 (ISOTYPE). ASU 


65093. 

Maianthemum mexicanum A. Garcia-Arevalo, 
Acta Bot. Mex.1 7:19. 1992. Mexico. Durango: 
16 Jul 1991, A. Garcia & S. Acevedo 1008 
(ISOTYPE). ASU 193190. 


POACEAE 
Bealia mexicana F.L. Scribner, In WJ. Beal, 
Grasses N.America 2:267, 1896, aad 
PM. Peterson in Madrono 36:262. | 
Mexico. Chihuahua: 7 Oct ie CG. ee 
819 (ISOLECTOTYPE). ASU 363 
Hordeum arizonicum G. pan Meciene 10:16. 
1949. USA. Arizona: 3 Apr 1904, J.J. Thornber 


Arts 43:21.1907.Mexico.Morelos:5 Sep 

1908, C.G. Pringle 13657 (isOoTYPE). ASU 50585. 
LILIACEAE 

Allium passeyi N.H.Holmgren & A.H.Holmaren, 


536 (ISOTYPES). ASU 221231, 226002. 

Muhlenbergia xerophila C.O. Goodding, J. 
Wash. Acad. Sci. 30:19. 1940. USA. Arizona: 22 
Sep 1937,L.N.Goodding 262-M (\soTyPe). ASU 
208587, 


ACKNOWLEDGMENTS 


The authors thank Elinor Lehto, Thomas Daniel, and Charlotte M. Christy for 
initiating data entry, and Robin Schoeniger and Ed Gilbert for help with for- 
matting of the data base. Raul Puente-Martinez helped with the Spanish trans- 
lation. We also thank Michael Chamberland and one anonymous reviewer for 
helpful comments on the manuscript. We also want to thank the Department 
of Plant Biology, Arizona State University for financial support. 


=— 


REFERENCES 

Anonymous. 1966. New herbarium abbreviations. Taxon 15:334. 

Anonymous. 1974,American Society of Plant Taxonomists, systematic resources in America, 
part 1, Survey and Preliminary Ranking. 

Davis, H.B. 1936.Life and work of Cyrus Guernsey Pringle. Free Press Printing Co.,Burlington, 
ae 

GreuTer, W., J. MCNEILL, FR. Barri, HM. Burpet, V. DEMOULIN, TS. Fitcueiras, D.H. Nicotson, PC. Siva 
J.E. Skoc, P. TREHANE, NJ. TurLAND, and DLL. HawskwortH (eds. and compilers). 2000. Interna- 
tional code of botanical nomenclature (St. Louis Code). Adopted by the Sixteenth In- 
ternational Botanical Congress, St. Louis, July-August 1999. Regnum Veg. 138. 

Hopkins, EJ.and A. THomas, Jk. 1960. The Arizona State University story. Southwest Publish- 
ing Co.,Inc., Phoenix, AZ. 

IcKERT-BOND, S.M. and DJ. Pinkava. 2000. Vascular plant type specimens in the Arizona State 
University Herbarium. J. Arizona-Nevada Acad. Sci. 35 (Proc. Suppl.): 9 (Abstr,). 

Kearney, T.H.,R.H. Peestes, and Coit asorarors. 1960. Arizona flora. University of California Press, 
Berkeley, Los Angeles. 


1059 


Mason, C.T., L.R. Lanorum, J.R. Reeoer, DJ. Pinkava, and R. Van Devenoer. 1992. A new flora of 
Arizona in preparation. J. Arizona-Nevada Acad. Sci. 26:1. 

Rusinore, M.W. 1973. From normal school to university. Unpublished report, Arizona State 
University, Hayden Arizona Collection. 

Tuomas, Jr, A. 1960. Arizona State University: A documentary history of the first seventy- 
five years 1885-1960. Vol. |: The formative years 1885-1900. Vol. Il: From normal school 
to teachers college 1900-1933. Vol. Ill: From teachers college to university 1933-1960. 
Vol. lV: Growth and development statistics and general information 1885-1960. Un- 
published report, Arizona State University, Hayden Arizona Collection. 

TRAUTH-Nare, A. and D.J. Pinkava. 2000. Frederick Mortimer Irish: Founder of the ASU her- 
barium. J. Arizona-Nevada Acad. Sci. 35 (Proc. Suppl.): 9 (Abstr). 


1060 BRIT.ORG/SIDA 19(4) 


Book Notices 
BeRNARD K. MaLoney. (editor). 1998. Human Activities and the Tropical Rainforest: 
Past, Present, and Possible Future. (ISBN 0-7923-4858-3, hbk.). Kluwer 
Academic Publishers, PO. Box 17,3300 AA Dordecht, The Netherlands. (Orders: 
Kluwer Academic Publishers Group, 101 Philip Drive, Norwell, MA 02061, 
U.S.A. US$142.00, hbk. 206 pp., figures, 61/2" x 91/2" 


led -ontributors; Foreword. Professor Sir Ghillean Prance 


Table of Contents: Pref 
1) Introduction. Bernard K. isloney 
2) Landscape sensitivity in the Humid Tropics: A ¢ 
3) Human activity and the Tropical Rainforest: Ae the Soils the forgotten component of the 

Ecosystem? Stephen Nortcliff 

4) The Long-Term History of Human Activity and Rainforest Development. Bernard K. Maloney 
5) Indigenous Knowledge of the Rainforest: Perception, Extraction, and Conservation. Roy F Ellen 
6) History and Density of Middle American Forests: The Inheritors of the Mayan Landscape. 


| isal. Michael F Thomas 


Peter A. Furley 
7) Social, Economic, and Political Aspects of Forest Clearance and Land-Use planning in Indo- 
nesia. Alastair I. F 
8) Diversity Destroyed? The Monoculture of | 
9) Modelling Climatic pees of Future Rainforest Destruction. Kenneth McGuffie et al 


10) Conclusion. Bernard K. Malon 


‘Eucalyptus. Christopher J. Barrow 


This isan importa gof the humani impact on Neotropi- 
i iology graduate courses. 


cal Rainforests. ea work gy and conse 
I broad hic scl d tal I bj acs an interdisciplinary 


Cr oO Oo a 
approach. Future editions would be improved by including a subject index. 1 would recommend this 
paok & to researchers interested in the human aspect to biodiversity and ecosystem conservation. 
nD. Janni, Botanical Research Institute of Texas, Fort Worth, TX 76102-4060, 


nee ore. 


S.HJ.V. Rarpanarivo, J.J. LAvRANos, A.J.M. LEEUWENBERG, and W. Roost. 1999. 
Pachypodium (Apocynaceae). Taxonomy, Ecology and Cultivation. (ISBN 
90-5410-485-6, hbk.). A.A. Balkema Publishers, Old Post Road, Brookfield, 
VT 05036-9704, wwwbalkema.nl, ball @ball nl, 802-276-3837 fax, 
31-10-4135947). $48.00, 128 pp, 80 color photos. 


This taxonomic monographic revision of eons m (Apocynaceae) is based on studies of living 
plants both in the wild and in cultivation. The genus is comprised of 23 species, eight of which are 
endemic to Madagascar. Other species occur in continental Africa. There is the generic descriptic 

followed by a key to species is followed by a treatment of species. Each species is treated systemati- 
cally and nee a line drawing, color ae: ae a distribution map. Some of the line drawings 
are a little on the rough side, however, the 80 Ss saree (pl. ates) are very nice. | find one new com- 
r) Raps 7o, comb. nov. The book is rounded 
catae, a nice index. 


— 


bination presented, Pachyupodium gracilius (H 
out with a chapter on Pachypodium in een followed by an index of e 
—Barney Lipscomb, Botanical Research Institute of Texas, Fort Worth, a 76102-4060, U.S.A., 


barney@britorg. 


SIDA 19(4): 1060. 2001 


PLANT SPECIES-AREA RELATIONSHIPS IN TEN 
NORTH CENTRAL TEXAS PROTECTED NATURAL AREAS 


Monica Granados Robert J.O’Kennon 
Department of Biology ‘Botanical Research Institute of Texas 
Texas Wesleyan University 9 Pecan Street 
1201 Wesleyan Fort Worth, TX 76102-4060, U.S.A, 
Fort Worth, TX 76105-1536, U.S.A bokennon@brig.org 


Bruce F. Benz! 
Department of Biology 
Texas Wesleyan University 
1201 Wesleyan 
Fort Worth, TX 76105-1536, U.S.A 
benzb@txwes.edu 


oe. 
= 


ABSTRACT 
The study of species-area fel anonstipo in protected natural areas (PNAs) is an effective tool for de- 
signing nature reserves and manag ical diversity. Ten North Central Texas PNAs were stud- 


ied to understand how plant species cine is related to PNA size. The species-area model was ap- 
plied to total plant species, native plant species, selected native speciose plant families and invasive 
plant species. Results indicate that area is a significant predictor of species richness (1° = > 0.60) for 
both total species and native species in North Central Texas PNAS. Habitat diversity as measured by 
topographic relief and topographic abruptness is also shown to be a significant predictor of plant 
species richness in North Central Texas PNAs. Introduced species richness could not be piediciee 
from area alone but was explained by perimeter and perimeter/area ratio and measure ol 
habitat diversity. The estimates of z values range from 0.15 to 0.30, while c ranges an 0.60 to 1.42 
for species-area relationships, both of which fall within the range of values estimated by previous 
research. Such estimates allowed us to evaluate effective management schemes for North Central 
Texas PNAs plant diversity. The results of this research pent us to examine thes invasion . exotic 
ae in Texas PNAs and to predict how such invasions will onser- 


E 


vation management practices are not implemented. 


RESUMEN 


El estudio de las relaciones entre el area y el 
herramienta efectiva para el diseno de reservas y el manejo ge la diversidad biologic. ne areas 
naturales protegidas del norte del estado de 


J 

especies vegetales esta relacionada con el tamano del area. El edie especies-area fue ey al 

numero total de especies vegetales, las especies nativas, ciertas familias de plantas con metas 
; ae Tih a daca! sd 


i L 
de la riqueza de especies (r? > 0.60) para el Hei total d el numero de especies nativas 
d d 4 les protegid diversidad del habitat que se midio por medio del total 
de relieve topografico identado d : cael también es un predictor significativo de la 


y! 
riqueza vegetal en estas areas. El ntimero de especies exdticas no se pudo predecir por el area 
pulcarenis sino quese pudo por medio de! Reno « sone area ae . prepercion pee: por area, y 


alor de z del 


SIDA 19(4): 1061 — 1072. 2001 


1062 BRIT.ORG/SIDA 19(4) 


nae de seceouens y Wilson son de 0. 5 a 0.30, mientras que la c varia desde 0.60 a 1.42 para las 
area. Ti os permiten evaluar los esquemas de manejo de la 


diversidad veseual de las areas naturales ce. del norte de Texas. Los resultados de esta 
investigacion nos permiten examinar las razones por las cuales las especies exoticas invasoras han 
colonizado las areas naturales protegidas y predecir como tales invasiones reduciran la riqueza de 
especies nativas si no se implementan pautas de manejo para su conservacion. 


INTRODUCTION 


The conservation of biodiversity is a priority issue for ecologists and conserva- 
tionists worldwide (Myers & Knoll 2001; Novacek & Cleland 2001). Asa result, 
protected natural areas (PNAs) have been created in order to protect, manage 
and monitor native and endemic biota from habitat destruction and the inva- 
sion of introduced and transient species. Such invasions have lead conserva- 
tionists and ecologists to engage in management programs to preserve natural 
habitats and prevent extinction. The theory of island biogeography (MacArthur 
& Wilson 1967) has been used on many occasions in order to measure species 
richness as well as recommending practices for ecological restoration (Hanski 
and Matts 1997; He and Legendre 1996; Lawrey 1991; Lomolino et al. 1989). Here 
we contribute to the existing body of island biogeographic research by exam- 
ining the floristic richness of ten protected natural areas in North Central Texas. 

Island biogeography attempts to document the equilibrium existing be- 
tween colonization and extinction rates of species on islands. The well-estab- 
lished theory developed by MacArthur and Wilson (1967) uses this equilibrium 
theory as a focal point for understanding biotic richness on islands and pro- 
tected areas surrounded by urban, agricultural and forestry developments. This 
equilibrium is based upon empirical evidence that demonstrates a positive cor- 
relation between island size and species richness (Meffe et al. 1997:132). Island 
biogeography theory explains island biological richness based on the degree of 
isolation usually measured as the distance from the nearest source habitat. Other 
factors that influence species richness on islands include colonization rates 
based upon distance from a source habitat and species turnover based on habi- 
tat availability. 

Variation in species richness is due in part to the rate at which species can 
successfully colonize PNAs. Colonization rates can differ and usually depend 
on the vagility of species able to occupy an area and the distance between is- 
land and source habitat. Short-lived species usually colonize areas that have 
been cleared or severely disturbed. Many PNAs experience high colonization 
rates because they are adjacent to disturbed source habitats. Colonists of pro- 
tected areas are commonly exotic species that are managed or introduced into 
adjacent areas subject to forest exploitation or agriculture (Alverson et al. 
1994:83) 

Species turnover refers to the balance of immigration and extinction of 
species continuously enlarged by the arrival of new species from mainland 


GRANADOS ET AL.. SPECIES. BEA Hes 


sources and their continuous reduction by extinction through competition for 
space (Brown & Lomolino 1989). In PNAs and on islands close to continental 
source habitats, species turnover is greater due to higher colonization rates. 
Oceanic islands that are the same size but more isolated support lower species 
richness because of lower species turnover due to lower colonization rates. 

The use of island biogeography in PNAs has been a subject of considerable 
interest for several reasons. The first is that many protected natural areas are 
effectively islands of a natural habitat in a sea of human dominated ecosys- 
tems. Increasingly, PNAs are located in areas where transient species have easy 
access for invasion. Secondly, these islands of natural habitat usually represent 
small isolated areas that vary in shape and size which are important criteria 
particularly relevant for designing and establishing reserves. Finally, the less 
isolated or the closer an island is toa mainland or source habitat, the more spe- 
cies that island is destined to harbor. Natural dispersal explains the diverse biota 
of oceanic islands while anthropogenic encouraged dispersal explains the sig- 
nificant encroachment on protected areas by invasive species leading to the 
loss of native species and the increasing numbers of naturalized exotic species 
in such areas. Island biogeography has also been used to determine the mini- 
mum effective size required for reserves, to characterize community structure, 
to manage species richness, to measure the effect of disturbance on communi- 
ties, to estimate extinction rates and for defining conservation biology. 

MacArthur and Wilson’s (1967:16) initial formulation of the theory of is- 
land biogeography (S=cA*, where S = species number, A= area and c and z are 
constants) indicated that the value of z usually lies between 0.2 and 0.35. Since 
then, studies have shown that areas located on continents or large islands have 
z values that range from 0.17 to 0.57. Studies done by Chown et al. (1998:564) 
empirically derived z values can range from 0.1 to 0.45 for species on oceanic 
islands. Crawley and Harral (2001) derived z values ranging from 0.18 and 0.57 
for spatial scales ranging from 0.01 m? to 110 ha for species in southern En- 
gland. We add empirical support of estimates of z that are in line with those 
obtained from other continental areas. 

Simberloff (1988) and others have argued that area is an effective predictor 
of species’ richness because it is a poey for habitat diversity. As area increases, 
so too does habitat diversity. Likewise, other researchers have demonstrated that 

sland elevation and island complexity are significant predictors of species’ rich- 
ness in addition to area . Habitats are an important component of protected 
areas because they represent the diverse combinations of resources and envi- 
ronmental conditions that plant species can inhabit. The number of differing 
habitats in a park or reserve can vary ines and can result in greater 
species’ number per area. Examples of this have been studied by Williams (1943) 
who showed that as area increases, so does the diversity of physical habitats. 
Simberloff (1988) also pointed out that the majority of the species-area rela- 


1064 BRIT.ORG/SIDA 19(4) 


tionships documented are in fact accounted for by the fact that larger sites have 
more species not only because the area is greater but because larger sites have 
more habitats than smaller sites and that habitat diversity by itself can explain 
species richness better than area alone. The results of these studies show that 
habitat diversity is an important component of area that determines species 
richness so we have made the effort to estimate habitat diversity in North Cen- 
tral Texas protected areas as well. 

Many PNAs have lost habitat diversity due to fragmentation and homog- 
enization. Fragmentation is generally accounted for by human activities that 
disturb or modify natural habitats resulting in the loss of native biota. Invasion 
of exotic species through habitat disturbance or modification tends to exacer- 
bate the effects on native biota causing further loss of native species. This en- 
croachment on natural landscapes is promoted by agricultural and forestry 
practices as well as urbanism. The result of such invasions leave PNAs progres- 
sively more disturbed as exotic species become more numerous and abundant 
and losses of native biota increase (Mooney & Cleland 2001). 

Edge effects are particularly important in the loss of native species in PNAs. 
The outer boundary of any habitat island is subject to external factors that 
modify natural conditions to create novel habitats that can be invaded by spe- 
cies with general habitat requirements (Meffe et al. 1997:294). Edge effects can 
be especially detrimental because these effects cause PNAs below a certain size 
or with a significant edge to area relation to lack sufficient natural habitat core 
area that many native species might need in order to survive. As the natural 
core area decreases in size, native species are lost and replaced by invasive spe- 
cies that dominate the edges and colonize the core. Invasive species’ habitat re- 
quirements are often more generalized which allow them to readily invade the 
protected area’s edge. A preponderance of evidence suggests that edge effects 
play a critical role in PNA management. We examine edge effects by examining 
PNA perimeter as a predictor of PNA floristic composition. 


METHODS 


The ten protected natural areas studied are located within the Cross Timbers 
and Prairies and Blackland Prairies regions in north Central Texas (Fig. 1). The 
Blackland Prairies consist of about 11,500,000 acres with dark-colored calcare- 
ous clays interspersed with gray acid sandy loams (Correll & Johnston 1970; 
Diggs et al. 1999). The flora typically consists of Bouteloua curtipendula, 
Bouteloua hirsuta, Sporobolus asper, Buchloe dactyloides, Bouteloua rigidiseta, 
Ulmus crassifolia, and Maclura pomifera and is considered to be true prairie. 
According to Diggs et al. (1999) the Cross Timbers and Prairies vegetation types 
encompass roughly 6,879,662 hectares (17,000,000 acres), with 404,686 of those 
hectares (1,000,000 acres) designated as the East Cross Timbers, 1,214,058 hect- 
ares (3,000,000 acres) designated as the West Cross Timbers and 2,630,459 hect- 


— 


GRANADOS FT AL.. SPECIES. ADCAC 1065 


— 


me 


eae 7 


hate + nf +h 


Fic. 1. Location of 10 protected natural areas in north central Texas that are the subject of the p t study. Scale is 
approximately 1:400,000 hi { the top of the page; gray ban develoy e chodad 
voirs, gray lines are highways, dash-dotted lines are streams. 


ares (6,500,000 acres) as Grand Prairie. These areas range from savanna to dense 
brush and which consist of Oaks and other woodland vegetation with neutral 
to slightly acidic clay soils over limestone. The flora consists of Elymus 
canadensis, Erioneuron pilosum, and other plants found in the Poaceae family 
(see Diggs et al. 1999). 

Existing plant lists from each of ten PNAs in North Central Texas were 
analyzed. Six variables were tabulated from each list: total species number, na- 
tive species number and introduced species number, as well as the number of 
native legumes, composites and grasses. The number of species in each of these 
three families was incorporated in our analysis because they are the most spe- 
cies-rich families of the North Central Texas flora and therefore might be sig- 
nificant predictors of habitat loss (Leach et al. 1991:34). These variables were 
considered the dependent variables in the regression analyses. 

Native plant species are those that have been present in a particular area 
before Columbus (Diggs et al. 1999-11). All other plants that have arrived since 
then and which are reproducing freely in nature are considered naturalized 
and were included in the total species count. Introduced species were defined 


1066 BRIT.ORG/SIDA 19(4) 


as any species that is not of North American origin and has been introduced 
since Columbus (Diggs et al. 1999:12). Introduced species recorded in these PNAs 
include exotic ornamentals (e.g. Iris spp. Narcissus spp., Nandina domestica, 
Photinia serratifolia, Ligustrum spp. and Wisteria spp.) that have been shown 
to be invasive and capable of becoming serious pests (Diggs et al. 1999:60). How- 
ever, horticultural species found around buildings or foundations were not in- 
cluded in the introduced species’ counts. 

Topographic maps were used to locate and examine habitat diversity in 
each PNA. The maps used are scale 1:24,000 USGS quads for Euless, Hurst, west 
Cleburne, Covington, Blum, Meridian, Mineral Wells, Benbrook, Haltom City, 
Lake Worth, Arlington, Duncanville, Cedar Hill, and Britton, Texas. Habitat di- 
versity was estimated using two proxy measures: overall topographic relief and 
the maximum topographic abruptness over 0.75 mile transects situated in or- 
der to maximize the number of contour intervals encountered. The perimeter 
of each PNA was measured on the same topographic maps. The size of each 
PNA was based upon data provided by both private and/or public landowners 
and land-managers. 

The model was estimated using linear regression after log transformation 
of dependent and independent variables. Statistical analysis was accomplished 
using SPSS and Sigma Plot. 


RESULTS 


The PNAs range in area from 20.2 hectares (50 acres) to 1416 hectares (3500 
acres). Their perimeters range from 1609 m (< 1 mile) to more than 33,796 m 
(>21 miles). The number of contour lines intersecting 1207 m (0.75 mile) 
transects ranged from | to 22 for topographic abruptness, and topographic re- 
lief ranged from 15.25 to 61 m (50-200 ft) (Table 1). 

The Asteraceae, Fabaceae and Poaceae were the most speciose families in 
all ten PNAs. Total species richness ranged from 160 species to 592. Native spe- 
cies richness ranged from 144 to 517; introduced species richness ranged from 
I5 to 75. Asteraceae species richness ranged from 31 to 76, Fabaceae ranged 
fromfour to 46 and Poaceae ranged from 15 to 59 (Table 2, Fig. 2). Total species 
richness increases with PNA size. 

Area is a significant predictor of total species richness in North Central 
Texas PNAs both for total and native species richness as well as for species num- 
ber in the Fabaceae, Poaceae and Asteraceae (r* = 0.4) (Table 3). Total species 
richness could be explained 52 percent of the time by area. The estimate of z for 
total species was 0.18 and c was estimated as 2.03. Tandy Prairie, Vivian Mal- 
one, and the Fort Worth Nature Center have higher than average species rich- 
ness, while other PNAs such as All Saints and Benbrook exhibit low species 
richness (Fig. 2). 


GRANADOS ET AL., SPECIES iy 


Taste 1.Physical and geographic characteristics of ten protected natural areas in north central Texas. 


PNA* Areain hectares Perimeterinmeters Elevationin Topographic Abruptness 

(acres) (miles) m asl Relief 

(AREA) (PERI) (ft asl) (REL) (ABR) 
All Saints 24.3 (60) 1287 (0.8) 229 (750) 20 ] 
Benbrook 20.2 (50) 1287 (0.8) 212 (694) 48 2 
Calloway Branch 40.5 (100) 3058 (1.9) 189 (620) 40 3 
Cedar Hills State Park = 739 (1826) 23,818 (14.8) 244 (800) 130 14 
FW Nature Center 1416.4 (3500) 34,601 (21.5) 181 (594) 101 10 
Lake Mineral Wells 1329 (3283) 18,829 (11.7) 259 (850) 160 12 
Meridian 204 (505) 6598 (4.1) 317 (1040) = 79 5 
River Legacy 393 (972) 11,265 (7.0) 183 (600) 73 8 
Tandy Hills 64.7 (160) 5150 (3.2) 177 (580) 77 13 
Vivian Malone 58.7 (145) 2253 (1.4) 229 (750) 72 9 


* Variable acronyms in parentheses. 


Native species number (Table 3) is explained by area 54 percent of the time. 
The estimate of z based on native plant species richness is 0.17, while cis L99. 
Both Tandy Prairie and Vivian Malone display high species richness in comparison 
to other areas, while Benbrook and All Saints display low native species richness. 

Variation in species’ number for grasses was explained by area 62 percent of 
the time. cis 1.34 and zis 0.14. The relationship between grass species richness 
and area indicates that Tandy Prairie has one of the highest numbers of grass 
species, followed by Calloway Branch, Vivian Malone and Meridian. Benbrook, 
All Saints and River Legacy have relatively few grass species by comparison. 

Asteraceae species’ richness could be explained by area 50 percent of the 
time. The estimate of cis 1.30, while zis 0.15. The Asteraceae are most abundant in 
Tandy, Vivian Malone, the Fort Worth Nature Center and Cedar Hills State Park. 

Variation in Fabaceae species’ richness was explained by area only 46 
percent of the time. The estimate of cis 0.60 and zis 0.30. PNAs such as Tandy 
Hills, Vivian Malone, Calloway Branch and Benbrook have a greater than an 
average number of legume species. 

Area was not a significant predictor of introduced species’ richness (F = 3.5, 
p< 0.1, r2 = 0.30) (Table 3); however, the number of introduced species could be 
predicted from perimeter (F=6.4, p<0.04, 1r7=0.45). A perimeter/area ratio vari- 
able wasalsoa significant predictor of introduced species richness (F=8.6, p<0.05, 
r2 =0.52). Protected natural areas that have high numbers of introduced species 
have a large perimeter and a high perimeter/area ratio (Figure 2). This indicates 
that areas possessing more ed unit area have greater numbers of introduced 


1068 BRIT.ORG/SIDA 19(4) 


Taste 2. Floristic characteristics of ten protected natural areas in north central Texas. 


PNA* Total Native Introduced Number Number Number 
Species Species Species Asteraceae Fabaceae Poaceae 
(TSP) (NS) (IS) (AS) (FS) (PS) 
All Saints** 160 144 16 29 4 15 
Benbrook 184 169 15 31 15 16 
Calloway Branch 260 230 30 42 20 28 
Cedar Hills State Park 434 375 59 64 33 44 
FW Nature Center 592 517 75 76 46 55 
Lake Mineral Wells 392 365 27 61 46 40 
Meridian 280 258 22 35 26 44 
River Legacy 277 250 27 40 18 26 
Tandy Hills 437 380 57 63 35 59 
Vivian Malone 382 344 38 54 27 37 


*Variable acronyms in parentheses. ** List of plants provided by R. Sanders (BRIT) 


species. The z values calculated from perimeter and perimeter/area ratio range 
from 0.3 to L1 respectively. Introduced species richness is greatest in Tandy Prairie, 
Cedar Hills State Park, Fort Worth Nature Center, and Vivian Malone, while 
the other areas have a lower than average number of introduced species. 
Habitat diversity as estimated by topographic abruptness and total topo- 
graphic relief was a significant predictor of both total species and native species 
richness. A topographic profile that included the number of contour lines in- 
tersecting 0.75 mile transects explained a significant amount of the variation in 
total species (F=37.8, p<0.001, r* = .82) and native species’ richness (F=40.9, p< 
0.001; r*=0.84). Overall topographic relief is alsoa significant predictor of total 
species’ richness (F=14.9, p<0.005, r2=0.65) and native species richness (F=17.4, 
p<0.003, r2=0.83).¢ ranges from 1.14 to 2.2 and z varies from 0.41 to 0.55. 


DISCUSSION 


The estimates of cand z obtained from the plant species and area of these ten 
protected natural areas in North Central Texas occur within the range estab- 
lished by previous r h. MacArthur and Wilson (1967) suggest that z should 
vary between 0.2 and 0.35 on isolated islands and from 0.17 to 0.19 in continen- 
tal areas. Other researchers (Chown et al. 1998) have shown that z ranges from 
0.10 to 0.45 for the flora and fauna on oceanic islands. Estimates of z for North 
Central Texas range from 0.14 to 0.30. Estimates of c for these ten North Central 
Texas protected areas range from 0.58 to 2.03. MacArthur and Wilson showed 
that cshould vary with the degree of isolation and the taxonomic groupings of 
organisms studied. 

Area predicts native species’ richness and total species’ richness. Habitat 


GRANADOS ET AL., SPECIES 1069 


FWNC @ 
ee 
3 Bs M@ Tandy Hills Cedar Hills 
9287 @ VivianMalone 
2. a Lake Mineral 
” 25 
SS : CallowayBrch = _ WM River Legacy 
2 2.4 = Meridian 
— = 
ey) = 
o 2.3— 
= 7 g Benbrook 
224 ~~ ge All Saints 
——— — 
2.0 2.5 3.0 3.0 
log (Area) 
~~ FWnc @ 
3g 1.8 . Tandy Hills Cedar Hills g 
Oo i y a 
rb) 7 
a. - a 
3 16- —— 
3 cal VivianMalone Bi a 
= . CallowayBrch a Lake Mineral 
ae. River Legac 
Oo 1.44 . gacy # & 
= : ce w Meridian 
>» | ~~, 
re} _| g All Saints 
= 12 J = w Benbrook 
¥ v | ¥ ad = be inl ¥ ¥ | i v T ¥ T ¥ ¥ 
0.0 0.1 0.2 0.3 
log (Perimeter/Area) 
Fic. 2.Ab f d R ion | provid jin Table 3. Below, regression 


plot of introduced species and perimeter/area. 


diversity as estimated by topographic abruptness and overall topographic re- 
lief explain species’ richness for both native and total species. Estimates of habi- 
tat diversity appear to better explain species richness than area. Our results 
confirm those of other researchers who show that habitat diversity is generally 
a better predictor of species richness presumably because greater topographic 


1070 BRIT.ORG/SIDA 19(4) 


Taste 3. Regression equations and test statistics for floristic richness indicators of ten protected 
natural areas in north central Texas. Variable acronyms can be found in Tables 1 and 2. 


TSP = 2.03 + 0.18 (AREA 


P=052 F=88 p<005 


NSP = 1,99 + 0.18 (AREA) r=055 F=96 p < 0.05 
ASP = 1.31 + 0.14 (AREA) r=046 F=68 p < 0.05 
LSP = 0.58 + 0.30 (AREA) r=046 F=69 p < 0.05 
PSP = 1.06 + 0.18 (AREA) r = 0.40 = 5:) p< 0.05 
ISP = 1.02 + 0.19 (AREA) r=030 F=3.5 ns 

ISP = 1.30 + 0.30 (PER) r=045 F=64 p < 0.05 
ISP = 1.30 + 1.09 (PER/AREA) r=052 F=86 p< 0.05 
TSP = 2.20 + 0.42(ABR) r=082 F=378 p<0.001 
NSP = 2.14 + 0.41(ABR) r=084 F=409 p<0.001 
ISP = 1.13 + 0.48 (ABR) r=062 =130 p<0.01 
TSP = 1.49 + 0.55(REL) r=065 F=149 p<0,.005 
NSP = 1.40 + 0.55 (REL) r=0.83 =174 p<005 
ISP = 0.51 + 0.54 (REL) rm=035 F=42 ns 


relief provides for a greater variety of habitat types offering more opportuni- 
ties for a greater number of species (Simberloff 1988). 

Introduced species’ number could not be explained by area. Both perim- 
eter and perimeter/area ratio explain a significant amount of the variation in 
introduced species across protected areas. The PNAs with a high number of 
introduced species have a high perimeter/area ratio suggesting some of these 
protected areas are fragments of natural habitat with permeable edges. High 
perimeter to area ratios increase PNA’s permeability to invasive species. The 
easy invasion of introduced species, many of which are weedy intruders that 
are often wind dispersed and/or horticultural exotics that produce an abun- 
dance of enticing seasonal fruits that are dispersed by birds and omnivorous 
mammals are not only invasive but pernicious. 

Our analyses demonstrate that management plans are necessary to con- 
serve the native species still existing in the protected natural areas of north 
central Texas by preventing the invasion of non-native species. It is possible 
that for every introduced species that is successfully surviving in these PNAs, a 
native species might be displaced. The ongoing competition between native and 
introduced species will continue until these areas are managed appropriately 
or the amount of edge relative to area diminishes. These areas were probably 
once subject to periodic fires (Leach & Givnish 1991) which allowed the native 
biota to colonize open areas and survive more abundantly than introduced spe- 
cies in north central Texas. Such periodic fires have worked in the past to create 
and maintain suitable environment for native species and a detrimental envi- 
ronment for most introduced species. Most of the invasive species that are now 


GRANADOS ET AL., SPECIES AREA 1071 


successfully colonizing these areas are not only successful, but they are natu- 
ralizing relentlessly because periodic fires are no longer allowed. 

It has long been argued that the optimal shape for a refuge should be circu- 
lar and that small linear protected areas have a greater chance of becoming 
significantly modified by invasive apecics (Simberloff 1988:479) which change 
protected natural areas into protected anthropogenic areas. Asa result, park or 
reserve shape, the amount of edge relative to area, and park size continue to be 
debated because although the model can predict species richness based upon 
area, it fails to incorporate park shape and edge-area considerations. In this study 
we have shown that many of the areas have an existing shape and size that grant 
increased access to invasive plant species. 

Appropriate management of these PNAs in North Central Texas should 
take two courses of action. One focus would be to increase park size, especially 
for those areas that exhibit high perimeter/area ratio, including Tandy Strat- 
ford Prairie, Vivian Malone and the Fort Worth Nature Center. By doing this, 
edge effects will diminish by increasing the core area where many native and 
some endemic species still thrive. Adding area to these PNAs could potentially 
buffer them from edge effects (boundary permeability) and enhance their abil- 
ity to maintain native species’ richness. Adding area is perhaps difficult because 
of competition from more lucrative land-use options. An alternative to increas- 
ing their size might be to impose zoning restrictions in land developments ad- 
jacent to these PNAs. Restrictions on planting native species in urban areas found 
near PNAs could diminish the influx of non-native species that typically in- 
vade and colonize such areas (e.g. Nandina spp., Photinia sp., Ligustrum spp). 
Coupling restrictions on urban plantings and increasing the size of PNAs would 
insure that native species in PNAs would be protected better than the smaller 
reserves in the urban landscape because larger, protected sites are more likely 
to be able to accommodate disturbance than small, unprotected reserves. 

In conclusion, the conservation of PNAsisan ongoing process. Disturbance 
takes place inside and out, succession follows and colonization by both native 
and introduced species will continue. One of the most critical concerns should 
be to determine which species will be successful in the long run. Incorporating 
appropriate conservation management schemes will make it possible to lower 
extirpation rates and insure the native species’ likelihood of survival. Protected 
natural area size, shape, habitat diversity, edge effects, proximity to disturbed 
communities, and distance from source habitats all play important roles in 
maintaining the natural biota in PNAs. We believe that the use of the species- 
area model will aid in managing and controlling PNAs’ species composition 
and give the public an idea of how invasion of introduced plant species can be 
harmful to such areas now and in the future. 


1072 BRIT.ORG/SIDA 19(4) 


REFERENCES 


Averson, W.S.,W. KUHUMANN and D.M. Watter. 1994. Wild forests: Conservation biology and 
public policy. Island Press, Washington, D.C. 

Brown, J.H. and M.V. Lomouino. 1989. Independent discovery of the equilibrium theory of 
island biogeography. Ecology 70:1954 -1957. 

CHown, S. L., N.J.M. Gremmen and K.J. Gaston. 1998. Ecological biogeography of southern 
ocean Islands: Species-area relationships, human impacts and conservation. Amer. 
Naturalist 152:562-575. 

Corrett, D.S. and M.C. JoHNsTON. 1970. Manual of the vascular plants of Texas. Texas 
Research Foundation, Renner 

Crawtey, MJ. and J.E. Harrat. 2001. Scale dependence in plant biodiversity. Science 291: 
864-868. 

Dices, G., B. Liescome, and R. O’Kennon. 1999. Shinners & Mahler's illustrated flora of north 
central Texas. Botanical Research Institute of Texas, Fort Worth. 

Hanski, |. and G. Mars. 1997. Uniting two general patterns in the distribution of species. 
Science 275:397-399. 

He, F.and P. Lecenore. 1996. On species - area relationships. Amer. Naturalist 148:719-737. 

Lawrey, J.D. 1991. The species area curve as an index of disturbance in saxicolous lichen 
communities. Bryologist 94:377-382. 

Leach, M.and TJ. GivnisH. 1996. Ecological determinants of species loss in remnant prairies. 
Science 273:1555=1558. 

Lomo.no M.V.,J.H. Brown, and R. Davis. 1989. Island biogeography of montane forest mam- 
mals in the American Southwest. Ecology 70:180-194. 

McArtHur, R. and E.O. Witson. 1967. The theory of island biogeography. Princeton Univer- 
sity Press, N.J. 

Merre, G.K,, CR. Carrot, and Contrisutors. 1997. Principles of conservation biology, 2nd ed. 
Sinauer Associates, Inc., Sunderland, MA. 


Mooney, H. A. and E.E. Cietanp. 2001. The evolutionary impact of invasive species. PNAS 
98:5446-5451. 

Myers, N. and A.H. Knott. 2001. The biotic crisis and the future of evolution. PNAS 98: 
5389-5392. 

Novacek, M. J. and E.E. Ctevano. 2001. The current biodiversity extinction event: Scenarios 
for mitigation and recovery. PNAS 98:5466-5470. 

SiMBERLOFF, D. 1988. The contribution of population and community biology to conserva- 
tion science. Ann. Rev. Ecol. Syst. 19:473-512. 

Wi.uiams, C.B, 1943. Area and number of species. Nature 152:264—267. 


STATUS OF QUERCUS x LEANA AND QUERCUS 
x RUNCINATA (FAGACEAE) IN ILLINOIS 


Gordon C.Tucker John E. Ebinger 
Department of Biological Sciences peas onsionegieal Sciences 
Eastern Illinois University rn ili iversity 
Charleston, IL 61920-3099, U.S.A. Giaieston IL 1L.61920- 3099, U. SA. 
eiu.edu cfee@eiu.edu 
ABSTRACT 


The oak a uereds x cane Nutt. (Q imomicaria x 9. velunnd) ane 2 x nungindia (A. isla 
Engelm 

Ilin Populati isti f many individuals of these two hybrid taxa are not ino in Ili- 
nois, oa ealy rarely have suspected backcross individuals been found. A Principal Components 
Analysis (PCA) involving the parent taxa and the supposed hybrids indicates that little introgres- 
sion occurs, each taxon appearing as a distinct cluster on the PCA graph. A study involving hybrid 
specimens encountered in the herbaria of the state indicates that Q. x leana is the most common 
taxon, being found in 31 counties, Q. x runcinata being known from only 11 counties. 


Q. 


RESUMEN 
bles hibridos O leana Nutt.(Q i bricania x Q. veld x runeiata, DO Eng 
} ) t Wh 
(Q. imbricariax Q. I I todo ILlinois. 
Se conocen en Illinois poblaci d hos individ 1 dos taxa hibridos 


i 1 [ 
+] ( i oe seme | | 


Un Analisis 


L £ t 
= | ] ] ge gg Ree | 


} 

de Componentes Principales (PCA p y I 
2 .e ] ] Tae ] Ae de] 

p g l | g PCA. Un estudio 

de | i hibrid los herbarios del estado indica que Q. x leana es el taxon 

31 condados, mientras que Q. x runcinata se conoce solo de 11 condados. 


irre 


me 
mas frecuente and 


INTRODUCTION 


Oak woodlands, whether savanna, open woodlands or mixed forests of oak and 
hickory or oak and pine are found throughout the northern hemisphere and 
southward into India and the higher elevations in tropical America (Stearns 
1991). In North America oaks dominate a variety of community types, from the 
extensive oak-scrub communities of California, to the oak savannas of the prai- 
rie peninsula, to the oak dominated forests of the eastern United States. Over- 
all, about 400 species are recognized in the western hemisphere, the United 
States claiming 90 species, with numerous hybrids being reported (Sargent 1933; 
Stearns 1991; Nixon 1997). 

Because oaks are wind-pollinated many hybridize readily. Extensive popu- 
lations of oak hybrids are uncommon, the individual hybrids being compara- 
tively rare as wild plants (Palmer 1948). In Illinois, both Quercus x leana Nutt. 


SIDA 19(4): 1073 — 1081. 2001 


1074 BRIT.ORG/SIDA 19(4) 


a hybrid involving Q. imbricaria Michx. and Q. velutina Lam.,and Q. x runcinata 
(A. DC.) Engelm., with the parents being Q. imbricaria and Q. rubra L., are occa- 
sionally found in the wild; Biagi and Jensen (1995) reported them to be the most 
frequently encountered oak hybrids in Indiana. Herbarium collections of these 
hybrids are usually from the edge of woods, along fencerows, in open pastures, 
and other relatively open, disturbed habitats. The present study was undertaken 
to determine the distribution of Q. x leana and Q. x runcinata in Illinois, their 
morphological variation, and their habitat preferences. 


METHODS AND MATERIALS 


The distribution of Quercus x leana and Q. x runcinata in Illinois was deter- 
mined by examining specimens from the following herbaria (A, DEK, EIU, F 
GH, KNOX, ILL, ILLS, ISM, MO, MOR, MWI, SIU), consulting literature sources, 
and contacting many of the state’s field botanists. Also, both authors made ef- 
forts to locate additional specimens on their numerous field trips through much 
of Illinois during the past two years. These specimens were studied to deter- 
mine the geographic ranges and morphological variation of the probable hy- 
brids. Where there was sufficiently detailed information, some sites where these 
hybrids had been recorded, were visited between 1998 and 2000 and searched 
for the presence of oak hybrids. In appendix 1 the Illinois counties where each 
hybrid taxon is known are listed with a representative specimen that we have 
annotated. For a few counties where Quercus x runcinata has been reported we 
have not seen a specimen, we list the specimen cited by Jones and Fuller (1955). 
Initially, the herbarium specimens were separated into probable hybrid 
taxa based on the morphological characteristics listed in Table 1. After removal 
of duplicate, incomplete, and immature specimens, 25 specimens of Quercus x 
leana and five specimens of Q. x runcinata were available for subsequent analysis. 
These specimens, as well as 22 randomly selected specimens of each of the prob- 
able parents (Q. imbricaria,Q. rubra,and Q. velutina), from throughout their known 
ranges in Illinois, were scored for the seven vegetative characteristics listed in 
Table 2. Only vegetative characters could be used as few of the hybrid specimens 
had fruits attached, and only specimens collected in late summer and with 
mature leaves were scored. All characters were measured for each specimen 
(usually three or more measurements) and the measurements from the prob- 
able parental species plotted to confirm that gaps existed in order to permit the 
use of scored characters. Of these characters, the number of bristles present 
was determined by picking the side of the blade with the most bristles, while 
the angle of the major vein to the midvein was determined using a major vein 
that extended into the blade lobe. The data were analyzed by principal compo- 
nents analysis (PCA) using NTSYS-pc version 1.70 (Rohlf 1993). All characters 
were standardized, the PCA was performed using a correlation matrix, and the 
data were transformed by a square root transformation (Sokal & Rohlf 1969). 


TUCKER AND EBINGER, TWO HYBRID OAK TAXA IN ILLINOIS 


1075 


Taste 1.A morphological comparison of Quercus x leana, Q. x runcinata, and their purported paren- 
tal taxa. 
Quercus Quercus Quercus Quercus Quercus 
Character velutina x leana imbricaria x runcinata rubra 
Terminal bud densely lightly glabrous& = glabrous & glabrous & 
pubescence pubescent pubescent ciliate ciliate ciliate 
Petiole length >30 mm 18-30mm <15mm 18-30mm >30mm 
Blade apex acuminate acute to obtuse to acute to acuminate 
acuminate acute acuminate 
Blade sinuses >25mm <25mm none <25mm >25mm 
deep deep deep deep 
Blade width/ 0.70-1.00+ 0.38-0.65 0.25-0.40 0.43-0.65 0.70-—1.00+ 
length ratio 
Bristles on one 15-26 2-7 0 2-7 15-26 
side of blade 
Major vein angle = 34-45° 34-45° 50-65° 35-45° 35-49° 


Taste 2. Characters scored for the principal component analysis ee specimens of Quercus 
imbricaria, Q. rubra, and Q. velutina and the probable hybrids of these tax 


1. Terminal bud scales Laseretas (1 = glabrous and ciliate, 2 = lightly pubescent and ciliate, 3 = 
densely pubescent and ciliate). 


2. Petiole length (mm). 
3. Blade apex shape (1 = obtuse to acute, 2 = acuminate). 
4. Blade lobing (1 = not lobed, 2 = shallowly lobed, the sinuses mostly less than 25 mm deep, 3 = 


deeply lobed, many sinuses more than 30 mm deep). 
5. Blade width at widest point/length ratio. 
Bristles present on one side of the blade (1 = a single terminal bristle, 2 = 2-6 bristles, 3 = 8 or 
more bristles 
7. Angle of major lobe vein to midvein (1 = >50°, 


Ov 


2 = <50°). 


RESULTS AND DISCUSSION 


Quercus x leana has an extensive geographic range throughout much of the 
eastern United States from Pennsylvania to Michigan, and south to Missouri, 
North Carolina, and Maryland (Palmer 1948). The type specimen was collected 
in Cincinnati, Ohio by T.G. Lea in 1842 (GH)). Jones and Fuller (1955) reported 
Quercus x leana for seven Illinois counties (Clay, Cook, Fulton, Peoria, Richland, 
Sangamon, and Will). The present study indicates that this taxon is relatively 
common, occurring in 31 Illinois counties (Fig. 1). Of the more than 100 her- 
barium specimens of this taxon examined, only three had sufficient label in- 
formation to allow relocation of the original tree. Also, hybrids were found at 


1076 BRIT.ORG/SIDA 19(4) 


Fig.1 Wm . k L . Lol At pecth at fa { (©) and Q. x runcinata (@). 


four new sites. At six of these sites, only one hybrid individual was found, while 
two trees were found at one site. All were in disturbed habitats (fence row, forest 
edge, open field) with the probable parents nearby. 

Quercus x runcinata also has an extensive geographic range, being reported 
from throughout most of the northeastern and north central United States in 
Kentucky, Illinois, Indiana, Maryland, Missouri and Pennsylvania (Palmer 1948). 
In Illinois this hybrid was reported for only six counties (Champaign, Johnson, 
Richland, St. Clair, Wayne, and Will) by Jones and Fuller (1955), while an up- 
date of this work added Peoria County (Winterringer & Evers 1960). During 
the present study specimens were recorded from four additional counties (Dou- 
glas, Moultrie, Saline, and Vermilion), bringing the total to 11 counties (Fig. 1). 
No living hybrids were found during the present study, but information on her- 


TUCKER AND EBINGER, TWO HYBRID OAK TAXA IN ILLINOIS 1077 


barium labels indicates that only individual trees were found, and that they 
were associated with disturbed habitats. 

In our PCA involving specimens of the three purported parental taxa (Q. 
imbricaria, Q. rubra, and Q. velutina), and all of the probable hybrid specimens 
scored, the first two principal components accounted for 72% and 15%, respec- 
tively, 87% of the total variance. In the resulting plot the 96 specimens separate 
into five distinct groups, three representing the parental taxa, the remaining 
two representing the two hybrid taxa (Fig. 2). The extent of blade lobing and 
blade width/length ratio (characters 4 and 5) are most important in determin- 
ing OTU scores on the first principal component. The extent of pubescence on 
the terminal bud and the number of bristles on the blade (characters | and 6) 
are most important in determining OTU scores on second principal component. 

The five clusters in the PCA plot are distinct, well-separated, and do not 
contain recognizable subgroups (Fig. 2). In this plot OTU’s representing Quercus 
x leana fall between those representing Quercus imbricaria and Q. velutina. The 
compact nature of this group suggests that the specimens are probably F1 hy- 
brids and probably do not represent backcrosses to either parent. Similar re- 
sults are obtained for Q. x runcinata, OTU’s representing that taxon are located 
between those representing Q. imbricaria and Q. rubra (Fig, 2). 

Morphological characteristics of the leaves were important in separating 
both Quercus x leana and Q. x runcinata from the parental taxa since nearly all 
of the specimens examined were sterile (Table 1). In both hybrid taxa the leaves 
are similar, with petioles between 18 and 30 mm long, blades shallowly 3- to 9- 
lobed with the sinuses mostly less than 25 mm deep, a width/length ratio of 
less than 70%, an acute to acuminate blade apex, and between 2 and 7 bristles 
on each side of the blade (Fig. 3). In Q. imbricaria the entire margined leaves are 
not lobed, the petioles are usually less than 15 mm long, and the obtuse to acute 
blade apex has a single terminal bristle. In contrast, the leaves of both Q. rubra 
and Q. velutina have petioles that commonly exceed 30 mm in length, deeply 
lobed blades with some sinuses more than 30 mm deep, a blade width/length 
ratio above 70%, an acuminate blade apex, and commonly 15 to 26 bristles on 
each side of the blade (Table 1). 

Though the leaves of Quercus rubra and Q. velutina are very similar, these 
two taxa are easily separated. Generally the clustered terminal buds of Q. 
velutina are 4-angled, 7-10 mm long, and with densely pubescent and ciliate 
scales; those of Q. rubra ovoid, less than 6 mm long, and the scales mostly gla- 
brous and lightly ciliate (Mohlenbrock 1986; Gleason & Cronquist 1991). Ter- 
minal bud characteristics are also useful in separating the two hybrid taxa. In 
both the terminal buds are fairly short, mostly less than 6.5 mm long. In Q. x 
leana the buds are commonly slightly 4-angled and the scales are lightly to 
densely pubescent and ciliate. Quercus x runcinata, in contrast, has ovoid buds 


1078 BRIT.ORG/SIDA 19(4) 


Q. velutina : 
kek o* a 


0.5- 
KK kkk kK 
* wk 
OQ. x leana x * 
wake KKK * * 
kK kkKK 
wAKKRKEK 
0.0- 
Q. imbricaria * 
KaKKEK * * ok 
Kk KEKE * 
RRR KK RR Q. xX runcinata 
* 
kek 
* 
* 
ak 
-0.5- * * 7 
k 
* 
* 
QO. rubra * 
kK *x 
* * 
* 
-1.0- - 
Tyr Sg: a a. ate gs et a. eet A ee Se e* Se a es ea cs ae gs 
£{Nn L 0 


Fig. 2. Plot of axis 1 vs. 2 of a principal component analysis using 7 variable for 22 sp 
rubra and Q ina al ith 30 speci f their bable hybrid 


with scales that are glabrous with lightly ciliate margins, similar to the termi- 
nal buds found in both Q. imbricaria and Q. rubra (Table 1). 

Few of the hybrid specimens bore mature acorns. Acorns of Quercus x leana 
were similar to, but slightly smaller than those of Q. velutina, the caps of both 
with loose, pubescent scales. Also, acorns of Q. x runcinata were similar to, but 
smaller than those of Q. rubra. Both taxa have glabrous to lightly puberulent, 
tightly appressed scales. 

The present study suggests that the oak hybrids Quercus x leana and Q. x 
runcinatd are uncommon, usually occurring as isolated individuals in disturbed 
habitats. Palmer (1948) mentions that these hybrids have been grown from seed, 
and that the mature foliage and fruits of these offsprings are intermediate be- 


TUCKER AND EBINGER, TWO HYBRID OAK TAXA IN ILLINOIS 1079 


tHan€ din Wings : £0 leana Tk left leaf Q. imbricaria, the one 


Fic. 3. Leaf vari 
at the lower left Q. velutina, tt inder st 


leana 


£ 4 ° * ££) 


1080 BRIT.ORG/SIDA 19(4) 


tween the purported parents. During the present study one tree was found that 
bore large numbers of acorns. Of the 250 acorns collected from beneath this 
tree, only 14 appeared to be of firm and of normal size, but none germinated. 
The remaining acorns were smaller, hada relatively soft pericarp, and lackeda 
fully developed embryo. The small number of viable fruit suggests that seed- 
lings rarely develop. 


APPENDIX 1 


Illinois counties where Quercus x leana and Q. x runcinata have been found 
with a representative collection from each county giving locality, date collected, 
collector, and herbarium where the specimen is deposited. 


Quercus x leana Nutt. Adams Co.: wooded bluff,W section Siloam Springs State Park, 29 Sep 1961, 
RA.Evers 71960 (ILLS). Champaign Co.: bluff top of Sangamon River, N of Mahomet, 18 Jun 1953, 
R.A.Evers 38112 (ILLS). Clark Co.: woodland border, SW of Moriah, 3 Sep 1970,R.A.Evers 103546 (ILLS). 
Clay Co.: woodland border,N of Louisville, 4 Jun 1953, R.A.Evers 37577 (ILLS). Coles Co.: EIU campus, 
Panther Trail, W of football stadium, Charleston, 9 May 2000, G.C. Tucker 12022 (EIU, ILLS, MOR). Cook 
Co.: woods, Columbia Woods Forest Preserve, Willow Springs, 17 Jul 1963, RAEvers 77205 (ILLS). 
Fayette Co.: edge of lowland old field, N side of Becks Creek, Dean Hills Nature Preserve, 8 mi E of 
sae, NE1/4 S10 T8N R2E, 29 Aug 2000, D.T. Busemeyer, pe Grr aila ale te 306 (ILLS). Fulton 

. Bemadolte, J Wolf s.n.(ILL). Hamilton Co.: roadside S of Dahlgren, 14 Sep 1951, RA.Evers 32566 
atin Hardin Co.: wooded bluff,4 mi S of Karbers Ridge, 8 Jul 1958, R.A.Evers pe (ILLS). Johnson 
Co.: open uplands, sandy soil, Simpson, 3 Oct 1919, £./.Palmer 16670 (MO). Livingston Co.: roadside, 
10 mi N of Rowe, 22 May 1945, G.D.Fuller 10357 (ISM).Macon Co.: Spitler Wood State Park, 14 Oct 
1965,R.A.Evers 86074 (ILLS). Madison Co.: woodland, Godfrey, 1 Oct 1953,H.E.Ahles 7599 (ILL). Massac 
Co.: floodplain eee River,6 mi N of Metropolis, near Rocky Branch, 18 Oct 1973, WMcClain 853 
(EIU).McD h top of dry ridge, Pleasant Valley, 18 May 1948, R.M.Myers 576 (ISM, MWI, SIU). 
Menard Co.: eee ee New Salem State Park, 23 May 1963, R.A. Evers 76017 (ILLS). Peoria Co.: 
wooded edge of parking lot, W of visitor center near power lines, Wildlife Prairie ae NW1/4 S29 
TON R7E, 2 Jul 2000, G.C. Tucker 12124 (EIU, ILLS). Perry Co.: edge of woods, 8 Jul 1 J.O.Neill s.n. 
(ILLS). Piatt Co.: woodland border, Lodge Park, S of Lodge, 3 Jun 1 971, RA. 105252 Ate Randolph 
Co.: bottomland woods, 1 mi S of Leanderville, 8 Jul 1953, R.A. Evers 38997 (ILLS). Richland Co.:: flat 
uplands, near Olney, 15 May 1923, £./ Palmer 22626 (MOR). Sangamon Co.: Washington Park, Spring- 
field, 15 Apr 1945, VS Fifert 9958 (ISM). Schuyler Co.: 6 Sep 1849, S.B.Mead s.n. (KNOX). St. Clair Co.: 
near Aldolf Engelmann’s farm, 4 Jun 1874, G.Engelmann s.n. (MO). Union Co.: roadside woods, 1 mi 
S of Jonesboro, 9 Jul 1956, F.B.Buser 6467 (ILLS). Vermilion Co.: edge of upland woods, Middle Fork 
Nature Preserve, W edge of Kickapoo State Park, NW1/4 S5 TION R12W, 16 Oct 2000, J.F.Ebinger 
29400 (EIU). Wabash Co.: edge of woods and road, Beall Woods Nature Preserve, 2 mi E of Keensburg, 
S11 12S R13W, 6 Jun 2000, J.E.Ebinger 28583 (EIU). Washington Co.: uncommon hybrid along rail- 
road tracks along IL 177 at Venedy, 26 Jun 1993, M.A. Basinger & ala 5878 (ILLS). Will Co.: 
edge of woods, Marley, 22 Jun 1987, A.Chase 26 (ILL, MO). Williamson Co.: pasture woodland N of 
Marion, 21 Sep 1950, R.A.Evers 27868 (ILLS). 


ed 


Quercus x runcinata (A. DC.) Engelm. Champaign Co.: woods, Crystal Lake Park, Urbana, 18 Oct 
1960, R.A.Evers 68121 (ILLS). Douglas Co.: edge of woods, Kaskaskia River near Chesterville, 13 Oct 
1970, AJones 2886 (ILL). Johnson Co.: upland woods, limestone outcrops, Simpson, 3 Oct 1919 
E.J.Palmer 16666 (MO). Moultrie Co.: tree 40-50 ft tall along edge of small woods by the Illinois 
Central RR, 1.5 mi NW of Allenville, NE1/4 SE1/4 $17 T13N RSE, 25 Jun 1973, L.Dennis 1547 (EIU). 
Peoria Co.: in Peoria, drive overlooking Illinois River, 27 Sep 1949, G.S.Winterringer 19434 (ISM). 


TUCKER AND EBINGER, TWO HYBRID OAK TAXA IN ILLINOIS 1081 


Richland Co.: Turkey Creek, R.Ridgway 26. Saline Co.: sandstone outcrop on Murray Bluff, S of Car- 
rier Mills, 30 Jul 1954,R.A.Evers 45369 (ILLS). St. Clair Co.: upper edge of bluff road, vic. Southern rr, 20 
Nov 1960, J.O.Neill 15529 (ISM, MO, SIU). Vermilion Co.: Kickapoo State Park, 24 Jul 1971, D. Seigler 
4726 (ILL). Wayne Co.: near Sannon Bridge, 21 Oct 1914, R.Ridgeway 97. Will Co.: property of Mrs. 
Koneeny, rt. 1, Mokena, Oct 1961,FA Swink s.n. (MOR). 


ACKNOWLEDGMENTS 


The authors would like to thank the herbarium curators for the use of the oak 
specimens used to complete the study. 


REFERENCES 


Biaci, A.and RJ. Jensen. 1995. The genus Quercus (Fagaceae)in Indiana: phytogeography 
and a key to the species. Proc. Indiana Acad. Sci. 104:11-24. 

Gteason, H.A. and A. Cronauist. 1991. Manual of the vascular flora of northeastern United 
States and adjacent Canada. 2nd ed. The New York Botanical Garden, Bronx. 

Jones, G.N. and G.D. Futter. 1955. Vascular plants of Illinois. The University of Illinois Press, 
Urbana. 

MOHLENBROCK, R.H. 1986. Guide to the vascular flora of Illinois, revised and enlarged edition, 
Southern Illinois University Press, Carbondale. 

Nixon, K.C. 1997. Quercus, In: Flora of North America Editorial Committee, eds. Flora of North 
America North of Mexico, Vol. 3. Magnoliophyta: Magnoliidae and Hamamelidae. Ox- 
ford Univ. Press, New York. Pp. 445-506. 

Patmer, E.J. 1948. Hybrid oaks of North America. J.Arnold Arbor. 29:1-48. 

Routr, FJ. 1993. Numerical taxonomy systems-pc (NTSYS-pc), Version 1.70, Applied Biosta- 
tistics, Inc, Setauket, NY. 

SARGENT, C.S. 1933. Manual of the trees of North America. Houghton Mifflin Co., Boston, 


Sokat, R.R. and F.J. RoHLF. 1969. Biometry. W.H. Freeman, San Francisco, CA. 

STEARNS, F. 1991.Oak woods: an overview. |n:G.V. Burger, J.E. Ebinger, and G.S.Withelm, eds. 
Proceedings of the oak woods management workshop. Eastern Illinois University, 
Charleston. Pp. 1—7. 

WiInTERRINGER, G.S. and R.A. Evers. 1960. New records for Illinois vascular plants. Illinois State 
Muis: SC. Pap. Ser i 1-135. 


1082 


BRIT.ORG/SIDA 19(4) 


Book NOTICE 


PETER FE FFoLuiott, Luts A. BojoRQUEZ-TapiA, and MARIANO HERNANDEZ-NARVAEZ. 2001 


Natural Resource Management Practices: A Primer. (ISBN 0-8138-2541-5 
hbk.). lowa State University Press, 2121 South State Street, Ames, [A 50014- 
8300, U.S.A. (Orders: www.isupress.com, 800-862-6657, 515-292-0155, 515- 
292-3348 fax). US$52.95, hbk., 250 pp., illustrated, 6" x 9" 


Table of Contents: 
I 


) Introduction 
2) Watershed and Water Management Practices 
3) Rangeland Management Practices 

4) Timber Management Practices 

5) Agroforestry Practices 

6) Wildlife Management Practices 

7) Fishery Management Practices 

8) Outdoor Recreation Practices 

9) Wilderness Management Practices 

10) Fire and Pest Ma oe Practices 

11) Soil Conservation Prac 

12) Rehabilitation of eee Land 

13) Integrated Natural Resources Management 


14) Importance of Research to Natural Resource Management 
lish to Metric Conversions, Appendix 2: Plot Studies; Appendix 3: Statistical Methods; 


Appendix 1: En 
a sis 4: Computer Simulation Models; Appendix 5: Geographic Information Syste 
s. As 


resource Management practice 


Index 


Ay 


and natural 


- 


This is a concise primer logical 
a quick reference for researchers or as a text for students in conservation classes, Natural Resource 
the subject. Covering aspects of watershed and 


Management Practices, is a much needed book o 
as the i of disturbed lands this book should g oe a a 
X76 


agroforestry practices, as W 
pe a in oe —Kevin D. Janni, Botanical Research Institute of Texas, Fort Worth, 1 


4060, ‘itorg. 


SIDA 19(4): 1082. 2001 


THE VASCULAR FLORA OF MADISON COUNTY, TEXAS 
Amanda K. Neill! and Hugh D.Wilson 


Department of Biology Herbarium (TAMU) 
Texas A&M University 
ollege Station, 1X 77843-3258, U.S.A. 


ABSTRACT 


A survey of the vascular flora of Madison County, Texas, was conducted from May 1996 to April 
2000. The flora consists of 985 species in 488 genera and 136 families. The largest families in the 
flora are the Asteraceae (127 species), Poaceae (123 spp.), Fabaceae (76 spp.), Cyperaceae (63 spp.), 
and PUppOrbiaeeae (32 spp.). The species are compiled in an annotated ie that includes for each 
name, collection citations, and a designation as to whether that species is con 


sidered native or introduced. Habitat ty where each species occurs are included in the list. Un- 
usual habitats, introduced species, and rare and endemic taxa are discussed. 


RESUMEN 


Una see de la note eames ge ‘Madison County, Texas, fue realizada desde mayo de 1996 
abril de 2000. La f1 488 géneros y 136 familias. Las 


familias mayotes son las Asteraceae (27 especies), Boacene (23 esp.), Bapacede (76 — J; ea 
(63 esp.), y 1 (32es 

para a epece el nombre sere ae de las regoleseiones yuna cai de sise peonsilcs a 
nativa o ntrocucica: Se mmeliyen en la lista los Epes de los habia en que vive cada especie. Estan 
comentado I taxones raros y endémicos. 


INTRODUCTION 


Madison County, like many other counties of Texas, has lacked an in-depth flo- 
ristic evaluation. A complete state-wide flora has long been a goal of Texas bota- 
nists. Lundell’s Flora of Texas (1942) was intended to bea ten-volume series, but 
the project was abandoned in 1955. Correll and Johnston’s Manual of the Vascu- 
lar Plants of Hage (1970) ncOnDeIatEG Lundell’s work, and remains the only 
reference with ing the entire state. Updated check- 
lists of the Texas flora have been compiled by Gould (1975a), Hatch et al.(1990), 
and Jones et al. (1997). Although these reference species by region or vegeta- 
tional area, no county-by-county species information is available due to the 
dearth of local floristic studies. Reed (1997) compiled the Manual of the Dicot 
Flora of Brazos and Surrounding Counties, technically including Madison 
County. However, less than 200 collections from this county were available for 
reference from Texas A&M University’s two herbaria at the time (FTC Her- 
barium Specimen Browser, internet reference). 

Almost a century ago, Texas botanist William Bray stated that “before the 


‘Present address: Institute of Systematic Botany, The New York Botanical Bronx, NY 10458-5126, U.S.A. 
SIDA 19(4): 1083 — 1121. 2001 


1084 BRIT.ORG/SIDA 19(4) 


flora of Texas suffers further radical changes, the schools of the state ought to 
cooperate in securing a complete and authentic list of species represented by 
carefully collected and well-preserved specimens” (Bray 1906). While the col- 
leges and universities of Texas have a great tradition of ecological and system- 
atic studies, a century has brought us but little closer to a modern state-wide 


flora. The radical floristic changes of which Bray spoke continue unabated and 
often unobserved, and the labor-intensive floras of Texas counties are still woe- 
fully few. 


The objectives of this county flora were to 1) collect, identify, and preserve 
specimens of all vascular plants occurring in Madison County, Texas; 2) gener- 
ate a species list for the county and delineate patterns of plant communities by 
habitat preference; and 3) analyze the flora for disjuncts, notable range exten- 
sions, endemics, recent invaders, and rare taxa. This article is provided asa sum- 
mary of the first author’s MS. thesis research (Neill 2000). Information is also 
available on the Madison County Flora Website at <http:// 
BIOCOURSE.BIO.TAMU.EDU/graduate-students/neilla/madhome.htm>. This 
site includes annotated species lists, searchable in several formats, as well as 
maps and links to images. 


Site Description 

Madison County comprises 302,451 acres (122,398 ha) in east-central Texas 
between 30°49' and 31'06'N latitude and 95°37' and 96°13' W longitude (Ramos 
1997). The county is bordered on the west by the Navasota River and Brazos 
County, on the east by the Trinity River and Houston County, on the north by 
Leon County, and on the south by Bedias Creek and Grimes and Walker coun- 
ties (Fig. 1). The county seat and largest city is Madisonville, situated about 90 
mi (142 km) NNE of Houston. 

The land surface is flat to gently rolling and generally slopes to the south- 
east, with elevations ranging from 420 ft (128 m) above sea level south of 
Normangee to 140 ft (43 m) above sea level along the lower Trinity River and 
Bedias Creek (a tributary of the Trinity) (U.S.G:S. 1962-1989). The eastern two- 
thirds of the county are drained by these watercourses, which converge at the 
southeastern corner. While the Trinity is a hydrologically tamed river with a 
historical floodplain 10 miles wide and now largely under cultivation, Bedias 
Creek is for most of its length a small, deep creek with extensive bottomlands 
of oxbow lakes, swamps, and frequently flooded ridge-and-swale topography. 
The Navasota River (a tributary of the Brazos River) drains the western third of 
Madison County, where it meanders through a wide and wooded floodplain of 
sloughs, natural lakes, terraces, and ravines. 

The geology of East Texas is the result of alluvium washed southward by 
rivers and deposited in layers upon the advance and retreat of coastal waters in 
the Gulf of Mexico. This deposition occurred in the region during the Eocene 


NEILL AND WILSON I TEXAS 1085 


N LEON COUNTY 


\ 
ROBERTSON 
COUNTY 


f{ WALKER COUNTY 


Legend 
ottomland soils 
tO é h 
0 er [| Post Oak Savanna! 
Madison County, TexaS 10, 3) (6 km == Blackland Prairie 


| Pineywoods 


Fic. ra " £1 I 1 1 r tate A-l 1 £, Cc la natal (10Z£Nn\ ne bk /(1004)\ 
5 


Epoch (58 to 37 mya) of the Tertiary Period (Spearing 1991). The Eocene layers 
deposited in Madison County are members of the Claiborne Group and are com- 
posed of clays shales, siltstones, sandstones, and lignites (Neitsch 1994). Neitsch 
(1994) lists 12 soil associations in the county, with six upland soils and three 
prairie soils, all loamy sands and clays. Three bottomland soils of alluvial clay 
cover modern floodplains and young terraces. 

The Madison County frost-free growing season averages 250 days, with a 
mean maximum July temperature of 96°F (35°C) and a mean minimum Janu- 
ary temperature of 38'F (3°C). Precipitation peaks in the spring and fall and av- 
erages 41.6 in (106 cm) annually (Ramos 1997). 


Cultural History 
Although no major archaeological sites have been found in Madison County, 
some Indian camp sites and kitchen-middens have been found that date from 
3000 B.C. (Madison County Historical Commission 1984). Tribes of the Caddo 
Confederacy formed permanent settlements in the eastern part of the county, 
where the people cultivated corn and squash. Nomadic tribes that occupied 
the county include the Bedias, who frequented Bedias Creek and its confluence 
with the Trinity River, and apparently had a village in this peninsular area 
(Neitsch 1994; Texas Almanac 1951). 

French and Spanish explorers, missionaries, and soldiers began leading 
expeditions through the area in the mid-sixteenth century. The first Spanish 


1086 BRIT.ORG/SIDA 19(4) 


settlement, Bucareli—where present-day Texas Highway 21 crosses the Trinity 
River—was established in 1774 as an outpost against French invasion. Bucareli 
was abandoned in 1779 due to flooding, epidemics, and raids by the nomadic 
Comanches (Madison County Historical Commission 1984). 

Madison County is most historically significant for containing the inter- 
section of two early roads. Originally Indian trails, these roads were used by 
missionaries, explorers, and eventually settlers. The Old San Antonio Road 
(OSR), also known as El Camino Real, or the King’s Highway, connected settle- 
ments in the East with those in San Antonio and Mexico. Today, this modern 
highway forms most of the northern border of Madison County. At the town of 
Midway, another road known as La Bahia Trail split from OSR and led south- 
west to the mission at Goliad (U.S. Dept. of the Interior 1998). 

Madison County was organized in 1853, and settlement increased rapidly 
toa population of 3000 by 1860 (Texas Almanac 1951). By 1900, the population 
had grown to 10,432, and 71,000 acres were being farmed (Richardson 1940). 
Over 20,000 acres of this was in cotton. Three railroads were built through the 
county, and the population peaked in 1930 at 12,227 inhabitants (Ramos 1997. 
Richardson 1940). Pasture and hayland for beef cattle production have replaced 
most cultivated crops today. Seventy percent of the county is used as rangeland, 
hayland, and pasture (Neitsch 1994). Fishing and hunting are important land 
uses in the county, with many properties leased for seasonal duck and deer hunting. 


Botanical History 

Every vegetation map available represents Madison County as an ecotone. Past 
ecologists and botanists in Texas recognized that the area around the 96" me- 
ridian, which divides the county in half, isa zone where western xerophytic 
grasslands mingle with eastern mesophytic forests (Bray 1906; Blair 1950; 
Cronquist 1982; Tharp 1926). Allred and Mitchell (1955), Correll and Johnston 
(1970), Cory and Parks (1937), and Gould et al. (1960) generally agree upon the 
boundaries of the three major vegetation zones converging in Madison County; 
these vegetation zones commonly are called the Pineywoods, Blackland Prai- 
ries, and Post Oak Savannah (Fig. 1). 

Correll and Johnston (1970) described the vegetational areas largely in ac- 
cordance with past descriptions. The Pineywoods region of East Texas repre- 
sents the most mesophytic area of Texas and is characterized by extensive pine 
and pine-hardwood forests with intermittent swamps. Madison County’s east- 
ern edge is one of the most westward-extending tongues of this vast vegeta- 
tional zone. The Blackland Prairies region lies in three diagonal bands trending 
NE-SW across east-central Texas. Of the two southern bands of Blackland Prai- 
rie, the San Antonio Prairie occupies the border of northwestern Madison 
County, and the Fayette Prairie is sometimes shown to extend into the south- 
central portion of the county. The Post Oak Savannah region interdigitates with 


NEILL AND wil SON, MAVIIVUIN LUUINTE I, TEXAS 1087 


the Blackland Prairies. A belt of Post Oak Savannah occupies most of central 
Madison County. 

Although no extensive vegetational studies have been conducted in Madison 
County, past research has included floristic aspects found within the county. 
Both Launchbaugh (1952) and Hightower (1987) investigated changes in veg- 
etation structure in response to disturbance in the San Antonio Prairie. Diamond 
(1980) described remnant plant communities of the Fayette Prairie, which is 
the southern disjunct of the Blackland Prairie proper, placing the northern limits 
of the Fayette Prairie in Grimes and Walker counties. McCaleb (1954) analyzed 
the vegetation of the southern Post Oak Savannah and included Madison County 
within his Oak-Hickory Forest Region. Allen’s (1974) vegetational study of the 
lower Navasota described plant communities along the watershed in Grimes 
County and points south. Apparently, none of the above investigators conducted 
research within Madison County. There are no resultant voucher specimens for 
the county in the Tracy (TAES) or Biology Department (TAMU) herbaria at Texas 
A&M University (FTC Herbarium Specimen Browser, internet reference). 


METHODS 


Vascular plants were collected at frequent and regular intervals in Madison 
County, from May 1996 to April 2000. Public lands (roadsides, railroad rights- 
of-way, city parks, and cemeteries) were investigated, and habitats identified as 
unusual were revisited over several seasons. With the assistance of a local news- 
paper article (Levey 1998), private landowners were contacted for permission 
to collect on their properties. Other private lands were chosen with the aid of 
aerial photographs, overflight in a small chartered plane, and topographic and 
soils maps, in order to survey both agricultural/livestock land-use areas and to 
investigate possible unusual or undisturbed habitats. Urban areas (Madisonville, 
Normangee, North Zulch, and Midway) provided disturbed upland sites sup- 
porting a unique, weedy flora. Riparian and lacustrine systems were accessed 
at bridges, on private property, and by canoe. 

At the time of plant collection, data were recorded on the specific habitat 
and relative abundance of the species in the area. Publications describing similar 
research were reviewed for exposure to floristic protocol. References include 
Allen (1974), McCaleb (1954), Reed (1997), Starbuck (1984), and Wilson (1972). 
References used for plant identification were Correll & Johnston (1970), Diggs 
et al. (1999), and Reed (1997). Grasses were identified with The Grasses of Texas 
(Gould 1975b), and identifications were verified by Stephan Hatch and Dale 
Kruse at the S.M. Tracy Herbarium in College Station, Texas (TAES). Sedges (Carex 
sp.) were identified with an unpublished key to the Carex of Texas provided by 
Stanley Jones, curator of the Botanical Research Center, College Station, Texas 
(BRCH), and identifications were verified by him. Other experts consulted were 
Monique Reed (TAMU), Theodore Barkley at the Botanical Research Institute 


1088 BRIT.ORG/SIDA 19(4) 


of Texas (BRIT), and Richard Rabeler at the University of Michigan Herbarium 
(MICH). Taxonomic classification essentially followed the Cronquistian system 
(Cronquist 1988), with the exception of familial treatment of the Tupelo family 
(Nyssaceae), Chickenspike family (Sphenocleaceae), and Dodder family 
(Cuscutaceae). Voucher specimens collected throughout the project were de- 
posited in the herbaria at Texas AGM University in College Station, Texas 
(TAMU, TAES), and are available for examination. 

A search of the Flora of Texas Consortium Browser (2000a) revealed some 
species that the authors did not find in the county, and these are included in the 
checklist. Only those specimens held at TAMU and TAES were seen by the au- 
thors. Numerous older specimens had been labeled “Madison County” but ac- 
tually were collected northwest of Normangee in Leon County. The location 
and identification of species collections held at ASTC, LL, TEX, SHST, and SMU 
have not been verified. 


RESULTS AND DISCUSSION 


Madison County vascular plant collections resulted in a total of 2254 speci- 
mens. The flora is comprised of 1041 taxa with 985 species in 488 genera and 
136 families. The five largest plant families are Asteraceae (127 species), Poaceae 
(123 spp.), Fabaceae (sensu stricto) (76 spp.), Cyperaceae (63 spp.), and 
Euphorbiaceae (32 spp.). These five families contain approximately 44% of the 
species found in Madison County. Further numerical distributions of the flora 
are found in Table 1. 


Major Vegetational Habitats 

Thirteen major habitat types classifiable in Madison County. These habitats 
are ially those described for Robertson County, Texas, by Starbuck (1984). 
corn habitats and lowlands include habitats in water, near water, and low 
areas within floodplains. Aquatic emergent (AEM), aquatic f loating (AFL ). and 
aquatic submerged (ASU) are habitat categories for obligat 

those plants that were found in standing water due to flooding, Habitats along 
the banks and frequently (two or more times a year) flooded areas of riverine 
and lacustrine bodies of water were divided into major riparian (RLM) and 
minor riparian (RUM) habitats. Major riparian habitat is that of the Navasota 
and Trinity rivers, whereas minor riparian habitats are those along creeks and 
ponds. Lowland habitats flooded once a year or less frequently were divided by 
woody cover into terrestrial lowland woods (TLW), terrestrial lowland open 
(TLO), and terrestrial lowland mixed (TLM). Drier upland areas were similarly 
divided into terrestrial upland woods (TUW), terrestrial upland open (TUO), 
and terrestrial upland mixed (TUM). Two of Starbuck’s habitat groups, parasite 
(PAR) and epiphyte (EPD, better describe plant habits, but were maintained for 
consistency in this text. Two of Starbuck’s habitats were not used. His bog (BOG) 


NEILL AND WIL SON VASCULAR FLORA NIAMIOUIN LUUITT ET, TEXAS 1089 


Taste 1. Distributional surmmary of the Madison County flora. 


Taxa Families Genera Species 
Ferns & Allies 11 13 14 
Gymnosperms 3 4 5 
Dicots 101 357 697 
Monocots 2] 114 269 
Total 136 488 985 


habitat applies to the acidic Sphagnum bogs found in the deep sands of 
Robertson County (Bryant 1977; Starbuck 1984). This habitat is not present in 
Madison County. Starbuck’s terrestrial prairie (TPR) habitat was used in his text 
solely for specimens collected by Hightower (1987) at a remnant prairie locality. 
Although certain plant species in Madison County could be characterized as 
prairie species, no undisturbed prairie habitats are intact in the county today. 

Aquatic habitats such as oxbow lakes, stock ponds, reservoirs, drainage 
ditches, and shallow creek and river margins supported a unique flora. Uncom- 
mon herbs found in these habitats include Juncus repens, Nelumbo lutea, Thalia 
dealbata, and Utricularia radiata. Lowland and riparian habitats contain the 
most diverse flora in Madison County. The extensive wooded floodplains of the 
Navasota River and Bedias Creek exemplify typical bottomland hardwood veg- 
etation. Trees most common in low, moist areas of the county include Carya 
aquatica, Fraxinus pennsylvanica, Nyssa sylvatica, Planera aquatica, Quercus 
lyrata, Q. nigra, Q. phellos, Tiliaamericana, and Ulmus crassifolia. Disturbances 
to this area include logging, flooding, and cattle grazing. Due to agricultural 
use and the construction of levees and dams, the Trinity River floodplain for- 
ests are less extensive. Some segregation of species was found when comparing 
collections from the two major riparian habitats (RLM). Species found only 
within the Navasota River floodplain include Amsonia illustris, Aristolochia 
tomentosa, Baptisia alba, Brunnichia ovata, Carex complanata, C. crus-corvi, 
Mikania scandens, Planera aquatica, Prunus caroliniana, Tillandsia recurvata, 
Urtica chamaedryoides, and Zizaniopsis miliacea. Species found only within 
the Trinity River floodplain include Amorpha fruticosa, Ampelopsis cordata, Carex 
retroflexa, Clematis reticulata, Equisetum laevigatum, Hibiscus moscheutos, 
Panicum virgatum, and Wood wardia areolata. Terrestrial lowland open habitats 
(TLO) were typically found in deforested areas or river floodplains, low pas- 
tures, moist roadside ditches, and other open areas near bodies of water. These 
habitats are often disturbed by grazing and mowing and thus support a variety 
of weedy invader species as well as native perennial herbaceous species. 

Level upland sites in Madison County are prime locations for pastures and 
haymeadows, as they were mostly cleared in the last century for cropland. Most 
wooded areas within what would be considered uplands are nearly always found 


— 


1090 BRIT.ORG/SIDA 19(4) 


along creeks, and so would be included in the terrestrial lowland classifications 
for this study. True wooded upland sites (TUW) and mixed upland habitats (TUM) 
do occur, and these may be categorized into three general association types: 

1. Post Oak — Winged Elm - Yaupon Association 

This association encompasses nearly all upland wooded sites in the county, and 
is typical of the Post Oak Savannah vegetation type. The association is found in 
areas with sandy loam topsoil and gray clay subsoils and is characterized by 
Quercus stellata, Ulmus alata, and Ilex vomitoria. 


2. Blackjack Oak — Post Oak — Winged Elm Association 

Characterized by the presence of Quercus marilandica, sites exhibiting this as- 
sociation are generally more poorly drained than those with the Post Oak - 
Winged Elm - Yaupon association. The aspect of these sites is also that of the 
Post Oak Savannah vegetation type but was frequently found with tall grasses 
seen in the Blackland Prairie vegetation type, such as Andropogon ternarius, 
Sorghastrum nutans, and Schizachyrium scoparium. 


3. Southern Red Oak — Sweet Gum — Post Oak Association 

This association is occurs in the eastern third of the county and is character- 
ized by a dominance of Quercus falcata and Liquidambar styraciflua. These 
sites have acidic sandy soils with good drainage. The best examples of this as- 
sociation are found in the uplands along Youngs Creek Canyon and Cobb Creek 
Canyon. These localities are discussed in the Unique hae pagaes The 
Padina series of deep sands is frequently found at these sites. Sas albid 
and Pinus taeda are also found in this association 


eb i 


Terrestrial upland open habitat (TUO) is ubiquitous and easily accessed in Madi- 
son County. Like open lowland sites, these generally roadside and pasture habi- 
tats are characterized by non-native herbs and grasses, although it is here that 
many of the showy and popular wildflowers of Texas also may be seen. Open 
upland habitats are first to suffer in summer drought conditions, when even 
narrow county roadsides are mown for hay in this county. It is difficult to find 
examples of this habitat type that have not experienced disturbance. Pasture 
sites are frequently invaded by Prosopis glandulosa and Acacia farnesiana. 

Epiphytes (EPI) found in the county are restricted to Pleopletis 
polypodioides, Tillandsia recurvata, and T. usneoides. Parasites (PAR) found in 
the county include Cuscuta cuspidata, C. glomerata, C. obtusifolia, Orobanche 
multiflora, and Phoradendron tomentosum. 


Unique Habitats 

Several localities in Madison County, primarily on private lands, exhibit un- 
usual characteristics that deserve special mention. Isolated populations of Pinus 
taeda are found at several places in the county, but the only area that deserves 


NEILL AND WILSON | TEXAS 1091 


designation as the East Texas Pineywoods vegetation type is found in south- 
eastern Madison County, on the peninsula formed by the confluence of the Trin- 
ity River and Bedias Creek and extending up Bedias Creek for several miles, 
although this area also contains some lowland hardwood forest. Species col- 
lected only in this habitat include Aristolochia serpentaria, Carex hyalina, 
Dioscorea villosa, Lygodium japonicum, and Spiranthes praecox. 

No intact, undisturbed pieces of the San Antonio Blackland Prairie are pre- 
served in Madison County. The narrow band of prairie that once followed OSR 
from the Navasota River to a few miles east of Interstate Highway 45 has been 
lost to grazing, crops, and encroachment of woody species. Although the road- 
sides are occasionally mown, some portions of this stretch of road contain pock- 
ets of prairie herbs along the fencelines and treelines. These sites support spe- 
cies including Astragalus distortus, Buglossoides arvensis, Eriogonum 
longifolium, Eryngium yuccifolium, Lythrum alatum, Orbexilum simplex, Pen- 
stemon australis, Rudbeckia grandiflora, R. maxima, and Silphium radula. 
Roadsides along OSR are the best places in the county for wildflower viewing 
in the spring and summer. 

The Navasota River will continue to host some of the most unique habitats 
in Madison County if the wild stretch of it south of Lake Limestone is allowed 
to remain unimpounded. The upper terraces along the river consist of deep 
sands where a high water table p prings and nearly bog-like seeps. Here, 
two springs supported a number of species not found elsewhere in the county: 
Baccharis halimifolia, Galium obtusum, Hydrocotyle ranunculoides, Juncus 
debilis, Polypogon monspeliensis, Ranunculus sceleratus, Saccharum baldwinii, 
Sagittaria platyphylla,and Taxodium distichum. The flows of these springs have 
been altered by landowners to provide water for livestock, but the habitat is 
still rich and unique. One site also had a spring-seep area surrounded by Myrica 
cerifera and Rubus argutus and harbored a few species typically associated with 
true bogs, such as Pluchea foetida, Rhexia mariana, and Fuirena squarrosa. 
Unfortunately, in 1999 the site was excavated to create a small tank, and the 
altered hydrology may not continue to support these unusual plants. 

Numerous natural lakes and oxbows are found along the Navasota River. 
Oxbows in low woods near the river host several aquatic and terrestrial species 
rarely or never seen elsewhere in the county. Some of these are Didiplis diandra, 
Eragrostis hypnoides, Juncus repens, Oldenlandia boscii, Saururus cernuus, and 
Utricularia radiata. Just south of OSR on the Navasota River lies a string of 
natural lakes in the open floodplain. These lakes have a unique flora differing 
from that of the oxbow lakes. Species found in and around these lakes include 
Bacopa rotundifolia, Eleocharis palustris, Justicia americana, Nelumbo lutea, 
Nuphar lutea, Sphenoclea zeylanica, and Zizianiopsis miliacea. Most of these 
species were found nowhere else in the county. Plants in these bodies of water 


1092 BRIT.ORG/SIDA 19(4) 


are somewhat protected from curious botanists by snapping turtles and nest- 
ing alligators. 

The Navasota River has created ancient high river terraces, which appear 
as a series of hills that are easily seen from OSR. These hills have bald tops with 
a white sand summit ringed by scrubby Post Oak woods, and rocky sides with 
rich prairie-like pockets. These unusual hills host species with distinctly xeric, 
western affinities. Species restricted to the sands include Argemone albiflora, 
Asclepias viridiflora, Croptilon rigidifolium, and Mirabilis linearis. Rivina 
humilis was found under the oaks near the summit of one of these hills. Species 
only seen on the rocky hillsides include Loeflingia squarrosa and Opuntia 
humifusa. The latter was covered with the scale insects (Homoptera: Coccoideae) 
from which the red pigment cochineal is extracted. The grassy areas of the hill- 
side were the only sites for Dalea compacta in the county. 

In the northern portion of the county are two creeks that drain into the 
Trinity River, Youngs Creek and Cobb Creek. These creeks have cut down into 
the ancient terrace of sand, clay, and mudstone to form deep canyons, creating 
a landscape unlike that in any other part of Madison County. However, the two 
creeks have very different histories, and this is reflected in their floras. Youngs 
Creek Canyon formed within the past 70 years as the result of an excavation by 
the owners in an attempt to divert the creek and reduce flooding of the sur- 
rounding cropland (Leon Wakefield, pers. comm. 1997). The easily eroded soil 
was quickly washed down to the ironstone bedrock, creating sculpted walls 
more than seven meters high, bizarre monolithic pinnacles of sandstone, and 
waterfalls. The surrounding land exhibits the uncommon Southern Red Oak - 
Sweet Gum - Post Oak association, but the canyon itself has not yet developed a 
unique flora. Only two species, Hydrocotyle umbellata and Najas guadalupensis, 
were found solely at Youngs Creek Canyon. Future monitoring of species re- 
cruitment at this easily accessible canyon could provide an interesting study 
of colonization in a situation of unusual disturbance. 

The uplands around Cobb Creek are similar to those around Youngs Creek, 
but the maze of spring-fed gullies and canyons was created without human 
interference. This locality supports a flora that is _ ied with age of ao 
ern Texasand has produced several species g lon 
along springs and low banks of Cobb Creek are Apios a mericana, Aralia spinosa, 
Carex emoryi, Onoclea sensibilis, Quercus shumardii, Triadenum walteri, and 
Woodwardia areolata. One very unusual habitat at this canyon wasa site witha 
hanging garden on the canyon wall that was continued in some aspects on the 

cliff above. The woods on the cliff were mostly composed of Quercus 
marilandica. Species found nowhere in the county other than this site include 
Areolaria grandiflora, Echinacea angustifolia, Lactuca hirsuta, Liatris elegans, 
Osmunda cinnamomea, and Spiranthes tuberosa. Other species unique to 


— 


NEILL AND WILSON , TEXAS 1093 


wooded upper banks of Cobb Creek include Dichanthelium ravenelti, 
Erythonium albidum, Hieracium gronovii, and Packera obovata. Species never 
or only rarely encountered elsewhere and occurring on high sandy banks and 
open areas include Alophia drummondii, Cuphea carthagenensis, Cuphea 
glutinosa, Drosera brevifolia, Euphorbia tetrapora, Hypericum gymnanthum, 
Juglans nigra, Lepuropetalon spathulatum, and Rotala ramosior. 

One area of the county is of particular and current interest because of its 
impending destruction. As reported in Hipp (2000), a 27,400-acre (11,089 ha) 
reservoir is planned on upper Bedias Creek at the junction of Madison, Grimes, 
and Walker counties. The impoundment will obliterate some of the most pris- 
tine and mature low woodlands along Bedias Creek. Species found only in the 
area scheduled to be inundated are Carex corrugata, Orbexilum pedunculatum, 
Poncirus trifoliata, and Liatris cymosa, an endemic discussed in the following 
section. The proposed reservoir will possibly flood the only area in the county 
where Spiranthes parksii, an endangered orchid, has been collected. This spe- 
cies is also discussed in the following section. Opponents of the Bedias reser- 
voir have expressed concern for the loss of wildlife habitat and possible archaeo- 
logical sites. We would add that the proposed reservoir threatens some of the 
greatest plant diversity and grandest natural beauty in Madison County. 


Range Extensions, Invaders, and Rare Taxa 
This project found 40 species, in 11 genera and 5 families, not previously col- 
lected in the counties surrounding Brazos County and Texas ASM University, 
which was the area covered by Reed (1997). The most notable range extension 
is that of Vicia lutea, or Yellow Vetch. Madison County collections of V. lutea 
constitute the first record of its presence in Texas, as discussed in Neill (1999). 
Already known to occur in North Carolina, Louisiana, and California, this an- 
nual Mediterranean weed now is to be expected in other Texas counties. Three 
dramatically invasive species with few local records were found in Madison 
County: Alternanthera philoxeroides, Lygodium japonicum, and Petrorhagia 
dubia. These exotics will undoubtedly become familiar weeds in the area. 
One hundred fifty non-Texas-native taxa were found in the county, or over 
14% of the total taxa of Madison County. The majority of these non-natives are 
roadside and pasture weeds, favored by disturbance and occasionally introduced 
for forage. A few non-native crop species were found as waifs or persisting small 
populations. Examples of these crops include Asparagus officinalis, Brassica 
rapa, Solanum esculentum var. cerasiforme,and Triticum aestivum. Non-native 
species introduced for horticultural purposes were found in varying stages of 
persistence at old abandoned home sites and at cemeteries throughout the 
county. Examples of species restricted to these sites and not appearing to be 
adventive are Canna x generalis, Ficus carica, Hedera helix, Hemerocallis fulva, 
Hibiscus syriacus, Hippeastrum bifidum, Lagerst roemia indica, Lantana camara, 


1094 BRIT.ORG/SIDA 19(4) 


Ligustrum lucidum, Liriope spicata, Oxalis rubra, Stenotaphrum secundatum, 
and Wisteria sinensis. Horticultural species appearing mildly adventive in that 
they occurred in areas where they were not likely to have been eae or they 
were witnessed to be reproducing in localized areas, include Aila 

Gladiolus italicus, Iris pallida, Mirabilis jalapa, Nandina domestica, Narcissus 
jonquilla, Pyruscalleryana, Rosa multiflora,and Solanum capsicastrum. A num- 
ber of horticultural escapes are well known invaders and are established and 
naturalized throughout the county, including Albizia julibrissin, Ligustrum 
sinense, Lonicera japonica, Melia azedarach, and Sapium sebiferum. 

Texas endemics found within Madison County are Cucurbita pepo var. 
texana, Krigia cespitosa f. gracilis, Lechea san-sabeana, Liatris cymosa, Lupinus 
subcarnosus, L. texensis, Palafoxia rosea var. rosea, Spiranthes parksii, and 
Valerianella florifera (Flora of Texas Consortium 2000b). 

Three species of special concern were collected and are discussed below. 
The source for federal information is Texas Parks & Wildlife (1991). 


Carex hyalina Boott.—This sedge was collected only at one site, in sandy up- 
land pine woods near the Trinity River-Bedias Creek confluence. Carex hyalina 
was considered rare and imperiled in 1990 and was a federal candidate under 
review for possible listing as Category 2 and either threatened or endangered in 
Texas. This species seems to be more common than previously thought and 
currently has no federal status. 


Liatris cymosa (H. Ness) K. Schum.—This endemic species is restricted to ap- 
proximately six counties in southeast Texas. Although its restricted range places 
it in danger of extinction, this species is not under federal review for listing. 
This species was collected only at one site, on eroded clay at the edge of low 
woods near Bedias Creek in south-central Madison County. 


Spiranthes parksii Correll —This endemic species was federally listed as rare 
and endangered in 1982, and is restricted to approximately eight counties in 
southeast Texas. This species has been collected twice in the south-central part 

of the county. Other likely habitats in the county were searched, but it was never 
encountered by the authors. 


ANNOTATED CHECKLIST OF THE SPECIES OF 
MADISON COUNTY, TEXAS 
Vascular plant families are alphabetically arranged within their respective di- 
visions. Angiosperm [families are subdivided into the Magnoliopsida (Dicots) 
and Liliopsida (Monocots). Taxa are listed alphabetically within families by 
genus, species, and subspecific epithets, and authorities are given for all spe- 
cies. Latin names are followed by common names. Parenthetical annotation in- 
cludes a designation of nativity to Texas, indicated by N (native) or | (intro- 


NEILL AND WILSON, TEXAS 1095 


duced), followed by abbreviations of the major vegetational habitats in which 
the taxon was collected. Alphabetically, these are: aquatic emergent (AEM), 
aquatic floating (AFL), aquatic submerged (ASU), epiphyte (EPD), parasite (PAR), 
major riparian (RLM), minor riparian (RUM); terrestrial lowland mixed (TLM), 
open (TLO), and woods (TLW); and terrestrial upland mixed (TUM), open (TUO), 
and woods (TUW). Voucher specimen numbers collected by Neill (N) are held 
at TAMU and TAES herbaria at Texas ASM University. Other collectors’ num- 
bers are given with the collector's surname and herbarium abbreviation: Stephen 
F Austin University (ASTC), Botanical Research Institute of Texas (BRIT and 
SMU), University of Texas (LL and TEX), and Sam Houston State University 
(SHST).Common names and nativity data are from Correll and Johnston (1970), 
Diggs et al. 1999), and Hatch et al. (1990). 


LYCOPODIOPHYTA sacs lunaroides (L.) Sw., Grapefern (N) Wil- 

ISOETACEAE Sl ceca yeee 
; oe oglossum a aires es Heartleaf 

lsoetes melanopoda J. Gay & Durieu ex Durieu, 
Adder's Tongue (h 68 (TAMU) 
Quillwort (N; AEM) N 2545 

OSMUNDACEAE 

EQUISETOPHYTA Osmunda cinnamomea L., Cinnamon Fern (N; 
TLM) N 2668 


EQUISETACEAE 
Equisetum laevigatum A.Br,Smooth Horsetail (N; |» POLYPODIACEAE 
LM) N19 oS podioides (L.) Andrews &Windham 
ichauxiana (Weath.) Andrews & 
ice Resurrection Fern (N; EPI) N 558 
SCHIZACEAE 
Lygodium japonicum (Thunb. , 7 Japanese 
Climbing Fern (1; TUW) N 


POLYPODIOPHYTA 


ASPLENIACEAE 
Asplenium platyneuron (L.) B.S.P., Ebony Spleen- 
ort (N; TLW, TUM) N 595, 1203, 1246 


AZOLLACEAE PINOPHYTA 
Azolla caroliniana Willd., Water Fern (N; AFL) N 
130, 981, 1997 CUPRESSACEAE 
SLECHNACER Juniperus virginiana L., Eastern Red Cedar (N; 
GEAE . TUM, TUO) N 83, 341, 2013, 2456 
Woodwardia areolata (L.) T. Moore, Chain Fern (N; 


Platycladus orientalis (L.) Franco., Arborvitae (|; 


RUM) N 2081, 2666 TLM) N 630 
DRYOPTERIDACEAE 
Onoclea sensibilis L., Sensiti 

N 581, 2667 
Woodsia obtusa (Spreng.) Torr., Common 

Woodsia (N; RUM, TLW, TUW) N 56, 596, 1587, 


PINACEAE 

Pinus echinata Mill, Shortleaf Pine (N; TUM) N 1776 

Pinus taeda L.,Loblolly Pine (N;TLW, TUW) N 1225, 
1453, 2054, 2520 


Fern (N;RLM,RUM 


=a 


660 TAXODIACEAE 
Taxodium distichum (L.) Rich., Bald Cypress (N; 
MARSILEACEAE RUM) N217 
Marsilea macropoda A.Br.,Water-clover (N;TLW) 
N 1934 


OPHIOGLOSSACEAE 
Botrychium biternatum sone Underw., 
Cutleaved Grapefern (N; TLW) N 2390 


MAGNOLIOPHYTA: MAGNOLIOPSIDA 


ACANTHACEAE 
Dicliptera brachiata (Pursh.) Spreng. (N;RLM) N825 


1096 


Hygrophila lacustris (Schlecht. & Cham.) Nees, 
Lake Acanthus (N; RLM) N 834 

Justicia americana (L.) Vahl, American Water-wil- 

; N 1888 

Justicia ovata (Walter) Lindau var. lanceolata 
(Chapm.) R.W.Long, Lanceleaved Water-wil- 
low (N; RLM, TLM, TLW) N 59, 284, 1655, 2196 

Ruellia humilis Nutt., Low Ruellia (N;TUO) N 203, 

24,982, 1684, 1799, 1806 
Ruellia nudiflora (Engelm. & A. Gray) Urban, Vio- 
uellia (N; TLW, TUO) N 985, 1998 
Ruellia strepens L., Limestone Ruellia (N; RLM, 
N 582, 1 

ACERACEAE 

Acer negundo L., Boxelder (N; RUM, TLW, TUM) N 
141, 1010, 1582, 27184 

AMARANTHACEAE 

Alternanthera philoxeroides (Mart.) Griseb,, Alliga- 

d (I; AEM) N 1915 
Amaranthus albus L., Tumbleweed Amaranth (N; 


<= 


N 269 
Amaranthus palmeri S. Wats., Palmer Amaranth 
(N; TUM, TUO) N 2242, 2262 
Amaranthus polygonoides L., Tropical Amaranth 
(N; TUCO) N 244 
ee rudis Gres Weedy dale (N; 
RLM, RUM, TLO) N 815, 963, 2182, 2 
Peers Spinosus ‘ Spiny pes (I; TUO) 
2308 


Amaranthus viridis L., Green Amaranth (I; TUO) N 
2241, 2269 


neseliciie drummondii Mo Drummond 
Snakecotton (N) Taabenhaus 2643 (TAES) 

Froelichia floridana (Nutt.) Moq,, var. tae 

Florida Snakecotton (N;TUO) N 

Froelichia floridana (Nutt.) Mog. var. ee: 
(Small.) Fern., Florida Snakecotton (N; TUO) 
N 727, 2216 

Gossypianthus lanuginosus (Poir.) Mog. in A. DC. 
var. lanuginosus, Woolly Cottonflower (N; 
TUO) N 1072 

lresine rhizomatosa Standl., Rootstock Bloodleaf 
(N; RLM, RUM) N 868, 1073 


ANACARDIACEAE 
Rhus aromatica Aiton var. serotina (Greene) 
ehd., Fragrant Sumac (N; TUM, TUO, TUW) 
N 529, 2098, 2561 
Rhus copallina L., Flameleaf Sumac (N;TLM, TUM, 
TUW) N 733, 1134, 1724 


BRIT.ORG/SIDA 19(4) 


Rhus glabra L., Smooth Sumac (N;TLM) N 147 
Toxicodendron radicans (L.) Kuntze. ssp.radicans, 
n Ivy (N;TLM, TUW) N 157, 1452, 1457 
APIACEAE 
Bowlesia incana Ruiz & Pav., Hoary Bowlesia (I: 
RUM) N 2503 


es tainturier! Hook. var. dasycarpum 
Hook. ex S. Wats., Hairyfruit Chervil (N; TLM 


— 
= 


N 1239, 157 

Baas tainturieri es var. tainturieri, 
Chervil (N; TLO, TLW) N 527 

Cicuta maculata L.var es Water- 
hemlock (N; RUM, TLM, TUM) N 75, 648, 1810, 


1953 

Cyclospermum leptophyllum (Pers.) Sprague, 
Slimlobe pa (I; RUM, TLM, TLW) N 1597, 

ace digitatum DC,, Finger Dogshade 
(N; RUM, TLO, TLW) N 52, 1669 1741, 1925 

Daucus pusillus Michx., Rattlesnake Weed (I;TLO, 
TUO) N 602, 1690, 1835, 2625 

Eryngium hookeri Walp., Hooker Eryngo (N;RLM, 
TLO, TUO) N 127, 583, 1750, 1882, 1956, 1979 

Eryngium prostratum Nutt. ex DC., Creeping 
Eryngo (N; dae TUW) N 2517, 1912, 2614 

Michx. var. yuccifolium, But- 

ton Snakeroot (N; TUM, TUO) N 

Hydrocotyle ranunculoides Ff, Lie Water- 
pennywort (N; AEM, RUM) N 1346, 1989 

Hydrocotyle umbellata L., Umbrella Water-pen- 
nywort (N; RUM) N 603 

Hydrocotyle verticillata Thunb. var. triradiata (A. 
Rich.) Fern. ee ar pennywort (N; 
RUM, TLO) N 1746, 

Hydrocotyle verti cillata an var. en 

Whorled Water-pennywort (N; RLM) NV 


1988 
Limnosciadium pinnatum (DC.) Math. & Const., 
ae aoe (N; RLM, TLW) N 71217, 


Press jum pumilum (Engelm. & A. Gray) 

be th. & Const. Prairie Dogshade (N; RUM, 
Ww 4, 1663 
rye nuttallii DC., Prairie Parsley (N; TLM, 
, UO) N 63,621, 764 

roto texana (J.M. Coult & Rose) Math. & 
Con xas Prairie Parsley (N; RUM) ae 

Ptilimnium di (DC.) Britton, Nuttall M 
Bishops Weed (N; TUM, TUO) N 23, 566, Be 
1797, 2650 


NEILL AND WILSON 


, TEXAS 


Sanicula canadensis L., Canada Snakeroot (N; 
TLW, TUW) N 599, 1412, 1557 

pasa divaricata (Walter) Raf.ex Ser.,Forked 
Scales (N;TUM, TUO) N 1789, 2624 

Sprae inermis (Nutt.ex DC.) Math.& Const, 
Spreading Scaleseed (N; TUO) N 2599 

Torilis ae (Huds.) Link., Hedgeparsley (I; TLM, 
TUM) N 753, 1587 

Trepocarpus aethusae Nutt.ex DC. (N; RLM, TLM) 


APOCYNACEAE 

Amsonia illustris Woods., Blue a (N; RLM, TLO, 
TLW) N 1347, 1657, 1995, 232 

Apocynum cannabinum L., ae (N; TUM, 
TUQ) N 1801, 2281 

Trachelospermum difforme (Walter) A. Gray, 
Climbing Dogbane (N; RLM, TLO, TLW, TUM, 
TUW) N 233, 610, 1675, 1745, 1919, 2323, 2612 

AQUIFOLIACEAE 

flex decidua Walter, Possum-haw (N; RLM, TUM, 
TUW) N 422, 665, 943, 1051, 1110, 1208, 1447 

llex longipes Chapm. ex Trel. var. hirsuta Lundell, 
Georgia Holly (N) Dixon 432 (SMU) (Lundell 

43) 
Ilex opaca Aiton, American Holly (N; RLM, TUW) 


wo 


Ilex vomitoria Aiton, Yaupon Holly (N; RLM, TUM, 
TUW) N 215, 942, 1050 

ARA 

ola a Devil's Walking Stick (N;RUM) N 


re ies L., English Ivy (1; TLM) N 


ARISTOLOCHIACEAE 

Aristolochia serpentaria L. var. hastata (Nutt.) 
Duchartre, Virginia Dutchman's Pipe (N;TLW) 
N 2382 


1244 


lochi Sims,Woolly Pipe-vine (N; 
RLM) N 2341 


ASCLEPIADACEAE 

Asclepias ee is i a leaf Milkweed 
(N; TLO, TUW) N 1662, 

Ase! pies obo Ell. kee (N; TUM) N 2058 

ham. & Schltdl. Prim- 

rose e Milkweed (N;TLO, TUM, TUO) N 238, 569, 

187 


ee tuberosa L., Butterfly Milkweed (N;TLM, 
UO) N 70, 1802, 1836 


1097 


Asclepias verticillata L., Whorled Milkweed (N; 
T UO) N 91,718, 1795, 1813 
Asclepias indie Raf.,Green Ant 
UM) 


| it NI 
p N; 


e 8 


70 

Asclepias viridis Walter, Antelope Horn (N; TUO) 

N 207, 279, 702, 1527 

Cynanchum laeve (Michx.) Pers., Smooth Swal- 
low-wort (N; RLM) N 900 

Matelea cyanchoides (Englem.) Woods, Sand 
Milkvine (N; TLW, TUM, TUQ) N 1682, 1727, 
1775 


Matelea gonocarpos (Walter) Shinners, Milkvine 


(N; TLW, TUM) N 214, 531, 1498, 1861 
ASTERACEA 
Achillea mill ium L. var. occidentalis DC., Yar- 
row (I; TUO) N87 


er artemisiifolia aud Ragweed (N; 
TUM, TUO) N 2086, 


Ambrosia psilostachya ae — stern Ragweed (N 
TLO, TUM, TUQ) N 266, 928, 1020, 2370, 2701 
Ambrosia trifida L., Giant Ragweed (N; RLM, TLO, 
UO) N 34, 820, 961, 2286 
Amphiachyris ene es DC., Annual 
oe (N; TUO) N 326, 911, 2180, 2713 
Arnoglossum plantagineum Raf., Indian Plantain 
area N 96, 1579 
i pert Nutt., Louisiana Sagewort 
(N; TUM, TUO) N 1118, 2153, 2696 
Aster ieeeie al var, texanus (Burgess) 
es, Texas Aster (N; TUM) N 1104 
stereos L., Heath Aster (N;TLW,TUO) N 986, 
1066, 
pid aa ale, Aster (N;TLM) N 1076 
ster lanceolatus Willd., Aster (N; TLO) N 
i lateriflorus (L.) Britton, California pene (N; 
RLM, TUM) N 949, 112 


Aster oolentangiensis Ridd., Azure Aster (N; TUM, 
TUO) N 1096, 2413 


Aster eee Aiton var. el Hook., Skydrop 
er (N; TUO) N 

aa Aiton var. an ae Aster (N; 
TUM, TUQ) N 329, 1137, 2368, 2 

Aster Bee Aiton var. eee, Lindl. ex 
DC) T.& G., Skydrop Aster (N; TUM) N 1793 

Aster ee Poir., Tall Aster (N; RLM) N 885 

Aster sericeus Vent. var. microphyllus DC., Silky 
Aster (N; TUM, TUO) N 1078, 1651, 2369, 2426 

Aster subulatus Michx. var. ligulatus Shinners, 
Lawn Aster (N;TLO, TUO) N 947, 1069 


—= 


a 


1098 


Astranthium integrifolium (Michx.) Nutt, Western 
Daisy (N; TUO) N 2605 
Astranthium integrifolium (Michx.) Nutt. ssp. 
ciliatum (Raf.) DeJong, Western Daisy (N;TLM) 
N 2578 
Baccharis halimifolia L., Eastern Baccharis (N; 
RUM) N 2329 
Baccharis neglecta Britton, Roosevelt-weed (N; 
UO) N 328, 1001, 1084 
Berlandiera pumila (Michx.) Nutt. Soft Greeneyes 
(N; TUM, TUQ) N 80, 2154 
Bidens bipinnata L.,Spanish Needles (N;TLM, TLW, 
M) N 774, 2391, 2424 
Bigelowia nuttallii L.C. Anders, Slender Bigelowia 
N; TLM) N 2044 


<< 


Boltonia asteroides (L.) L’'Her., White Boltonia (N; 
TUM) N 1776 
Boltonia diffusa Ell., Small Boltonia (N; TUM, TUO) 
2087, 
Calyptocarpus vialis Less., Horseherb (N; TUO) N 
2239 
Centaurea americana at American 
Basketflower (N; TLO, TUO) N 
Chaetopappa asteroides (Nutt.) a Comm 
Least Daisy (N; TLO, TUO) N 1222, 1706, en 
Chrysopsis pilosa ba al ea (N; TLO, 
TUM) N 104, 178, 1 1996, 225 
Cirsium altissimum (L ( an ae Thistle (N) 
Cory 54578 (LL) 
Cirsium engelmannii Rydb., Engelman Thistle (N; 
TUO, TUW) N 698, 1952 
Cirsium horridulum Michx., Bull Thistle (N; TLM, 
551, 149] 
Cirsium texanum Buckl., Southern Thistle (N; 
M) N 1866 
Conyza canadensis (L.) Cronq. var. canadensis, 
Horsetail (N; TUO, TUW) N 1047, 2710 
Conyza canadensis (L.) Cronq. var. glabrata (A. 
oe Cronq., Horsetail (N; TLM, TUO) N 2295, 
2693 


agen basalis (Dietr.) Blake, Goldenmane Tick- 
eed (N; TLO, TUM, TUO) N 33, 1845, 2616 
ie grandiflora Hogg. e 
longipes (Hook,) T.& G,, oe eee 
(N; TLO, TUM, TUO) N 463, 1466, 1798, 2568 
Croptilon divaricatum (Nutt.) Raf, Slender Scratch 
Daisy (N; TUM, TUO) N 277, 775, 2251, 2328 
Croptilon rigidifolium (E.B. Smith) E.B. Smith, 
Scratch Daisy (N; TUM, TUO) N 1873, 2392 


BRIT.ORG/SIDA 19(4) 


Dracopsis amplexicaulis (Vahl) reed ak 
Coneflower (N; TLW, TUO) N 
Echinacea ene: ia DC. var in Black 
n (N; TLM) N 2671 
Echinacea sanguinea Nutt., Purple Coneflower 
(N; TLM, TUM) N 1710, 1784 
Eclipta prostrata (L.) L., Yerba de Tago (N; RLM, 
RUM, TLM) N 839, 980, 2198, 2683 
Elephantopus carolinianus Raeusch, Leafy 
oo eae RUM, TLW) N 294, 774, 


Ear peste (Raf.) Goodman & CA. 
elmann’s Daisy (N; TLO, TUO) N 

ue 1826, 2567 

Erigeron strigosus Muhl. ex Willd. var. beyrichii 
(Fisch. & Mey.) T.&G. ex A. oi Prairie Flea- 
bane (N;TLO, TUO) N 565, 2 

Erigeron strigosus Muhl. ex ra var. Strigosus, 
Prairie Fleabane (N; TLM, TLO, TUM, TUW) N 
1807, 1899, 2046, 2598, 2613 

Erigeron tenuis T.& G., Slender Fleabane (N;TLM, 
TLO, TLW, TUO) N 39, 1302, 1515, 1622, 2636 

Eupatorium capillifolium (Lam.) Small, Dog-fen- 
nel (N; TUM, TUO) N 773, 1113, 2387 

Fupatorium coelestinum L., oo (N; RLM, 
au TLW) N 738, ene hee 


“(N: TUM, TUO) N 984, 2185, 2299, 2431 
Eupatorium incarnatum Walter, Pink Boneset (N; 
TLM, TLW) N 1026, 2388, 2417 
Eupatorium semiserratum DC., Boneset (N; RUM, 


Eupatorium serotinum Michx., Late Boneset (N; 
RUM, TUM, TUO) N 225, 293, 739, 2119 


Eurybia hemispherica (Alex.) Nesom, Low Aster 
(N; TLM, TUM, TUO) N 2412, 2420, 2695 

Euthamia leptocephala (T.&G.) Greene (N; TLM, 
TUO) N 2 430 


Evax candida (1.&G.) A. Gray, Silver Rabbit-to- 
acco (N; TUM, TUO) N 547, 1852, 2564 
Evax prolifera DC., Bighead Rabbit-tobacco (N; 
TLW) N 1263 


Evax verna Raf, Manystem Rabbit-tobacco (N; 
TUO) N 1520 

Facelis retusa (Lam.) Schultz-Bip. (|; TLW, TUO) N 
1511, 2 9/7 


, 


g.,Indian Blanket (N; TUO) 


Gaillardia pulchella F 
N 208, 2393 


NEILL AND WILSON TEXAS 


Gnaphalium falcatiuim Lam., Cudweed (I; TLW) N 


1248, 1927 

Gnaphalium obtusifolium L., Fragrant Cudweed 
(N; TLM, TLW, TUO) N 2294, 2423, 2697 

Gnaphalium pensylvanicum Willd., Pennsylvania 
Cudweed (N; TLW, TUO) N 387, 1252, 1574 

Gnaphalium purpureum L., Purple Cudweed (N; 
TUO, TUW) N 655, 1965 

Helenium amarum (Raf.) Rock var. a 
Sneezeweed (N; RLM, TUW) N 166, 6 

Helenium oo So an 

weed - at 2,261 

Heleniu ee um 06, Smallhead 

Sneezeweed (N; TUO) N 
el § Ls eae (N; TLO, TUO) N 


73,2204 

Helianthus debilis Nutt.ssp.cucumerifolius (T.&G.) 
Heiser, Cucumberleaf Sunflower (N; TUO) N 
1957, 2113 

Helianthus grosseserratus Martens, Sawtooth 

Sunflower (N; TLM, TLO) N 1G, 1045 

ianthus ae Raf., Hairy Sunflower (N;RUM, 

TUM) N 2011, 2045 

Heterotheca mariana (L.) Ell., Maryland 
Camphorweed (N; TLO) N 1825 (Chrysopsis 
mariana (L,) Ell) 

Heterotheca subaxillaris (Lam.) Britton & Rusby 
var. latifolia (Buckl.) Gandhi & Thomas, 
Camphorweed (N; TLO, TLW, TUM) N 939, 
1132,2395 

Heterotheca subaxillaris (Lam.) Britton & Rusb 
var. subaxillaris, Camphorweed (N; TLO) N 638 

Hieracium gronovii L., Hawkweed (N; RUM) N 


He. 


g 


Hymenopappus scabiosaeus L'Her. var. 

artemisiifolius (DC.) Gandhi & Thomas, Rag- 
Woolly-white (N; TLO, TUO) N 1514, 

1608, 2593 

Hymenopappus scabiosaeus L'Her. var. 
scabiosaeus (T.&G.) B.L. Turner, Flattop 
Woolly-white (N; TUM) N 505, 535 

lva angustifolia Nutt. ex DC., Narrowleaf 
Marshelder — TUM, TUO) N 983, 1083, 
2230, 2254, 233 

lvaannua L., paneer (N:-RLM,TLM, TLO) N 845, 


64, 2206 
Krigia cespitosa (Raf.) Chambers f. cespitosa, 
eedy Dwarf Dandelion (N; TLM, TLW) N 
1295, 2531, 2583 


1099 


Krigia ee (Raf.) Chambers f. gracilis (DC.) 
Kim, Dwarf ae (N; TLM, TUO) N 514, 
1540, oe 

Krigia dandelion re Nutt., Tuber Dwarf Dande- 
_tion (N; TUM es N 536, 1320, 2558 

I id Nutt., Western Dwarf Dande- 
me (N; TUO) N 2494 

Krigia virginica aie Virginia Dwarf Dande- 
lion (N; TUQ) N 

Lactuca etecale : ve Lettuce = TUM, 
TUO) N 1783, 2027, 2090, 2109 

Lactuca ae (L.) Gaertn., a ia (N; 

W) N 1664, 1970 

Lactuca hirsuta Muhl. var. albiflora (T.&G.) 
S Hairy Lettuce (N; TLM) N 2672 

Lactuca ludoviciana (Nutt.) Ridd., Wild Lettuce (N; 

UO) N 2651 

Lactuca serriola L., Prickly Lettuce (1; RLM) N 852 

Liatris cymosa (H. Ness) K. Schum., Branched 
Gayfeather (N; TUM) N 2298 

Liatris elegans ae ae 
Gayfeather (N; TLM) N 

Liatris mucronata DC.,, ie (N; TLM, TUM, 

UO) N 2231, 2314, 2429 

Liatris squarrosa (L.) Michx., Gayfeather (N; TUO) 
N 719, 1087, 2692 

Liatris squarrosa (L.) Michx. var. alabamensis 
(Alex.) Gaiser, Gayfeather (N; TLM, TUO) N 
2014, 2051 

Mikania scandens (L.) Willd., Climbing Hemp (N; 
RLM, RUM) N 2217, 847, 937 

peor nitidum (T.&G.) ot Flat-top Gold- 

rod (N; TUO) N 1049, 238 
eee obovata (Muhl. ex ae ) eee & Cove, 
olden Groundsel (N; TLW) N 2 

Packera tampicana (DC.) Jeffrey, eae (N; 
RLM, TLW, TUO) N 72, 425, 1272, 2521 

Palafoxia rosea (Bush) Cory var. rosea, Rose 
Palafoxia (N; TUM, TUO) N 709, 1796, 2089, 
2229, 2414 

Palafoxia texana DC. var.ambigua (Shinners) B.C. 

rner & Morris, Texas Palafoxia (N; TUO) N 


= 


— 


Pinkscale 


315 
Palafoxia texana DC. var. texana, Texas Palafoxia 
(N; TUM) N 
fuchea camphorata (L.) DC., Camphor Marsh- 
fleabane (N;RLM, TLM) N 867, 954, 2205, 2332 
Pluchea foetida (L.) DC., Stinking Marsh-fleabane 
(N; RUM) N 239 


me) 


1100 


Pluchea odorata (L.) Cass., Purple Marsh-fleabane 
(N; RLM) N 297, 2331 

Pterocaulon oe .) OC.,Wand Blackroot (N; 
TUO) N 23 


~~ cae anus (Walter) DC., Carolina 
e Dandelion (N; TUO) N 1556, 1904, 2634, 


Pyrrhopappus pauciflorus (D. Do 
Manystem False Dandelion (N; a N 2582 
Ratibida columnifera (Nutt.) Woot. & Standl., 

exican-hat (N; TUO) N 2654 
Rudbeckia grandiflora (D. Don.) J.F.Gmel ex DC. 
var. alismifolia ee oo Rough Cone- 
er (N; TUM) N 2 

ere grandiflora ie eel J.F.Gmel. ex DC. 
grandiflora, Rough Coneflower (N;TUM ) 


eee hirta L.var. hirta, Black-eyed Susan (N; 


TUO) N27 
Rudbeckia hirta L. var. angustifolia (T.V. Moore) 
Perdue, Black-eyed Susan (N; TUM) N 503 
Rudbeckia hirta L. var. pulcherrima Black- 
eyed Susan (N; TUM, TUOQ) N 1756, 1859 
wee maxima Nutt., Great nin eee (N; 
, TUO) N 177, 564, 2010 
oa. ampullaceus Hook., Texas Groundsel (N: 
LM, TLW, TUO) N 48, 1265, 1484, 2453 
ilo N CaraveeSimn HR atv 


TLO, TUO) N 646, 1612, 2646 
ee ey ,,} NI R Wer 


aianoas N 1809, ae 
4 


a 


Rydb,, 
mon Galdenied (N: TUOQ) N 1003 
ie canadensis L.var.scabra T.&G.Common 
Goldenrod (N; TUM) N 1730 
a gigantea Aiton var. serotina a 
Giant Goldenrod (N; TLM) N 2448 
a. nemoral is Aiton var.nemoralis, ae 
Goldenrod (N;TUO) N 989, 1901 
Solidago radula Nutt., Rough Goldenrod (N;TUO) 
N 1030, 2316 
Solidago ee P. Mill, Wrinkled Goldenrod (N; 
TUO) N 
Solidago ens EIl., Willowleaf Goldenrod (N; 
TUM, TUO) N 784, 1080, 2694 (S. patula Muhl. 
ex Willd. var. strictula Torr. & A. Gray) 
Solidago speciosa Nutt. var. Glee T.&G., 
Noble Soaat (N; TUO) N 
Solidago ulmifolia Muhl. ex Willd, a ese: 
Sid (N;TLM, TUO) N 2297 2415 


BRIT.ORG/SIDA 19(4) 


i few ee KA Lol 


A 


illd. var. microphylla 
A. coh ee (N; TUM, TUO) N 
268, 784, 2233 

Soliva sessilis Ruiz & Pavon, Burweed (1; TLW, TUO) 
N 449 1575, 1740 

Soliva stolonifera (Brot.) Loud., Burweed (I; TLW) 
N 1262 


Sonchus asper (L.) Hill, Prickly Sowthistle (I; TUO) 
N 1613, 2496 

Taraxacum officinale G.H. Weber ex Wiggers, 
Dandelion (|; TUO) N 

Tetraneuris scaposa (DC.) Greene, Bitterweed (N) 
Gonzales 15 (TAES) 

Verbesina encelioides (Cav.) Benth. & Hook f.ex A. 
Gray, Cowpen Daisy (N; TUO) N 2367, 2389 

Verbesina virginica L., Frostweed (N; RLM, RUM, 
TUW) N 1048, 2181, 2227, 2321 

Vernonia baldwinii Torr., Baldwin lronweed (N; 
RLM, RUM) N 292, 862, 2156 

Vernonia missurica Raf., Missouri Ironweed (N) 
Jo nes & Jones 407 (TAES) 

Vernonia texana (A. Gray) Small, Texas Ironweed 
(N; RUM, TLW, TUM, TUW) N 1973, 2088, 2228, 
2246, 2428, 2656 

Xanthium strumarium L., Cocklebur (N; RLM, 

N 163, 831, 2285 


BERBERIDACEAE 
Nandina domestica Thunb., Heavenly Bamboo (I; 
TUO) N 1623 


a 


BETULACEAE 

Betula nigra L., River Birch (N; RUM) N 1591, 1730 

BIGNONIACEAE 

Campsis radicans (L.) Seem.ex Bureau, Trumpet- 
creeper (N; RLM, TLM, TUM) N 89,671, 945 

Catal dis ee joides ese cad Cigar-tree 
(N; 


, TUQ) N 1800, 
edie 
igh des arvensis (L.) LM. Johnst., Bugloss (I; 
“TUO) N 363 


Heliotropium indicum L., Ga (N: RLM, TLO, 
TLW) N 126, 865, 1893, 1 

Heliotropium procumbens a Fourspike Helio- 

ope (1; RLM, TLM, TLO) N 883, 2188, 2659 

sabeceue incisum Lehm., Narrowleaf Puc- 
coon ( ioe 1199, ] 7 

rene macrosperma ngelm. — var. 
macrosperma, Spring rorgetmeno (N; 
RUM, TLW) N 4517, 1299, 1 


NEILL AND WILSON, , TEXAS 


Myosotis verna Nutt., Early Scorpion-grass (N; 
TLW) N 1300 

Onosmodium bejarense DC. ex A.DC. var. 
occidentale (Mack.) B.L. Turner, Western 
Marbleseed (N; TUW) N 1416 

BRASSICACEAE 

Brassica rapa L., Turnip (I; TUO) N 1321 

Capsella bursa-pastoris (L.) Medic, Shephard’s 
Purse (1; TLM, TUO) N 347, 1228 

Cardamine hirsuta L., Hairy Bittercress (N;TLM) N 


Cardamine parvifl icola (Britton) OLE. 
see cl Bittercress (N; TLM, TUM) N 384, 


Oab0 on Nutt. ex T.&G., Shortpod 
raba (N;TLM) N 2469 

a densiflorum 

pcppennies) (N; pea 


Schrad., 
1503 


Prairie 


Lepidium virginicum L. L. 
Hitchc.,Virginia eae (N: TLM) N 1230 

Lepidium virginicum L. var. virginicum, Virginia 
Pepperweed (N;TLM) N 57 

Neobeckia aquatica (Eat.) Greene, Water Horse- 
radish (N; AEM) N 1402 

Rapistrum rugosum (L.) All. (|; TUM, TUQ) N 1323, 


Rorippa sessiliflora (Nutt.) A.S. Hitchc., Stalkless 
Yellowcress (N; RLM, TLW) N 444, 1348, 1827, 


BUDDLEJACEAE 

aes procumbens L., Juniperleaf (N; RLM, 
M, TUM, TUO) N 185, 884, 2037, 2145 

CACTACEAE 


Opuntia engelmannii Salm-Dyck. var. linguiformis 
(Griffiths) Parfitt & Pinkava, Cowtongue 
Prickly-pear (N;TLO) N 

Opuntia humifusa (Raf.) Raf. var. humifusa, East- 
ern Prickly-pear (N; TUM, TUC) N 1779, 1855 

Opuntia macrorhiza Engelm. var. macrorhiza, 
Plains Prickly-pear (N; TUO) N 1698 

CAESALPINIACEAE 

Cercis canadensis L. var. canadensis, Eastern Red- 
bud (N;TLM, TLW, TUM) N 154, 368, 1529, 2522 

Chamaechrista fasiculata (Michx.) Greene, Prai- 
rie Senna (N;TLM, TUO) N 44, 908, 1029, 1853, 
2003 


= 


Gleditsia aquatica Marsh.,Water Locust (N; RLM) 
N2711 


1101 


Gleditsia ae L., Honey Locust (N; RLM, 
TLM, TLW, TUM, TUW) N 194, 584, 1739, 1282, 


Parkinsonia aculeata L., Retama (|; TUO) N 2632 
Senna ie ifolia (L.) Irwin & Barneby, Sicklepod 
a (I; TLM, TUO) N 2293, 2405 
a ae (L.) Link, Coffee Senna (|; TUQ) 
N 241, 2406 


CALLITRICHACEAE 

Callitriche heterophylla Pursh. (emend. Darby), 
Larger Water-starwort (N; AFL, RUM, TLW) N 

437, 1257, 1356, 1552 

Callitriche nuttallii Torr. Nuttall Water-starwort 
RUM, TLW) N 446, 1267 

Callitriche palustris L., Common Water-starwort 
(N: RLM, RUM, TLW) N 1553, 1937, 2539 

Callitriche peploides Nutt., Mat Water-starwort (N; 
RLM, TLO) N 2476, 2540 

CAMPANULACEAE 

Lobelia appendiculata A. DC., Earflower Lobelia 
(N; TUO, TUW) N 658, 1620, 1652, 1722 

Lobelia cardinalis L., Cardinal Flower (N; RLM) N 
859 


a 


N; 


=> 


Lobelia puberula Michx. var. pauciflora Bush, 
Downy Lobelia (N; TLM) N 2296 

Lobelia puberula Michx. var. puberula, Downy 
Lobelia (N; TUM) N 778 

Triodanis perfoliata (L.) Nieuw. var. biflora (R. & P.) 

adley, Small Venus’ Looking-glass (N;TLM, 

TUO) N 54, 518, 1487 

Triodanis perfoliata (L.) Nieuw. var. perfoliata, 
Clasping Venus’ Looking-glass (N; TLO, TUO) 
N 1426, 1536, 2566 

CAPPARACEAE 

Polanisia erosa (Nutt.) IItis ssp. erosa, Large 
Clammyweed (N;TLO) N 2155 


CAPRIFOLIACEAE 

Lonicera japonica Thunb., Japanese Honeysuckle 
=e. N 36,325, 1097, 

1126, 1133, 1584 
Lonicera sempervirens L., Scarlet isle (N; 
, TUM) N 135,357, 2025 

Sambucus nigra L. ssp. Ae 4 .) R. Bolli, El- 
derberry (N; RLM, TLW, TUM) N 78, 598, 858, 
1596, 1647 


Cc L : i / NA 


ip L h, Coralberry 
(N;TLM, TUM, TUW) N 43, 1722, 1138 


1102 


Viburnum rufidulium Raf,, Rusty Blackhaw Vibur- 
num (N;TLM, TLW, TUW) N 7149, 1578, 2072 


CARYOPHYLLACEAE 
Arenaria drummondii Shinners, Drummond 
rt (N; TUM, TUO) N 546, 1316, 1629 

( tia drummondii (Shinners) McNeill 

a Michx., Sandwort (N;TLW) N 1273 
(Minuartia patula (Michx.) Mattf.) 

Arenaria patula Michx. var. robusta Steyerm., 
Sandwort (N; TLM) N 2577 (Minuartia 
muscorum (Fassett) Rabeler) 

Arenaria serpyllifolia L., Thyme-leaved Sandwort 
(I; TLO, TUO) N 3617, 1195, 1223 

Cerastium bractypealum Pers, Shortpetal Chick- 
weed (1; T 

Cerastium acho (Engelm. & A. Gray 
Robins, Shortstalk Chickweed (N;TUO) N 365 

Cerastium eed Thuill., Mouse-ear (1; TUO 


= 


tL 


a 


7 


Loeflingia squarrosa Nutt., Spreading Loeflingia 
N 1856 
Paronychia lindheimeri Engelm ex A. Gray, 
Lindheimer Nailwort (N;TLM) N 2421 
Petrorhagia dubia (Raf.) G. Lopez & Romo, 
_ lang au (Il; TLO, TUM) N 1463, 1477 
(Ell.) T.&G., i Pearlwort 
RUM, TLM) N 2470, 273 
Silene ele L., Sleepy ier (N;TLO,TUO) 
N 257 
Ilene gala : Forked Catchfly (I; TUO) N 573 
1329, 


Stellaria red. (L.) Vill, Chickweed (I; TUM, TUW) 
N 335, 2 


CHENOPODIACEAE 

are see ides L., Epazote (I; RLM, 
RUM, TLM, TUO) N 869, 1709, 2157, 2199 

cheropodu cei Mog,., Pitseed Goose- 
foot (N; TUM) NV 

Chenopodium ae ane Aellen Standley 
Goosefoot (N; TUM) N 2371 

CISTACEAE 

Helianthemum georgianum Chapm. , Georgia 
Sunrose (N; TUM, TUO) N 1872, 1958, 2621 

Helianthemum rosmarinifolium Pursh., Rosemary 
Sunrose (N; TLO, TUO) N 1697, 1707, 2644 

Lechea mucronata Raf., Hairy Pinweed (N; TUM, 
TUO) N 1817, 700, 1843, 1959 

Lechea san-sabeana (Buckl.) Hodg., San Saba Pin- 
weed (N;TUO) N 706, 2620 


BRIT.ORG/SIDA 19(4) 


Lechea tenuifolia Michx., Narrowleaf Pinweed (N: 
TUM, TUO) N 705, 1686, 1879, 2069 


CLUSIACEAE 

Hypericum drummondii (Grev. & Hook.) T.&G., 
Drummond St. John's Wort (N; TLM, TUM, 
TUO) N 259, 1079, 1107, 2225, 2292 

Hypericum gymnanthum Engelm. & A. Gray, 
Clasping St. John’s Wort (N;TLO) N 2688 

Hypericum hypericoides (L.) Crantz. ssp. 
Lee icoides, St. Andrew's Cross (N; RUM, TLO, 

N 276, 747, 938, 2224 

Sau, mutilum L., Dwarf St. John’s Wort (N; 

TLM) N 2386 


Triadenum walteri (Gmel.) Gl., Marsh St. John’s 
rt (N; RUM) N 2079 


CONVOLVULACEAE 

Dichondra carolinensis Michx., Ponyfoot (N;TLW, 
TUO) N 389, 1555 

Evolvulus sericeus Sw., Silky Evolvulus (N; TUM, 
TUO) N616, 757, 1500 

Ipomoea cordatotriloba 
cordatotriloba, Bindweed (N;TLO, HS ees 
607, 955, 1824 

Ipomoea lacunosa L., Pitted Morning-glory (N; 
RLM, RUM, TLM) N 8517, 948, 2311, 2385 

Ipomoea pandurata (L.) G.EW.Mey, Bigroot Morn- 
ing-glory (N;TLM, TUM) N 2074, 2689 

Jacquemontia tamnifolia (L.) Griseb., Hairy 
Clustervine (I) Taubenhaus 2642 (TAES) 


CORNACEAE 
Cornus drummondii C.A.Mey, eae Dog- 
ood (N; TLM, TLW) N 49, 632, 164 
is florida L., Flowering ee (N; TLW) 
N 1331, 2103 


CRASSULACEAE 
Crassula aquatica (L.) Schoenl., Water Pygmy- 
weed (N;TLM,TLO, TLW) N 1427, 1549, 2478 


CUCURBITACEAE 

Cucumis melo L., Muskmelon (1; TLM) N 2384 

— pepo L.var. texana (Scheele) D. — 

as Gourd (N; RLM) N 288, 813,214 
ae pendula L., Melonette (N; a4 RUM, 
TLM, TUW) N 160, 703, 933, 2023, 2291 

CUSCUTACEAE 

Cuscuta Sees Engelm., Cusp Dodder (N; 
PAR) N 7 

Cuscuta pines Choisy, Cluster Dodder (N; 
PAR) N 2201 


NEILL AND WILSON 


1 TEXAS 


Cuscuta obtusiflora Kunth. in H.B.K. var. 
glandulosa Engelm., Red Dodder (IN; PAR) N 
134, 2202 


DROSERACEAE 
Drosera brevifolia Pursh., Annual Sundew (N;TLO) 
N 2473, 2512 


EBENACEAE 
a virginiana L., Virginia Persimmon (N; 
TLO, TUO) N 140, 922, 1455, 1648 


ELATINACEAE 
Flatine brachysperma A. Gray, Shortleaf 
Waterwort (N;TLM, TLW) N 1288, 1428, 1551 


ERICACEAE 
Vaccinium arboreum Marsh. ae - TUM, 
TUW) N 100,211,557, 1074, 1128, 1 


EUPHORBIACEA 
— gracilens a Gray, Slender Copperleaf (N; 
_TLM, TUO) ieee 2159, 2190 

lyph ) elm. ex A. Gray) L.Mill 
7 Gandhi One- a Copperleaf (N; TUM, 
TUOQ) N 265, 704, 788 

Acalypha rhomboidea_ Raf., 
Copperleaf (N; RLM) N 840 

Acalypha virginica L., Virginia Copperleaf (N;RUM, 

QO) N 931, 2158, 2684 

Cnidoscolus texanus (Muell.-Arg.) Small., Texas 
Bullnettle (N; TLM, TUW) N 550, 1674 

Croton argyranthemus Michx., Silver Croton (N; 
TUM) N 2056 

Croton capitatus Michx.,Woolly Croton (N; RLM, 
TLO) N 800, 962 


Rhomboid 


Croton capi Michx. var. lindheimeri (Engelm. 
& A.Gray) Muell.-Arg., Woolly Croton (N;TUQ) 
/ 


Croton glandulosus L.var.glandulosus, Tropic Cro- 
ton (N;RLM, RUM, TUO) N 745, 896, 2409, 2452 
Croton glandulosus L. var. Cees ie 
eats Croton (N; TUO) N 
nal 1s L. var. septentrionalis Muell.- 
Arg., Northern Croton (N; TUM) N 210 
Croton lindheimerianus Scheele — var. 
lindheimerianus, Three-seed Croton (N;RLM) 


N 799 
Croton monanthogynous Michx., One-seed Cro- 
n (N;TLO, TUM) N 927, 21717, 2283 
Croton texensis (Klotzch.) Muell.-Arg., Texas Cro- 
ton (N; TUQ) N 1924 


1103 


— michauxii G.L. Webster, en Cro- 
n (N; TUM, TUO) N 257, 708, 2057, 2304 
ee bicolor Engelm. & A. a on, 
the-prairie (N; TUO) N 787, 2288 
Euphorbia corollata L.,Flowering Spurge (N; TUO) 


Euphorbia dentata Michx., Toothed Spurge (N; 
RLM, TLO, TUM, TUO) N 570, 936, 1948, 2128, 
2217 

Euphorbia maculata L., Spotted Spurge 
TLO, TUO) N 204, 798, 952, 2129 

Euphorbia nutans Lag., Eyebane (N; RLM, TLM, 
TLO, TUO) N 886, 1025, 1046, 2130, 2243 

Euphorbia prostrata Aiton, Prostrate Spurge (N; 
TLO, TUM) N 754, 1827, 2284 

meee serpens Kunth. in H.B.K., Mat Spurge 
(N;TLO) N 2374 


— 


N;RLM, 


pr (ata Lam. ane ale (N; 
UO) N 1448, 1749, 
on ae sa tote (N; 
TLO) N 2514 


Euphorbia texana Boiss., Texas Euphorbia (N;TLO, 
M, TUO) N 418, 1539, 1900 
a bil abnormis Baill. var. abnormis, 
Drummond Leaf-flower (N; TUQ) N 2407 
ae eae ne Walter ssp.caroliniensis, 
Carolina Leaf-flower (N; TLM, TUM) N 2408, 
2685 
Phyllanthus pudens L.C.Wheeler, Birdseed Leaf- 
flower (N;TLM, TLO) N 2789, 2373 
Ricinus communis L., Castorbean (1; TUQ) N 237 
Sapium sebiferum (L.) Roxb., Chinese Tallowtree 
(l; RUM) N 143, 629, 1028, 1125 
Stillingia sylvatica - sea Delight (N; TLO, 
TUM, TUO) N 37, 1794 
gs betonicielia — Betony Noseburn (N; 
, UO) N 644, 690, 2053 
rage mine Engelm. & A. Gray, Shortspike 
urn (N; TLO, TUO, TUW) N 926, 1475, 


Tragia ramosa Torr., Catnip Noseburn (N;TUO) N 
2700 
Tragia urticifolia Michx., Nettleleaf Noseburn (N; 
M) N 2223 
FABACEAE (sensu stricto) 
Aeschynomene indica L., Joint Vetch (N) McLeod 


Amorpha fruticosa L.,Indigobush (N;RLM) N 1398 
Apios americana Medik., American Potato-bean 
(N; RUM) N 2084, 2680 


1104 


fs distortus T.&G.var.engelmannii (Sheld.) 
es, Bentpod Milkvetch (N; TUO) N 367, 


se 
Astragalus joes T.&G,, Slimpod Milkvetch 
(N; RUM) N 2733 
Baptisia alba (L.) Vent var. macrophylla (Larisey) 
Isley, White Wild Indigo (N; RLM, TLO) N 462, 
1838 
Baptisia bracteata Muhl. ex Ell. var. glabrescens 
(Larisey) Isley, Leafy Wild Indigo (N;TLM, TUM) 
136,414 
Baptisia nuttalliana Small, Nuttall Wild Indigo (N; 
TLO, TUQ) N 572, 1478 
Baptisia sphaerocarpa a Green Wild Indigo 
(N; TLO, TUM) N 527,6 
a Benth. Butterfly Pea 
N. TUM, TUW) N 254 691 
itoria mariana he Atlantic Butterfly Pea (N;TUW) 
N 1909, 266 


C 


— 


Crotalaria | rae les ats nl (N; TLO, 
TUO, TUW) N 650, 694, 

Dalea compacta Spre ak cee (A. Gray) 
Barneby, Plains Prairie Clover (N; TUM) N 1867 

Dalea multiflora (Nutt.) Shinners, Roundhead 
Prairie Clover (N; TUM, TUQ) N 131, 2050 

Dalea phleoides (T.&G.) Shinners var. microphylla 

T.&G.) Barneby, Longbract Prairie Clover (N; 
TUM) N 250 

Dalea purpurea Vent. var. purpurea, Purple Prarie 
Clover (N) McLeod s.n. (TAES) 

Dalea villosa (Nutt.) Spreng. var. grisea (T.&G,) 
Shinners, Oklahoma Prairie Clover (N; TUO) 
N 2635 


— 


Desmodium canescens (L.) DC., Hoary Tick-clover 
(N; TUM, TUW) N 666, 2660 

Desmodium ciliare (Muhl. ex Willd.) DC., Littleleaf 
Tick-clover (N; TLW, TUM, TUO) N 997, 2327, 


2432 

Desmodium ane um (Michx.) DC,, Tick-clover 
(N; TLW) NV 

Desmodium pao (L.) DC. var. 
paniculatum, Panicled Tick-clover (N; TLO, 
TLW) N 959, 240? 


Desmodium sessilifolium (Torr) T.&G., Sessileleaf 
Tick-clover (N; RUM, TUM, TUW) N 209, 594, 


2022 
Desmodium Ca oe DC., Velvetleaf Tick- 
clover (N; TLW) N 2 
Erythrina herbacea L., Crea (N; TLW, TUM, 
TUW) N 530, 1672, 1 


BRIT.ORG/SIDA 19(4) 


Galactia regularis (L.) B.S.P, Downy Milkpea (N; 
TUM, TUO) N 643, 2648, 2686 

Glottidium vesicarium (Jacq.) Harper, Bagpod 
Sesbania (N;RUM, TUO) N 278, 921, 1068, 2320 

Lathryus hirsutus L., Singletary Pea (I; TUO) N 21, 


1607, 2570 
Lathyrus pusillus Ell, Low Pea (N;TLO, TUO) N 1509, 
2557 


Lespedeza cuneata (Dumont) G. Don., Sericea 
Bushclover (I; TUO) N 1632 

Lespedeza hirta (L.) Hornem.,Hairy Bushclover (N; 
TLM, TUM) N 2245, 2676 

Lespedeza repens (L.) W. Bart., Creeping 
Bushclover (N; RUM, TUO) N 680, 1088 

Lespedeza striata (Thunb.) Hook & Arn., Japanese 
Bushclover PA Sah N 730, 2307 

Lespedeza stuevei Nutt., Stueve oo (N; 
TLM, TUM, TUO) N 777, Bi 205 

aan virginica (L.) Britton, ae er 
Bushclover (N; TUM, TUO) N 779, 1081, 2249 

Lotus che (Hook.) Benth. var. unifoliatus, 
Pursh Deervetch (N; TLO) N 626, 2638 

Lupinus eee Hook., Texas Bluebonnet (N; 
TLO,T 


<7 


Lupinus texensis Hook., Texas Bluebonnet (N; 
N 1752 
Medicago arabica (L.) Huds., Spotted Medic 


= 


N 1236 
Medicago lupulina L., Black Medic (1; TLO) N 353 
Medicago orbicularis (L.) Bartal., Button Clover (I; 
TLO)N: 5 


’ 


Medicago polymorpha L., Medic (1; RUM, TLO) N 
1822, 2505 

Medicago sativa L., Alfalfa (I; TUO) N 1650 

Melilotus indicus (L.) All, Yellow Sweetclover (I; 
TLO, TUO) N 1680, 1823 

elilotus officinalis (L.) Lam.,, Yellow Sweetclover 

(1) Fowler 129 (SHST) 

Orbexilum pedunculatum (Mill) Rydb. var. 
pedunculatum, Sampson's Snakeroot (N; 
TLW) N 

Orbexilum simplex (Nutt. ex T.&G.) Rydb., 
Singlestem Snakeroot (N;TUM, TUO) N 7808, 
2006 


cb) 


Pediomelum hypogaeum (Nutt. ex T.&G,) Rydb. 
var. subulatum (Bush) Grimes, Edible Scurfpea 
(N; TUM) N 552, 2732 

Pediomelum rhombifolium (T. & G.) Rydb., 
Roundleaf Scurfpea (N; TLO, TUM, TUO) N 
1868, 1933, 2 


NEILL AND WILSON, TEXAS 


Psoralidium tenuiflorum (Pursh.) Rydb., Slimleaf 
Scurfpea (N) Clark 415 (TAES) 

Pueraria lobata (Willd.) Ohwi, Kudzu (1; TUW) N 
1420 


Rhynchosia latifolia Nutt. ex T.&G., Broadleaf 
Sno sei ue TUM, TUO, TUW) N 593, 
2049, 2 
Robinia ie ae ' — Locust (|; TUM) N 2750 
Robinia ae acacia L., Black Locust (N; TUM, 
TUO) N 1242, 1619 
Sesbania piace (Rydb.) Cory, Rattlebush 
(N; RLM, RUM) N 977, 2165 
i herbacea (P. Mill) Pe Bequilla (N; 
M, RUM, TLO) N 974, 1038, 2319 
a dffinis T.&G., Eve's Necklace (N;RLM,TLM, 
TUO) N 580, 882, 1469, 1 &9 
Strophostyles helvula (L.) Ell. Trailing oo 
RLM, RUM, TUM) N 95, 683, 878, 232 
Strophostyles lei (T.&G,) Piper, fies 
Wildbe ean (N; TLO, TUOQ) N 699, 2008, 2639 
Stylosanthes biflora (L.) B.S.P., Pencilflower 
TUOQ) N 205, 1685, 1711 
i onobrychoides Nutt., 
Hoary Pea (N; TUM) N 79, 2048 
Tephrosia virgit ana (L.) Pers., Virginia Hoary Pea 
(N; TLW) N 2450 
Trifolium campestre Schreb. in Sturm. Low Ho 
Clover (I; TLO, TUO) N 1538, 1805, 2594, 7615 
Trifolium carolinianum Michx.,Carolina Clover (N; 
,TUO, TUM) N 431, 1274, 1541, 2546 
Trifolium incarnatum L., Crimson Clover (|; TUO) 


N ; 


— 


Multibloom 


N519 

Trifolium polymorphum Poir., Peanut Clover (N; 

W,TUW) N 1473, 2551 

Trifolium resupinatum L., Persian Clover (1; TLO, 
TUO) N 2556, 2601 

Trifolium vesiculosum Savi., Clover (I; TUO) N 16817, 
1758 

Vicia ludoviciana Nutt. ssp./udoviciana, Louisiana 
Vetch (N; TLM, TLW, TUM) N 420, 1266, 2575 

Vicia lutea L., Yellow Vetch (|; TUO) N 1458, 2618, 


2630 

Vicia minutiflora Dietr., Small-flowered Vetch (N; 
RUM, TLM, TLW) N 397, 1226, 12 

Vicia sativa L. ssp. nigra (L.) Ehrh., Narrow-leaved 

Vetch (I; RUM, TLO) N 1193, 2506 

Vicia villosa Roth, Hairy Vetch (1; TUO) N 1220 

Vicia villosa Roth. var. ee labrescens Kuch., Hairy 
Vetch (|; TUO) N 


1105 


Vicia villosa Roth. ee a (Host.) Corb., Hairy 
Vetch (I; TUO) N 

Vigna unguiculata ‘kr ales Black-eyed Pea (I; 
TLO) N 2203 

Wisteria sinensis (Sims) Sweet, Chinese Wisteria 


(1; TUW) N 735 


a= 


FAGACEAE 

Quercus falcata Michx., Southern Red Oak (N;TLO, 
TUM, TUW) N 77, 1424, 1738, 1788, 1908 

Quercus lyrata Walter, Overcup Oak (N; RLM, TLM) 
N 287 855, 426, 1644, 1898, 2209 

Quercus macrocarpa Michx., Bur Oak (N; TLM, 
TLO) N 


— 


01,2585 
Quercus marilandica Muench., et _ Oak (N; 
TUO, TUW) N 1494, 1691, 2096, 


Quercus nigra L., Water Oak at aon 
270, . 1999 


Quercus phellos L., Willow Oak (N; RLM, TLW) N 
471, 1450 

Quercus shumardii Buckl., Shumard Oak (N; RUM) 
N 2097 


Quercus stellata Wang. var. margaretta (Ashe) 
Sarg., Sand Post Oak (N; TUM) N 1787, 1841 

Quercus stellata Wang. var. paludosa Sarg., Bot- 
tomland Post Oak (N; TLW, TUW) N 1086, 2131 

Quercus stellata Wang. var. stellata, Post Oak (N; 
TUM) N 73, 2376 

a Mill, Live Oak (N;TUO) N 1872 


UEICUS virgl 


FUMARIACEAE 
Corydalis micrantha (Engelm.) A. Gray var. aus- 
tralis (Chapm.) Shinners, Southe 
Scrambled Eggs (N; TLW, TUM, TUO) NV ph 
1201, 1278 


GENTIANACEAE 

Sabatia campestris Nutt., Prairie Rose-gentian (N; 
TUO) N 28, 612, 1714 

GERANIACEAE 

Geranium carolinianum L., Carolina Crane’s-bill 
(N: TLO, TUO) N 458, 1408, 2645 

Geranium dissectum L., Crane’s-bill (I; TLO) N 1446 


HAEQRAGACENE 


tl 


[r) NAG 


Changeleaf 
Parrot’s Feather (N: AFL, ici “ 1550, 1936, 
001, 25276 

Myriophyllum pinnatum (Walter) B.S.P., Gree 
Parrot's Feather (N; AFL, ASU) N 410, ok 
] 5 


1106 


HAMAMELIDACEAE 
Liquidambar styraciflua L., Sweet Gum (N; TLW) 
N 669, 2524 


HIPPOCASTANACEAE 
Aesculus pavia L.var.pavia, Red Buckeye (N;TLM) 


HYDROPHYLLACEAE 

Hydrolea ovata Nutt. ex Choisy., Hairy Hydrolea 
(N; AEM, RLM, RUM) N 187, 686, 803, 1101 

Nemophila aphylia (L.) Brumm, Smallflower Ne- 
mophila (N; TUW) N 1245 

Phacelia glabra Nutt., Smooth Phacelia (N; TLM) 
N 1227 

Phacelia hirsuta Nutt., Hairy Phacelia (N; TLO) N 


2550 
Phacelia patuliflora (Engelm. & A. Gray) A. Gray, 
Sand Phacelia (N; TLW) N 1667 


JUGLANDACEAE 

Carya alba (Mill) K.Koch., Mockernut Hickory (N; 
TLW, TUQ) N 2095, 2627 

Carya aquatica (Michx.f) Nutt, Water Hickory (N; 
RLM, RUM) N219 856, 1012, 1886 

Carya illinoinensis (Wang.) K.Koch., Pecan 
TLO) N 200, 902 

Carya texana Buckl., Black Hickory (N; RLM, TUM, 

JO) N 174,218, 785, 1615 
Juglans nigra L., Black Walnut (N; TLO, TLW) N 
4 


— 


N-TLM, 


KRAMERIACEAE 

Krameria lanceolata Torr, Rattany-vine (N; TUM 
TUQ) N 710, 1860 

LAMIACEAE 

Hedeoma hispida Pursh., Rough Mock Penny- 
bd a hs. aay N 1502, 1653, 1688, 2597 

Lam e L., Henbit (N; TLO, TUQ) NV 


Lycopus rubellus Moench, Arkansas Bugleweed 


Monarda citriodora Cerv. ex Lang. ssp. citriodora 
Lemon Beebalm (N; TLO, TUO) N 622, 1817, 
20270 


Monarda clinopodioides A.Gray, Basil Beebalm (N; 
TUO) N 2602 
dao L., Wild el (N;TLM, TLO, 
M) N 138, 647, 206 
ee lindheimeri on x A.Gray ex A.Gray, 
Lindheimer Bergamot (N; TUM, TUO) N 76, 


BRIT.ORG/SIDA 19(4) 


a punctata L.var.intermedia (McCI.& Epl.) 
erfall, Spotted Beebalm (N; TLO) N 37 

Mond palaaia - var.lasiodonta A. Gray, Spot- 

W,TUM,TUQ) N 1864, 1930, 


— 


1960 
Perilla frutescens (L.) Britton, Beefsteak Plant (| 
RLM) N 2334 
ie ie ifolia Fern. Obedient-plant (N; 
M) N 


Ps id nem (Nutt.) Engelm.& A.Gray, 
media a plant (N; TLM, TLO) 
i oe oe 
Physostegia en Lundell, Obedient-plant (N; 
TLM, TLO, TUM, TUO) N 41, 1436, 1638, 2559 
Prunella vulgaris L. var. vulgaris, Self-heal (I; TLW, 


M) N 507, 1518 
Prunella vulgaris L. var. hispida Benth. Self-heal (|; 
TUM) N87 


Salvia azurea Lam., Blue Sage (N; TUM, TUCO) N 
330, 2248, 2418 


Salvia coccinea PJ. Buchoz ex Etlinger, Tropical 
arlet Sage (N; TLW, TUM) N 678, 767 
Salvia lyrata L., Lyreleaf Sage (N; TLW, TUM) N 106, 
509, 1254 


ne 


Scutellaria cardiophylla Engelm. & A. Gray, 
nea ea ialieap (N; nae stl N oe 679 


ap (N; TLW) NV 
Scutellaria ovata sleet Skullcap (N; TUW) N 
197] 
Scutellaria parvula Michx.var./eonardii (Epl.) Fern., 
kullcap (N; TLO, TUW) N 1430, 2596 
Scutellaria parvula Mich.var. parvula, Skullcap (N; 
N 1961 
Stachys crenata Raf, Shade Betony (N;TLM) N 2528 
~~ aes Willd., Slenderleaf Betony (N; 
M)N1 


= 


etd ne L. var. os German- 
N; TLW, TUO) N 640, 

oe dichotomum L., nae Bluecurls (N; 
UM) N 2253, 2289 

LAURACEAE 

ean (Nutt.) Nees., Sassafras (N;TLW, 

TUW) N 1717, 403, 1419 
LENTIBULARIACEAE 
Utricularia gibba L., Cone-spur Bladderwort (N; 
9] 


Utricularia adete Small, Whorled Bladderwort 


(N; AFL) N 1969 


NEILL AND WILSON 


MIAVIOVUIN LYVUINT I, TEXAS 


LINACEAE 

Linum medium (Planch.) Britton var. texanum 
(Planch.) Fern, Texas Flax (N; RUM, TLO, TUO) 
N 600, 1744, 2040, 2603 


LYTHRACEAE 

Ammannia x coccinea Rottb., Purple Toothcup 
(N; RLM, RUM, TLM) N 876, 894, 2161, 2193 

Cuphea carthagenensis (Jacq.) J.F. Machr., (N; 


M) N 2682 

Cuphea glutinosa Chapm. & Schlecht., (1; TUM, 
TUO) N 2633, 2681 

Didiplis diandra (Nutt. ex DC.) Wood., Water- 
purslane (N; ASU) N 1970 

Lagerstroemia indica L., Crepe Myrtle (1; TLM) N637 

Lythrum alatum Pursh.var.lanceolatum (Ell.) T.&G., 
La S al (N; TLO, TUM, TUQ) N 
183,21 690 

Rotala ramosi i .) Koehne, Tooth-cup (N; TUM) 
N 2677 


MALVACEAE 

Callirhoe Use (Torr.) A. Gray. var. 
involucrata, Low Wine-cup (N;TLO) N 32 

Callirhoe renee (T.&G.) A. Gray var. 


lineariloba (T.&G.) A. Gray, Low Wine-cup (N; 
TUO) N 382, 1495 

Ca ies Soest ee Gray, Woods Wine-cup 
(N; TUM, TUO) N 


Toe ee All., aber leaved Hibiscus (N; 
RUM 


— 


Hibiscus Resa ee ssp. lasiocarpus (Cav.) O.J. 
Blanchard, Swamp Rosemallow (N;RLM, TLO) 
N 191,899, 1916 
Hibiscus syriacus L., Rase-of-Sh 
Malvaviscus arboreus Dill ex Cav.var.drummondii 
(T.&G.) Schery, Drummond Turk’s-cap (N; 
, U0, TUW) N 750, 1905, 2120, 2318 
Modiola caroliniana (L.) G.Don.,Carolina Modiola 
(N; TLO, TUO) N 169, 1407, 1818 
Sida rhombifolia L., Arrowleaf Sida (N; TUM) N 


(1-TINAL) ALA ZA 
t 


2085 

Sida spinosa L., Prickly Sida (N; RLM, TLO, TUM) N 
822,897, 2264 

MELASTOMATACEAE 

Rhexia mariana L. var. mariana, Meadow Beauty 
(N; RUM) N 267 

MELIACEAE 

Melia azedarach L., Chinaberry (1; RUM, TUM) N 


1107 


MENISPERMACEAE 
Cocculus carolinus (L.) DC., Carolina Snailseed (N; 
RLM, TLM, TUO) N 1517, 728, 935, 2024 


MIMOSACEAE 
Acacia farnesiana (L.) Willd., Huisache (N; TUO) N 


1397, 2137 
Albizia julibrissin Durazz., Mimosa (1; RUM, TLW) 


161,628 
Desmanthus illinoensis (Michx.) MacM. ex Rob- 
ins & Fern,, Illinois Bundleflower (N; RUM, TLO, 
UO) N 162, 620, 1834, 2111, 2623 
Mimosa nuttallii (DC.) B.L. Turner, Catclaw Sensi- 
tive-briar (N; TLO, TUM) N 1462, 2106 
Mimosa Strigillosa T. & G., Herbaceous Mimosa 
(N; RLM, TLO) N 174, 957 
Neptunia lutea (Leavenw.) Benth., Yellow 
Neptunia (N; TUM, TUO) N 109,613, 1692 
Neptunia pubescens Benth. var. poe Prai- 
rie Neptunia (N; RLM, TUO) N 909, 1833 
P is glandulosa Torr. var. sR osa, Honey 
Mieceaie (N; TLO, TUM, ie N 93, 524, 762, 
1877 


MOLLUGINACEA 
-_ a (R. és P) Rohrb,, (I; RLM, TLW) N 79335, 


ies verticillata L.,Green Carpetweed (|; RLM, 
TLM, TUM, TUO) N 260, 693, 892, 1862, 2197 

MORACEAE 

Ficus carica L., Fig (I; TLM) N 1400 

Maclura pomifera (Raf.) Schneid., Bois D’Arc (N; 
TLW, TUM) N 263, 7 

Morus rubra L., Red Mulberry (N; TLW, TUW) N86, 
442, 1640, 2093 


Ow 
GH 
ion 


MYRICACEAE 
oe cerifera L., Southern Wax-myrtle (N; RUM, 
M)N 419, 591, 1085, 1782 


NELUMBONACEA 
Nelumbo lutea aie Pers., Yellow Lotus (N; 
AEM) N 2146 


NYCTAGINACEAE 
Mirabilis albida (Walter) Heimerl., White Four- 
O'Clock (N; TLM, TUM, TUO) N 46, 273, 994, 
94/7 


Mirabilis jalapa L., Four-O'Clock (|; TLM) N 144 

Mirabilis linearis (Pursh.) Heimerl., Linearleaf Four- 
O'Clock (N; TUM) N 2169 

Mirabilis nyctaginea (Michx.) MacM., Wild Four- 
O'Clock (N; TUM) N 533, 1704 


1108 


NYMPHAEACEAE 

Nuphar lutea (L.) Sm. ssp. advena Kartesz & 
Gandhi, Spatterdock (N; AFL) N 1829, 2148 

Nymphaea odorata Aiton ssp.odorata, American 
Waterlily (N; AEM) N 721 

NYSSACEAE 

Nyssa sylvatica Marsh. var. sylvatica, Black Tupelo 
(N; RLM, RUM) N 1405, 2099, 2399 


Forestiera acuminata (Michx.) Poir. Swamp Privet 
M) N 400, 447, 2547 

Forestiera ligustrina (Michx.) Poir, Privet Forestiera 
(N; RLM, RUM, TLW, TUW) N 289, 743, 1052, 
1131, 2047, 2102 

Fraxinus americana L., White Ash (N; TLM, TLW, 
TUW) N 376, 377, 2101, 2211 

Fraxinus pennsylvanica Marsh., Red Ash (N; RLM, 

M) N 234, 435, 2588 

Ligustrum lucidum Aiton f., Wax-leaf Ligustrum 
(I; TLW) N 673 

Ligustrum sinense Lour., Chinese Privet (|; TLM, 
TUM, TUO, TUW) N 40, 336, 756, 1129, 1621 


NAGRACEAE 
Calylophus berlandieri Spach. ssp. pinifolius (A. 
Towner, Sundrops (N) Clark 474 (TAES) 
Calyophus serrulatus (Nutt.) Raven, Yellow Sun- 
drops (N; 
Gaura brachycarpa Small, Plains Gaura (N; TLO, 
TUQO) N 523, 1477, 2626 
Gaura longiflora Spach., Kearny Gaura (N; TLO, 
TUO) N 267, 2125, 2265 
Gaura sinuata Nutt. ex Ser, Wavyleaf Gaura (N; 
M) N S62 
Gaura suffulta Engelm. ex A. on suffulta 
oe adside oe (N; Heit 


| It 


L., Bushy seb i RUM, 
TLM, TUO, TUW) ee 734, 2009, ? 
Ludwigia decurrens Walter, Primrose- iey (N; 
AEM, RLM, RUM) N 841, 972, 2163 
Ludwigia knee Walter ssp. brachycarpa 
(T.&G.) Peng., Torrey Seedbox (N; RUM) N 


‘ 


wa 


2039 
Ludwigia glandulosa Walter ssp. glandulosa, 
seedbox (N; RLM, TLO) N 793, 1883 
Ludwigia palustris (L.) Ell, American Seedbox (N; 
RLM, RUM, TLO) N 1554, 1747, 1987, 2038 
Ludwigia peploides (Kunth. in H.B.K.) Raven, 


mooth Water-primrose (N; AEM, RUM) N 


sm 
212, 1717, 2164, 2595 


BRIT.ORG/SIDA 19(4) 


Oenothera heterophylla Spach. ssp. heterophylla 
Varileaf Evening Primrose (N;TLM, TLO, TUO) 
N 30, 1968, 2191 

Oenothera laciniata Hill, Cutleaf Evening Prim- 
rose (N; TLO, TUM, TUO, TUW) N 393, 1417, 
1816, 1863 

Oenothera linifolia Nutt., Threadleaf Sundrops (N; 
TUM, TUO) N 1521, 1718, 2735 

Oenothera spachiana T.&G., Spach Evening Prim- 
rose (N;TUO) N 2610 

Oenothera speciosa Nutt., Showy Evening Prim- 
rose (N; TLO, TUO) N 457, 1480 

OROBANCHACEAE 

robanche multiflora Nutt., Largeflower Broom- 

rape (N; PAR) N 520, 1649 

OXALIDACEAE 

Oxalis corniculata L., Creeping Woodsorrel (1; 


YN 1251 
Oxalis dillenii Jacq., Dillen Woodsorrel (N;TLM) N 


164 
Oxalis rubra St.-Hil., Pink Shamrock (I; TLM) N 1243 
Oxalis stricta L., Yellow Woodsorrel (N; TUM, TUO) 


Oxalis violacea L., Violet Woodsorrel (N;TLM, TUM) 
N 1516, 2290 


PAPAVERACEAE 
Argemone albiflora Hornem. ssp. texana G. 
wnbey, White Prickly Poppy (N; TUM) N 
1858 
Argemone polyanthemos (Fedde.) G. Ownbey, 
White Prickly Poppy (N; TLO) N 563 


PASSIFLORACEAE 
Passiflora incarnata L., Maypop (N; TLO, TUO) N 
) 2114 
Passiflora lutea L., Yellow Passionflower (N; TLW, 
TUO, TUW) N 272, 725, 1676, 241 


PHYTOLACCACEAE 

Phytolacca americana L., Pokeweed (N; RUM, 
TU W) N 145, 729, 1659 

Rivina humilis L., Rougeplant (N; TUM) N 274 


PLANTAGINACEAE 
Plantago aristata Michx.,Bottlebrush Plantain (N; 
O, TUO) N 35, 1609, 1693 
Plantago heterophylla Nutt., Simplespike Plantain 
(N;TUQ) N 1517 
Plantago hookeriana ie : CA. Mey., Hooker 
Plantain (N; TLW) N 


NEILL AND WILSON 


TEXAS 


Plantago patagonica Jacq, Bristlebract Plantain 
(N; TUM) N 567 

Plant hodosperma Dcne., Redseed Plantain 
(N; TLM) N 2529 

Plantago virginica L., Paleseed Plantain (N; TLM, 
TUM) N 45, 441, 538 

Plantago wrightiana Dcne., Wright Plantain (N; 
TUO, TUW) N 649, 2617 


PLATANACE 
Platanus occidentalis L., American Sycamore (N; 
RUM, TLW) N 893, 1586 


POLEMONIACEAE 

Ipomopsis rubra (L.) Wherry, Standing Cypress (N; 
TUO) N 186 

Phlox cuspidata Scheele var. cuspidata, Pointed 
Phlox (N; TUO) N 344, 348, 1499 

Phlox drummondii Hook. var. peregrina Shinners, 
Drummond Phlox (N; TUO) N 158, 2560 

Phlox pilosa L., Downy Phlox (N; TUM) N 1492 


POLYGALACEAE 
Polygala incarnata L., Pink Milkwort (N; TLO) N 
5/3 


= 


Polygal / Walter, Bitter Milkwort (N;TLO, 
TUO) N 1716, 1737, 2607 2655 

Polygala verticillata L,Whorled Milkwort (N;TUO) 
N 732 


POLYGONACEAE 

Brunnichia ovata (Walter) Shinners, oo 
Vine (N; RLM, TLO) N 116, 223, 956, 

Eriagonum longifolium Nutt., eee ae Buck- 
wheat (N; TUM) N 2220, 2272 

Polygonum hydropiperoides Michx. mp 
Smartweed (N; RLM, RUM, TLO, TUO) ‘i 692, 
968, 1611, 1851 

Polygonum lapathifolium L., Curltop Smartweed 
(N; TLO, TLW) N 7922, 2200 

olygonum pensylvanicum L., Pennsylvania 

Smartweed (N; RLM, TLO) N 295, 969, 2336 

ce ead persicaria L., Lady's Thumb (1; RLM) N 


Pol yonumpunett Ell, Water Smartweed (N; 
RLM, TLO) N 296, 843, 2335 
Polygonum ramosissimum Michx., Bushy Knot- 
weed (N;RLM) N 2333 
Polygonum setaceum Baldw., Smartweed (N; 
M) N 189 


Rumex chrysocarpus Moris, Pale Dock (N;TUO) N 
1772, 2268 


1109 


Rumex crispus L., Curly Dock (1; TLO) N 1241, 1897 

Rumex hastatulus Baldw., Heart Sorrel (N; TUM, 
TUO) N 24, 537, 1472 

Rumex pulcher L., Fiddle Dock (1; TLM, TLO, TUO) 


, 578, 1 
Rumex verticillatus L., Swamp Dock (N; RLM) NV 
113 


PORTULACACEAE 
Claytonia virginica L., Spring Beauty (N; TUO) N 
3,1198 
tul | L. Common Purslane (1; TLO 
951 
Portulaca pilosa L., Chisme (N; TUO) N 2179, 2309 
Portulaca umbraticola H.B.K.,Wingpod Portulaca 
M,TUOQ) N 240 
Talinum parviflorum Nutt., Prairie Flameflower (N; 
711 


PRIMULACEAE 
Anagallis arvensis L., Scarlet Pimpernel (1; TUO) NV 


<= 


2 = 


tunculus minimus L., Chaffweed (|; TLM, TLO, 
TLW, TUM) N 1261, 1328, 1548, 1755, 2477 
(Anagallis minima (L.) E.H.L. Krause) 

Samolus valerandi L.ssp.parviflorus ee Hultén, 
Thinleaf Brookweed (N; TLW) N 


RANUNCULACEAE 

Anemone berlandieri ia Tenpetal Anemone 
(N; TUO) N 357,250 

Anemone caroliniana te Carolina Anemone 
(N; TLW, TUO) N 390, 392, 2500 

Clematis pitcheri T.&G. var. pitcheri, it Clema- 
tis (N; RLM, TUM) N 585, 802, 932, 

Clematis reticulata Walter, Netleaf ae (N; 
RLM, TUM) N 1885, 2140 


Delphinium carolinianum Walter ssp. vimineum 
(D.Don) Warnock, Carolina Larkspur (N; TLO, 
TUO) N 38, 510, 1210 

Delphinium carolinianum Walter ssp. virescens 
(Nutt.) R.E. Brooks, Plains Larkspur (N) Clark 
362 (TAES) 

Myosurus minimus L., Tiny Mousetail (N; RLM, 
TUQ) N 405, 2532 

Ranunculus fascicularis Muhl. ex Bigel., Tufted 
Buttercup (N;TLO, TLW) N 391, 1931, 2513 

Ranunculus hispidus Michx. var. nitidus (Chapm.) 

n, Bristly Buttercup (N; TLW) N 71250 

Ranunculus laxicaulis (T.&G.) Darby, Small Butter- 

cup (N;TLW) N 1249, 1475 


1110 


Ranunculus muricatus L., Roughseed Buttercup 
(I; RUM) N 2502 

Ranunculus fee L. Sticktight Buttercup (I; 
RLM, TLM, TLW, TUM, TUO) N 411, 1421, 1545, 


1831, 248 
Ranunculus ae Poir,Weak Buttercup oe 
R M, TUOQ) N 412,540, 1531, 253 
Raaunael us sardous Crantz., Buttercup A RUM, 
TLM, )N 65,345, 395 


Ranunculus sceleratus L., Blister Buttercup (N; 
RUM) N 1984 


Thalictrum dasycarpum Fisch. & Ave.-Lall., Purple 
dow-rue (N) Cory 54579 (LL) 
RHAMNACEAE 
Berchemia a (Hill) K.Koch., Rattanvine (N; 
LM, TL M, TUW) N 42, 667, 1064, 1537 

Rhamnus ees Walter, Carolina Buckthorn 
(N; TLW) N 2070, 2136, 2738 

ROSACEAE 

Crataegus marshallii Eggl., Parsley Hawthorn (N; 
TLW, TUM) N 264, 1140, 1324, 1522, 2454 

Crataegus spathulata Michx., Littlehip Hawthorn 
( ) N 1297 


Crataegus viridis L., Green Hawthorn (N;RLM, TLW, 
)N 402, 427,895, 1296, 2537 

Geum canadense Jacq. var. camporum (Rydb,) 
Fern, White Avens (N; RUM, TUW) N50, 1411 

Geum canadense Jacq. var. texanum ie & 

eath., White Avens (N; TLW) N 7155 

Prunus peee ia Marsh, Chickasaw eile (N; 

TUM) N 


Prunus oe niana : Aiton, Cherry-laurel (N; 
RLM) N 1196, 2 

Prunus gracilis Eng a & es Gray, Oklahoma Plum 
(N; TUO, TUW) N 398, 

Prunus mexicana S.Wats os Plum (N;TLW, 
TUM, TUO, TUW) N 358, 375, 1504, 2015, 2274 

Prunus persica (L.) Batsch., Peach (|; TUM) N 757 

Prunus serotina Ehrh. var. serotina, Black Cherry 
(N; TLW, TUM) N 1464, 2104 

Pyracantha coccinea M.J.Roem., Scarlet Firethorn 
(; TUM, TUO) N 752, 11.20 

Pyrus calleryana Decne., Callery Pear (I; TUM 


N 


<7 


59, 
Pyrus communis L., Common Pear (|; FUM) N 2493 
Rosa carolina L., Carolina Rose (N; TUM) N 2757 
Rosa multiflora Thunb. ex Murr., Japanese Rose 
(I; TUQ) N 1467, 1636 


BRIT.ORG/SIDA 19(4) 


Rubus aboriginum Rydb., Blackberry (N;TLW,TUO) 
N 1761, 1814 

Rubus argutus Link., Blackberry (N; RUM) N 2737 

Rubus flagellaris Willd., Northern Blackberry (N; 
TLM, TLW) N 1260, 2604 


— trivialis Michx., Dewberry (N; TUO) N 350, 
214 


oe ia Siebold.& Zucc., Bridal-wreath 
Spiraea (1; TLM) N 1938 


RUBIACEAE 
Cephal ee occidentalis L., Buttonbush (N; 
AEM,R M)N 159, 592 871, 1041, 1626 
Diodia teres Walter, Rough Buttonweed (N; TUM 
UO) N 202, 253,712, 770, 1869, 2260 
Diodia virginiana L. Virginia Buttonweed (N;RLM, 
RUM, TLM, TLO) N 611,689,872, 2194 
Galium sal ee Plant (N; RUM, TLM, TLO) 
N 464, 1 
Galium circaezans ie Woods Bedstraw (N; 
TUW) N 1445, 2662 
Galium scone Bigel., Bluntleaf Bedstraw (N; 
RUM) N 19 
Galium on Aiton, Hairy Bedstraw (N; TLW, 
TUM, TUO) N 664, 707, 1508, 1754, 2021, 2135, 
553 


Galium tinctorium (L.) Sco | (1; TLM, 
Hee TUM, TUO) N 417, 545, 1264, 1533, 2527, 
255 


ae virgatum Nutt., Southwest Bedstraw (N; 
TUO) N 1683 
Hedyotis nigricans (Lam.) Fosb., Prairie Bluets (N; 


Houstonia micrantha (Shinners) Terrell, White 
Bluets (N; TLM, TUO) N 1231, 2471, 2498 
Houstonia pusilla Schopf., Small Bluets (N; TLW, 

1221,1277 
Houstonia rosea (Raf.) Terrell, Rose Bluets 


— 


N; TLM) 
N 338, 2472 

Mitchella repens L., Partridge-berry (N; TLW) N 
753, 2078 

Oldenlandia boscil (DC.) Chapm. (N; RLM, RUM) 
74, 2036, 


Olden| andia uni ae L.(N; RLM) N 2339 
sis L., Field-madder (I; TLM, TUOQ) 


N 333, 1224 
Spermacoce glabra Michx., Smooth Buttonweed 
(N; RLM, RUM, TLO) N 195, 283, 1917, 2340 


NEILL AND WILSON 


WAU TOUIN LUUINE I, TEXAS 


RUTACEAE 
Poncirus trifoliolata (L.) Raf. Bitter Orange (|;TLW) 


thoxyl lava-herculis L., Hercules-club (N; 
TUM, TUO) N 262, 641, 724, 1459 


ACEAE 
Populus deltoides Marsh.,Eastern Cottonwood (N; 
TUM) N 1460 


Salix nigra Marsh., se Willow (N; RLM, RUM, 
TLO) N 459, 1483, 1736 


SAPINDACEAE 
Cardiospermum Hpieacer| L., Balloonvine 
RLM, TLO) N 196,835, 9 
— swan L. var. nee (Hook. & 
nson, See Soapberry (N; RLM, 
ae N 1011, 2117,2355 


SAPOTACEAE 
Sideroxylon lanuginosum Michx. ssp. albicans 
(Sarg.) Kartesz & Gandhi, Gum Bumelia (N; 
TUOQ) N 767, 2138 
Sideroxylon lanuginosum  Michx. — ssp. 
oblongifolium (Nutt. T.D.Penn.,Gum Bumelia 
(N; TLW, TUM) N 176, 1333, 1844, 2105 


SAURURACEAE 
Saururus cernuus L., Lizard-tail (N; RLM) N 124 
2000 


— 


SAXIFRAGACEAE 

Lepuropetalon spathulatum (Muhl.) Ell, Petite- 
plant (N; TLO) N 2482, 25171 

Penthorum sedoides L., Ditch-Stonecrop (N;RUM, 
TLM) N 222, 2187, 2411 

Saxifraga texana Buckl., Texas Saxifrage (N; TLO, 
TUO) N 2465, 2480 


SCROPHULARIACEAE 
Agalinis fasciculata (Ell.) Raf., Beach Gerardia (N; 
UO) N 791 
ile Use ae (Nutt.) Small ex Britton, Prai- 
rardia (N; TLO, TUO) N 904, 958, 2324 
bea inis purpurea (L.) Penn., Purple Gerardia (N) 
Jones & Jones 433 (TAES) 

Aureolaria grandiflora (Benth.) Penn. var. grandi- 
flora, Bigflower Oakleech (N; TLM) N 2665 
Bacopa monnieri (L.) Penn., Coastal Water-hys- 

sop (N; RUM) N 723 
Bacopa rotundifolia (Michx.) Wettst., Disc Water- 
hyssop (N; RLM) N 2142 


1111 


Castilleja indivisa Engelm., Texas Indian Paint- 
brush (N;TUQ) N 387, 1317 

Gratiola flava Leavenw., Golden Hedge-hyssop 
(N; TLM) N 2466 


Gratiola neglecta Torr., Yellowseed Hedge-hyssop 
(N:RLM,RUM, TLM, TUQ) N 544,826, 1513, 2534 

Gratiola virginiana L., Virginia Hedge-hyssop (N; 

_TLW, TUO) N 445, 1279 2535, 2548 

Leucospora multifida (Michx.) Nutt., Narrowleaf 
Conobea (N; RLM) N 891 

Linaria canadensis (L.) Dum. var. canadensis, 
Oldfield Toadflax (N;TLM) N 1235 

Linaria canadensis (L.) Dum. var. (Scheele.) 
Penn., Texas Toadflax (N; TUM, TUQ) N 438, 
1530 

Lindernia dubia (L.) Penn. var. anagallidea 
(Michx.) Cooperrider, Clasping False Pimper- 
nel (N; RLM, RUM, TLW) N 1932, 1972, 2736 

Mecardonia acuminata (Walter) le Sawtooth 
Water-hyssop (N;TLM) N 238 

Mecardonia procumbens (P. Mill) oat Prostrate 
Water-hyssop (N; TLW) N 2549 

Penstemon australis Small ssp. laxiflorus (Penn.) 
Be outhern Beardtongue (N; TUM) N 

179, Py 65 

Verbascum thapsus L.,Common Mullein (1; TUO) 
N 1906 

Veronica agrestis L., Wayside Speedwell (1; TLO) N 
1192 

Veronica arvensis L.,Common Speedwell (1; TLM) 


we 


Veronica peregrina L. ssp. peregrina, Purslane 
Speedwell (N; TLM, TLW) i" 1233, 1281 

Veronica peregrina L. ssp. xalapensis (Kunth. in 
H.B.K.) Penn., Purslane Speedwell (N; TLW) N 
450 


SIMAROUBACEAE 
Ailanthus altissima (Mill) Swingle, Tree of Heaven 
(Il; TUO) N 790 


SOLANACEAE 

Datura wrightii Regel., Jimsonweed (N; TUM, 
TUO) N 2175, 2300 

Physalis cinerascens (Dun.) A.S. Hitchc. var. 
cinerascens, Beach Groundcherry (N; TUO) N 
274, 903, 1061 

oe heterophylla Nees. var. heterophylla, 

my Groundcherry (N; TUM, TUQ) N 532, 

ee 


a 


1112 


Physalis pubescens L. var. pubescens, Downy 
Groundcherry (N; TLM, TUQ) N 1403, 2192 
Physalis pubescens L. var. integrifolia (Dun.) Wa- 

hae Pe Groundcherry (N; RLM, RUM) 


= 


So an Suen a False Jerusalem- 
cherry (1; TUW) N 17 055 

Solanum carolinense eae Horse-nettle (N; 
TLM, TLO, TLW, TUO) N 606, 930, 1673, 1954, 
2195 

Sol di Raf., Western Horse-nettle 
(N; TUO) N 1618, 2149 

Solanum elaeagnifolium Silverleaf Horse- 

nettle (N;TLM, mae 


lantim rif trim 


ar. Cm (Dun.) 
A. Gray, Cherry ee (1; TUO) N 2698 

Solanum ptycanthum Dun. ex DC., American 
Nightshade (N;RUM, TLO,TUM, TUW) N 7955, 
1760, 1914, 2176 

Chl trot 
2178 

Solanum triquetrum Cav., Texas Nightshade (N; 
TUW) N 701 


Dun., Buffalo-bur (N; TUO) N 


SPHENOCLEACEAE 
Sphenoclea zeylanica Gaertn., Chicken-spike (I: 
RLM) N 214] 


TILIACEAE 

Tiliaamericana L.var.americana, American Bass- 
wood (N; RLM, RUM) N 7592, 1646 

Tilia americana L. var. caroliniana, Carolina Bass- 
wood (N;TLM, TLW) N 152,627, 2080 


ULMACEAE 
Celtis laevigata Willd., Sugar Hackberry (N; TLM, 
TUM) N 369, 465, 755 
Planera aquatica (Walter) J.F. Gmel., Water Elm 
(N; RLM) N 443, 1645 
a ai oe Elm (N; TUM, TUW) 
1,378, 1209 


ieee americana L., American Elm (N;TLM, TLW, 
TUO) N 332, 349, 2092, 2523 
Ulmus Hea Nutt., Cedar Elm (N; RLM, TLW, 
TUM, TUO) N 898, 929, 2343, 2375 


URTICACEA 
gassiies tel .) Sw., False Nettle (N;RLM, 
RUM) N 870, 2091 
Parietaria pensylvanica Muhl.ex Willd. var. obtusa 
(Ryd Small) Shinners, Pellitory (N; TUO) 
N 1406 


BRIT.ORG/SIDA 19(4) 


Parietaria pensylvanica Muhl. ex Willd. var. 
Sees aly el Pellitory (N; RLM, 
RU W) N 57, 440, 1253, 2504 

Urtica eats ae ou leaf Nettle 
(N; RLM, TLW) N 342, 439 


VALERIANACEAE 

alors florifera Shinners, Cornsalad (N; TUCO) 
N 156 

ee radiata (L.) Dufr., Cornsalad (N; TLM, 
TLO, TLW, TUM, TUO) N 364, 416, 461, 1269, 
1303 


VERBENACEAE 

Callicarpa americana L., American Beautyberry 
(N; TLM, TUM) N 156, 920 

Lantana camara LWestindiaa Lantana (I; TUM) 


Lantana urticoides Hayek, Lantana (N; TUM) N 
2152 


Phyla lanceolata (Michx.) Greene, Lanceleaf 
Frogfruit (N; RLM, RUM, TLO) N 128, 687, 966, 


1820, 1975 

ae nodiflora (L.) Greene var. nodiflora, Com- 

rogfruit (N;TLM, TUM) N 137, 2273 

ee pani Nutt., Dakota Vervain (N) 
Huss 91 (TAES) (Glandularia bipinnatifida 
ives Nutt.) 

Verbena brasiliensis Vell., Brazilian Vervain (I; TLO, 
TUO) N85, 1213, 1635, 1819 1976 

Verbena canadensis (L.) Britton, Rose Vervain (N; 
TUO) N 1404 (Glandularia canadensis (L.) 


Nutt.) 

Verbena officinalis L. ssp. halei (Small.) Barber, 
Slender Vervain (N;TLO, TUO) N 74, 925, 1493, 
2126 


Verbena scabra Vahl, Harsh Vervain (N; RUM) N 
222) 

Verbena stricta Vent., Hoary Vervain (N) 
33014 (TEX) 


Verbena urticifolia L. var. urticifolia, White Vervain 
(N;TLM) N 1940 


Albers 


Verbena xutha Lehm., Gulf Vervain (N; TLO, TUO) 
N 198, 2112, 2619 


VIOLACEAE 

Viola bi ee bee, Field Pansy (N; TLM, TUO) N 
ie ee Li i (N; TLW,TUM) N 1204, 2468 

Viola sororia Willd. var. missouriensis (Greene) 
McKinney, Missouri Violet (N; RUM, TLM, TUM 
N 337, 396, 1207 


Kare 


NEILL AND WILSON TEXAS 


Viola sororia Willd. var. sororia, Sister Violet (N; 


VISCACEAE 
Phoradendron tomentosum (DC) oT ex A, 
Gray, Mistletoe (N;PAR) N 7715, 1136,1141, 1205 


VITACEAE 

Ampelopsis arborea (L.) Koehne., Peppervine (N; 
RUM, TLM, TUM) N 88, 633, 748 

Ampelopsis cordata co 
(N; RLM, RUM) N 1947, 

Cissus incisa (Nutt.) Des eck Ivy Treebine (N; 
RLM, Ten N 749,919, 1857 


P, th rinarictolia (L.) Pla 


sais 
Creeper (N:TLM,TLW,TUM) N 155, 1111, 1298, 
] 


Vitis aestivalis Michx. var. lincecumii (Buckl.) 
Munson, Summer Grape (N; TUM) N 1777 
Vitis cinerea (Engelm. in A. Gray) Engelm. ex 

Millard., Sweet Grape (N; RUM, TLW, TUM) NV 
231, 1489, 2649 
Vitis mustangensis Buckl. deo Grape (N;TLM, 
TLO, TUM) N 175,625,217 
Vitis palmata Vahl, Catbird aaa 


=> 


N; RLM, TLM) 


Vitis riparia Michx., Frost Grape (N; RLM) N 1978 
Vitis rotundifo ia Michx. var. rotundifolia, Musca- 
e Grape (N; RLM, TUM, TUW) N 230, 1335, 
ee 790 


MAGNOLIOPHYTA: LILIOPSIDA 
AGAVACEAE 
Manfreda virginica (L.) Rose, Rattlesnake Master 
(N; TUM) N 2107 
Yucca arkansana Trel., Arkansas Yucca (N; TUO) 
N 1616 


= 


yucca louisianensis Trel., Louisiana Yucca (N; TUO) 
N 2565 


= 


Yucca treculeana Carr., Spanish Dagger (N;TUM 
N 213 


ALISMATACEAE 

Echinodorus cordifolius (L.) Griseb., Burhead (N; 
RLM, TLW) N 1894, 1 

Sagittaria graminea Michx. ssp. gra 

ead (N; AEM, RLM) N 424, 688, 1479 
Sait eats Engelm.,Delta Arrowhead 
AEM, RUM) N 224, 1985 

ARACEAE 

Arisaema dracontium (L.) Schott., Green Dragon 
(N; TLM, TLW, TUW) N 142, 1414, 1572 


1113 


ARECACEAE 

Sabal minor (Jacq.) Pers., Small Palmetto (N;RLM, 
TLW) N 110,677 

BROMELIACEAE 

Tillandsia recurvata L., Ball-moss (N; EPI 
2170 


— 


N 282, 


Tillandsia usneoides (L.) L., Spanish-moss (N; EPI) 
86, 807, 1034, 1496 


CANNACEAE 
Canna x generalis LH. Bailey, Canna (I; TLW) N 676 


COMMELINACEAE 

Commelina diffusa Burm. f. var. diffusa, Spread- 
ing Dayflower (N;TLO, TUW) N 970, 2663 

Commelina erecta L. var. angustifolia (Michx.) 
Fern., Narrowleaf Dayflower (N; TUM, TUO) N 
256, 172 

Commelina erecta L. var. erecta, Erect Dayflower 
(N;TLO) N 971 

ails ee L., Virginia Dayflower (N; 
RLM, TLW, TUO) N 853, 1594, 1980, 2344 

mee hirsutiflora Bush, Hairyflower Spi- 
derwort (N;TLM, TLW) N 1603, 2486 

eset ohiensis Raf, Ohio Spiderwort (N; 

RLM, TLO, TLW, TUW) N 355, 1435, 1654, 1832 

Tradescantia reverchonii Bush, Reverschon Spi- 
derwort (N) MacQueen 58 (TAMU) 

Tradescantia subacaulis Bush., Stemless Spider- 

ort (N; TLM, TUM) N 413, 2586 

CYPERACEAE 

Carex albicans Willd. ex Spreng var. australis 
(Bailey) J. Rettig, Sedge (N; TLW ) N 2517 

Carex arkansana Bailey, Arkansas Sedge (NN; TUO) 
N 2544 


23 


Carex austrina (Small) K. Mack., Southern Sedge 
(N; RUM) N 1966 


Carex bulbostylis Dew., Sedge (N; TUW) N 1437 
Carex bushii Mack., Bush Sedge (N; TUO) N 2629 
Carex caroliniana Schwein., Carolina Sedge (N; 


| Pe ip’ H CI | c | N; 
RUM, TLO, TLW,TUO) N 1791, 1286, 1770, 2541 
Carex complanata T.&H., Sedge (N; RLM) N 407, 


Carex corrugata Fern., Corrugated Sedge (N; 
RUM) N 1771 

Carex crus-corvi Schuttlw. ex Kuntze., Crowfoot 
Sedge (N; RLM, TLM) N 428, 1307, 1344 

Carex emoryi Dew., Emory Sedge (N;RUM) N 2516 


1114 


Carex flaccosperma K. Mack., Thinfruit ee (N; 
RLM, TLM /TLW) N 549 1283, 1342, 

Carex frankii Kunth., Franks Sedge ie 
N 1951, 1993 

Carex hyalina Boott., Whitesheath Sedge (N; 


Carex hyalinolepis Steud., Thinscale Sedge (N; 
9?, 1562 
Carex leavenworthii Dew., Narrowleaf Sedge (N; 
RLM) N 408 
Carex longii Mack., Longs Sedge (N;TLW) N 1569 
Carex louisianica Bailey, Louisiana Sedge (N:TLW) 


N 1285 

Carex lupulina Muhl., Hop Sedge (N; RLM, RUM, 
TLW) N 848, 1742, 1991, 2600 

Carex oxylepis T.& H., Sharpscale Sedge (N; TLW) 


N 1287 

Carex retroflexa Muhl., Reflexed Sedge (N; RLM) 
N 2590 

Carex tetrastachya G. Scheele, Sedge (N; RLM, 
TUM) N 1880, 2591 


Carex texensis (Torr.) L.H. Bailey, Texas Sedge (N; 
TLW) N 1291 
Carex triangularis Boeckl., Sedge (N; RLM) N 409 
arex vulpinoidea Michx., Fox Sedge (N; RLM) N 
1343 


Cyperus acuminatus |. & H., Taperleaf Flatsedge 
(N; AEM) N 179, 713 

Cyperus articulatus L., Jointed Flatsedge (N;TLO) 
N 609 


Cyperus croceus Vahl, Baldwin Flatsedge (N;TUM, 
TUO) N 780, 2703 

Cyperus echinatus (L.) A.Wood, Globe Flatsedge 
(N; TUM) N 2434 

Cyperus erythrorhizos Muhl., Redroot Flatsedge 
(N; RLM) N 812, 2349 

Cyperus esculentus L. var. esculentus, Yellow Nut- 
sedge (N; TUM) N 2457 

Cyperus haspan L., Sheathed Flatsedge (N; RUM) 
N714 


Cyperus odoratus L. var. squarrosus Jones, Wi 
& Carter, Fragrant Flatsedge (N) Jones & Jones 
404 (TAES) 

Cyperus polystachos Rottb., Flatsedge (N;RLM) N 
809 


Cyperus pseudovegetus Steud., Flatsedge (N;TLM, 
71,1095 
Cyperus reflexus Vahl, Bentawn Flatsedge (N; 
TUM) N 2458 


BRIT.ORG/SIDA 19(4) 


Cyperus retrorsus A. Chapm., Flatsedge (N; TUM, 
ae N 258, 6 


Cyperus retroflexus Buckl. var. retroflexus, 
Oneflower Flatsedge (N) Duncan s.n. (TAES), 
MacQueen 73 (TAM 

Cyperus rotundus L.,Nutgrass (|; RLM, TUO) N 1884, 
2267 


Cyperus strigosus L., False Nutgrass (N; TUO) N 
2134 
Cyperus virens Michx., Green Flatsedge (N; TLO) 


N 1036 

Eleocharis fallax Weath., Spikerush (N) Kessler 
5839 (SHST) 

Eleocharis lanceolata Fern., Lanceleaf Spikerush 
(N) Herrick 120 (TAMU) 

Eleocharis microcarpa Torr., Smallseed Spikerush 
(N; RLM) N 7347 


(N; AEM, RUM, TLM) N 423, 1305, 1565, 1986 

Eleocharis obtusa (Willd.) Shult., Blunt Spikerush 
(N; AEM) N 1476 

Eleocharis palustris (L.) Romer & J.A. Shultes, 
Largespike Spikerush (N; RLM) N 1887 

Eleocharis quadrangulata (Michx.) R.&S., 
Squarestem Spikerush (N; AEM) N 685 

Fimbristylis autumnalis (L.) R.&S., Slender Fimbry 
(N; RLM, RUM) N 252 811 

Fimbristylis vahlii (Lam.) Link., Vahl Fimbry (N; 
RLM) N 236, 808, 2353 


Fuirena squarrosa Michx., Hairy Umbrellagrass (N; 


Isolepis carinata Hook. & Arn. ex Torr., Small Bul- 
rush (N; RLM, TLM, TUO) N 541, 1340, 1485 
Isolepis molesta (M.C. Johnst.) $.G. Smith, Small 
Bulrush (N; TLM) N 542, 7 

Kyllinga Cee Vahl, Fragrant Flatsedge (N; RUM, 
TLO) N 574, 232 

wee caduca Ell, Anglestem Beakrush 
(N; RLM) N 240 

Rhynchospora corniculata (Lam.) A.Gray, Horned 
Beakrush (N; AEM) N 1946 

Rhynchospora globularis (Chapm.) Small var. 
recognita Gale, Beakrush (N; RLM) N 125 

Rhynchospora glomerata (L.) Vahl, Cluster 
Beakrush (N; TLW) N 1451, 1547 

Rhynchospora Gane Torr., Tall Beakrush 
(N; AEM, RUM) N 775, 

Schoenoplectus ca roe € ae J. Sojak., 
Giant Bulrush (N; AEM) NV 


NEILL AND WILSON , TEXAS 


Scirpus cyperinus (L.) Kunth., Woolgrass Bulrush 
(N) Kessler 5849, 5866 (SHST) 

Scleria ciliata Michx., Fringed Nutrush (N; RUM) 
N 1625 

Scleria oligantha Michx., Littlehead Nutrush (N; 


DIOSCOREACEAE 
sass villosa L., le Yam (N;TLW, TUW) NV 
1658, 1907, 2 


IRIDACEAE 

Alophia drummondii (Graham) Foster, Purple 
Pleatleaf (N; TLO) N 2678 

Gladiolus italicus Mill, Corn-flag (1; TUO) N 2741 

Iris hexagona Walter var. flexicaulis (Small) Foster, 
Dixie Iris (N;TLM, TUQ) N 1311, 2481, 2574 

Iris pallida Lam., Bearded Iris (1; TUM) N 768 

Sisyrinchium albidum Raf.,White Blue-eyed Grass 
(N; TUM) N 556 

Sisyrinchium exile Bickn., Yellow Blue-eyed Grass 
(N; TLW, TUO, TUW) N 1268, 1433, 1600 

Sisyrinchium langloisii Greene, Pale Blue-eyed 
Grass (N; TUO) N 1602 

Sisyrinchium minus Engelm. cn Gray, Least Blue- 
eyed Grass (N; TUO) N 160 

Sisyrinchium A ipoun a eee Blue-eyed 
Grass (N; TLO, TUO) N 468, 1454 

Sisyrinchium rosulatum Bickn., Spreading Blue- 
eyed Grass (I; TLO, TUM, TUO) N 567, 1703, 2664 


JUNCACEAE 
Juncus acuminatus Michx., Knotleaf Rush (N; 
TUW) N 1440 


a 


Juncus acuminatus Michx. f. sphaerocephalus 
rm., Knotleaf Rush (N; RUM) N 1637 
Juncus brachycarpus Engelm.,Whiteroot Rush (N; 

RUM) N 1728 
Juncus bufonius L., Toad Rush (N) Gould 8560 
(TAES) 


— 


Juncus capitatus Weigel. Capped Rush (N; TUO 


Juncus debilis A.Gray, Weak Rush (N; RUM) N 19817 

Juncus dichotomous Ell., Forked Rush (N; TLM, 
TUO) N 1217, 1308 

Juncus — eels Slimpod Rush (N; 
AEM, TLM) N 

Juncus ice, ae Rush (N; RLM, TUW 
N 


3; 


VS 


Juncus ae L. var. solutus Fern. & Weig., Com- 
mon Rush (N; AEM, ASU, RLM) N 247, 432, 
114, 1560 


1115 


— 


Juncus interior Wieg., Inland Rush (N; RUM, TLW) 
N 1962, 1992 

Juncus marginatus Rostk.,Grassleaf Rush (N;TLM, 
TLO, TUO) N 26, 575, 2643 

Juncus repens Michx., Creeping Rush (N; ASU) NV 
1971 


el Lam IN 1 me) kh (N-RLIM) 


N 1103 
Juncus tenuis Willd., Slender Rush (N; RUM) N 58 
Juncus validus Coville. var. validus, Roundhead 

Rush (N; AEM, RLM, TLW, TUM) N 7122, 1270, 

1701, 1780, 1944 
Luzula bulbosa (Wood.) Rydb., Bulb Woodrush (N; 

TLM, TLO) N 2479, 2519 
aria) 

mna min Phil., Duckweed (N; AFL) N 1042 
Sree ie cn (L.) Schleid., Duck-meat (N; 


we nae (Meyer) Thomps., Duck-meat 
(I; AFL) N 1043, 2489 

Wolffia columbiana Karst.,Water-meal (N; AFL) N 
2490 


LILIACEA 
Allium conan L. var. canadense, Wild Onion 

(N; TLM, TLO, TUW) N 460, 1434, 1589, 2579 

Allium ee L. var. mobilense (Regel.) M 
Ownbey, Wild Onion (N; TLM, TUM, TUO) N 
404, 502, 1725, 2576 

Asparagus officinalis L., Asparagus (1; UO) N 1877 

Cooperia drummondii Herb., Cebolleta (N; TUO) 
N 271 


Cooperia pedunculata Herb., Giant Rainlily (N; 
2173 
Crinum americanum L., Swamp-lily (N; TUO) N 


1639 

Erythronium albidum Nutt., White Fawn-lily (N; 
TLW) N 2510 

Habranthus tubispathus (L'Her) Traub, Copper 
Rainlily (l; TUO) N 206, 2172 

Hemerocallis fulva L., Daylily (I; TW, TUM) N 675, 


766 

Hippeastrum bifidum (Herb.) Bak., Oxblood Lily 
(I; TUM) N 2244 

Hymenocallis caroliniana (L.) Herb., Carolina 
Spiderlily (N; AEM, TLW) N 1641, 2538 

Hymenocallis eulae Shinners, Late Spiderlily (N; 
TUM, TUO) N ee) oe 2377 


Shinners, Spiderlily (N; 


AEM) N 1313 


1116 


Hypoxts hirsuta (L.) Cov., Hairy Goldstar-grass (N; 
TUM) N 415, 1461 


ce spicata Lour.,, Creeping Lilyturf (1; TLW) N 


hrs joni L., Al (I; TUO) N 2525 
Nothoscordum bivalve (L.) Britton, Crow-poison 
(N; ar TUO) N 334, 910 
ii A.Gray, Nutall Deathcamas (N; 


~ TUM) N 1325 


MARANTACEAE 

Thalia dealbata Roscoe, Powdery Thalia (N; AEM) 
N 132, 2572 

NAJADACEAE 

Najas ees (Spreng.) Magnus, South- 
ern Naiad (N; AFL) N 682 


ORCHIDACEAE 
Corallorhiza wisteriana Conrad.,Wister Coralroot 
(N; TUW) N 1456 
Spiranthes cernua (L.) Rich var. cernua, Nodding 
Ladies’ Tresses (N; TLM, TUM, TUO) N 1000, 
1024, 1063, 1117, 2422 
Spiranthes gracilis (Bigel.) Beck var. gracilis, Slen- 
r Ladies’ Tresses (N; TUO) N 1005, 1062 
Spiranthes parksii Correll, Navasota Ladies’Tresses 
(N) Nixon 15571 (ASTC), Parker 1858 (BRIT) 
acs praecox (Walter) Wats., oe leaved 
Ladies’ Tresses (N; TUW) N 167 
Spiranthes tuberosa Raf. Gray's oe Tresses (N; 
M) N 2670 
Spiranthes vernalis Engelm. & A. Gray, Spring La- 
dies’ Tresses (N; RUM, TLO, TUO) N 604, 608, 
1804, 2606 


POACEAE 
Agrostis elliottiana Schult, Elliot Bentgrass (N; 
TUM) N 1354 

rele niet (Walter) B.S.P.,Winter Bentgrass 
(N; W,TUO) N 55, 1410, 1526, 1762 

Aira ee rae Hairgrass (|; UO) N 1535, 1687 

Alopecurus sally Walter, Carolina Foxtail 
(N;TLM, TLW, TLO) N 64, 1409, 1558 

Andropogon oe (Walter) B.S.P., Bushy 
Bluestem (N; RLM) N 976 

Andropogon ternarius Michx., Splitbeard 
Bluestem (N; TUM, TUO) N 1009, 1023, 1092, 
1109 

Andropogon virginicus L., Broomsedge Bluestem 
(N; RUM, TUO) N 2035, 2464 


BRIT.ORG/SIDA 19(4) 


Aristida desmantha Trin. & Rupr., Curly Threeawn 
(N; TUO) N 1008 

Aristida longespica Poir. var.geniculata (Raf.) Fern, 

ins Threeawn (N;TLM) N 2444 
Aristida — Poir.var. longespica, Slimspike 
Threeawn (N; TUO) N 1075 

Aristida slbanthe els bis Threeawn (N; 
TUM, TUO) N 914 

Arundinaria ia gigantea ee ne Giant Cane 
(N; W) N 833, 1 8 

Arundo nee L., Giant Reed (|: TUO) N 880, 881 

Avena fatua L. var. fatua, Wild Oats (I TLO, TUO) N 


455, 1393 
Avena fatua L. var. sativa (L.) Hausskn., Common 
ts (I; TLO) N 167 
Bothriochloa ischaemum (L.) Keng. var.songarica 
(Fisch. & Mey.) Cel. & Harl., King Ranch 
Bluestem (I:Tl JM, TUO) N 789, 1108, 1525, 2276 
Bothriochloa laguriodes (DC.) Herter ssp. 
torreyana (Steud.) Allred & Gould, Silver 
Bluestem (N; TUM, TUO) N 7105, 1007, 1093 
ete longipaniculata (Gould) Allred & 
ould, oe, a Bluestem (N; TUM, 
ao N97 
Bouteloua eu area (Michx.) Torr. var. 
curtipendula, Sideoats Grama (N;TUO) N 696 
Briza minor L., Little Quaking Grass (I; TLO, TLW, 
N517, 1332, 1449 
Bromus catharticus M.A.Vahl, Rescuegrass (I; RUM 
M,TUO) N 516, 1352, 1394, 2507 
Bromus japonicus Thunb., Japanese Brome (I: 
TUO) N 1726, 2005 
Bromus pubescens Muhl. ex Willd., Brome (N) 
Nixon 4712 (ASTC) 
Bromus secalinus L., Rye Brome (|; TLM, TUO) N 


6/7, 1677, 1763 

Buchloe dactyloides (Nutt.) Engelm., Buffalograss 
(N; TLO) N 579 1889 

Cenchrus spinifex A. Cav., a (N; TUM, TUO) 
N 249, 737, 2016, 2270, 239 

Chasmanthium latifolium ioe Yates, Broad- 
leaf Woodoats (N;RLM,RUM) N 112,795, 1765 

Chasmanthium laxum (L.) Yates var. laxum, 

oodoats (N; RLM, TLW) N 228, 2400 

Chasmanthium laxum (L.) Bae var. sessiliflorum 
(Poir.) Wipff & S.D. , Narrowleaf 
Woodoats Aaa pee 2658 

Chloris canterai Arech., Windmillgrass (|; TUO) N 
2124 


NEILL AND WILSON, 


TEXAS 


Chloris verticillata Nutt., Tumble Windmillgrass (N; 
TUM) N 2259 
Chloris virgata Sw., Showy Windmillgrass (N;TLM) 


15 
Chloris x eileen Mueller, Shortspike 
Windmillgrass (N; TUM, TUO) N 1105, 2122 
Cynodon dactylon (L.) pare: Bermuda Grass (I; 
TUM, TUO) N 973, 1033, 1327 
Dactyloctenium aegyptium (L.) Willd., Durban 
Se done ( RLM, TUO) N 849, 912, 2310 
Dichantheliu (Poir.) Gould & Clark var. 
ees Rosettegrass (N; TLO, TUQ) N 996, 
8 


197 
Dichanthelium acuminatum (Sw.) Gould & Clark 
var. acuminatum, Woolly Rosettegrass (N; 
RUM, TUO, TUW) N 71, 716, 1422, 1766 
Dichanthelium acuminatum (Sw,) Gould & Clark 
var. lindheimeri (Nash) Gould, Lindheimer 
Rosettegrass (N;RLM, RUM, TUW) N 746, 875, 


2708 
Dichanthelium clandestinum (L.) Gould, 
eertongue Rosettegrass (N; TUW) N 2709 
Dichanthelium commutatum (Schult.) Gould, 
Variable Rosettegrass (N; TLM, TLW, TUW) N 
1289, 1429 1524, 2077, 2518 
Dichanthelium dichotomum (L.) Gould var. 
um, Rosettegrass (N; TUM) N 539 
Dichanthelium oligosanthes (Schult.) Gould var. 
oligosanthes, Rosettegrass (N; TUM) N 553 
es (Schult.) Gould var. 
scribnerianum one Gould, Scribners 
Rosettegrass (N; TUM, TUO, TUW) N 25, 1850, 
2/07 


dichotom 


Dichanthelium ravenelii (Lamson-Scribn.) Gould, 
Rosettegrass (N;TLM) N 2673 

Dichanthelium scoparium (Lam.) Gould, Velvet 
Rosettegrass (N; RUM, TUM, TUO) N 227, 508, 
1773, 1778, 1903 

Dichanthelium sphaerocarpon (Ell.) Gould & Clark 
var. spl | R | 1R 
(N;: TLW) N 1294, 1505 

Digitaria cognata (Schult. Pilger ssp. cognata, Fall 
Witchgrass (N; TUO) N 1057, 1617 

Digitaria sanguinalis (L.) Scop., Hairy Crabgrass 
(Il: TLO, TUM, TUO) N 2019, 2123, 2212, 2261 

Digitaria villosa (Walter) Pers., Shaggy Crabgrass 
N; TUM) N 782 

Digitaria violascens Link., Violet Crabgrass (1) Wells 
36 (TAES) 


1117 


Echinochloa colonum (L.) Link. Junglerice (1; AEM) 


N 2460 

Echinochloa crusgallii (L.) Beauv. var. crusgallil 
sles a (|; AEM,RLM, RUM) N 876, 2032, 
2082, 2 

Echi tees muricata (Beauv.) Fern., Barnyard 
Grass NpNewinan 18 (1. ES) 


L/ h) Hell | 
cE Uist) 


(N; AEM, RLM, TUO) N 290, 22] a 2347 
Fleusine indica (L.) Gaertn (1; TUO) N 


Elymus canadensis L. var. canadensis, Canada 
Wildrye (N; TUM, ee N 107,618, 1733 

ar virginicus L.,Virginia Wildrye (N;TLO, TL 

TUO) N 1840, 1920, 2018 
Eragrostis curtipedicillata Buckl., 
rass (N; TUM) N 2280 

Eragrostis hirsuta (Michx.) Nees, Bigtop Lovegrass 

(N; TUM) N 2186 


Gummy 


Eragrostis hypnoides (Lam.) B.S.P., Teal Lovegrass 
(N; RLM) N 235 
Eragrostis intermedia A.S. Hitchc. var. intermedia, 
Plains Lovegrass (N; TUM) N 1099, 2237 
eae reptans (Michx.) Nees, Creeping 
ass (N; RLM) N 887, 1939, 2354, 2592 
crags secndion a hae oxylepis (Torr.) 
D.Koch,R N: TUM, TUO) N 246, 


a 2263, 2305, 24 433 
Eragrostis spectabilis (Pursh.) Steud., Purple 
Lovegrass (N; RLM, TUM) N 771, 2352, 2438 
Friochloa sericea (Scheele) Munro ex Vasey, Texas 
ss (N) Nixon 4737 (ASTC 
Eustachys retusa (Lag.) Kunth.,Windmillgrass (N; 
TUO) N 196 


Festuca subverticillata (Pers.) E.B. Alexeev, Nod- 
ding Fescue (N;TUOQ) N 1585 

Glyceria septentrionalis Hitchc. var. septentrionalis, 
Eastern Mannagrass (N; AEM) N 1945 

Gymnopogon ambiguus (Michx.) B.S.P, Bearded 
Skeletongrass (N) Wells 45 (TAES) 

Hordeum leporinum Link., Hare Barley (|; TLO) N 
1442 


Hordeum pusillum Nutt., Little Barley (N; TUO) N 


— 


1396 
Leersia lenticularis Michx.,Catchfly Grass (N;RLM 
N 866, 978, 2348 


Leersia virginica Willd., Whitegrass (N; RLM, RUM) 
N817, 2674 
Lolium nne L., Perennial Ryegrass (N; TLM, 


Mm per 
TLO, Tu) N 454, 1237, 1395 


1118 


Melica mutica Walter, Twoflower Melic (N; RUM, 
TLW) N 148 1259 

Muhlenbergia capillaris (Lam.) Trin., Hairyawn 

hly (N; TUM) N 1098, 2440 

Muhlenbergia schreberi Gmel., Nimblewill (N) 
Rich & Briley 1713 (ASTC) 

Nassella fees (Trin. & Rupr.) Pohl, Texas 

ss (N; TLO, TLW) N 453, 1490 

bie Fe (L.) ee Basketgrass (I; 
TLM, TLW) N 1593, 2303, 

Panicum anceps Michx., aa Panicum (N; 

M) N 2235, 2441 

Panicum brachyanthum Steud., Pimple Panicum 
(N;TLM, TUM) N 2252 2302 

Panicum dichotomiflorum Michx., Fall Panicum 
(N; RLM, TLW) N 888, 2459 

Panicum diffusum Sw., Spreading Panicum (N) 
Waller & Lonard 1973 (TAES) 

Panicum gymnocarpon Ell, Savannah Panicum 
(N; RLM) N 850 

Panicum hallii Vasey var. hallit, 
Wells 65 (TAES) 

Panicum hians Ell, Gaping Panicum (N;RLM,TLM, 
TUO) N 120, 543, 818, 1055 

Panicum rigidulum Nees, Redtop Panicum (N; 
RLM, TLW) N 297,819, 890, 1892, 1994 

Panicum virgatum L., Switchgrass (N; RLM, TUO 
N 889, 1089, 2712 


Halls Panicum (N) 


<a. 


Paspalidium geminatum (Forssk.) Stapf. var. 
geminatum, Paspalidium (N; RUM) N 2030 

Paspalum dilatatum Poir., Dallisgrass (I; TUM) N 
17 


Paspalum floridanum Michx. var. glabratum 
ngelm.ex Vasey, Sat Paspalum (N; TUM, 
vo 5, 1032, 243 
rl ae (EIL) on Water Paspalum 
(N; M) N 863, 2166, 2350 (P repens 
Be crates Val. ie. (Ell) Wipff & S.D. Jones 
Paspalum laeve Michx., Field Paspalum (N) New- 
man 2 (TAES) 
Paspalum notatum Flugge var. notatum, 
Bahiagrass (|; TUM, TUO) N 245, 1849 
Paspalum notatum Flugge var. saurae Parodi, 
Bahiagrass (|; TLM) N 634 
Paspalum plicatulum Michx., Brownseed 
pe Gaal (N; TUM, TUO) N 69, 1950, can 
258, 2 


rh Seen Rupr. ex Fourn., Hairyseed 
Paspalum (N) Nixon 7604 (TAES) 


BRIT.ORG/SIDA 19(4) 


Paspalum setaceum sas var. setaceum, Thin 
Paspalum (N; TUOQ) N 1708 
Paspalum setaceum Mic ri . var, Stramineum 
(Nash.) D. Banks, Thin Paspalum (N; TUO) NV 
48 


18 
Paspalum urvillei eel , Vaseygrass (I;TLO, TUO) 


Phalaris Seth ae aes Carolina Canarygrass 
(N;TLM, TLO, TUM) N 60, 507, 1425 

Piptochaetium avenaceum (L.) Parodi, Blackseed 
Needlegrass (N; RLM, TLM, TLW, TUM) N 555, 
1293, 1345, 1580 

Poa annua L., Annual Bluegrass (I; TLW) N.2739 

Poa autumnalis Muhl. ex Ell, Autumn Bluegrass 
(N; TUO) N 1567 

Poa chapmaniana eal ale Chapman 
Bluegrass (N; TUM) N4 

Polypogon monspeliensis t ) Desf., Rabbitfoot 

pomeogen tl RUM) N 


a 


Spreng., a RUM) N 2345 

een scoparium (Michx.) Nash var. 
divergens (Hack.) Gould, Little Bluestem (N; 
TUM, TUO) N 1058, 2306 

Schizachyrium scoparium (Michx.) Nash var. 
scoparium (C.E.Hubb.) Gould, Little Bluestem 
(N; TUM, TUO) N 993, 2238 

Secale cereale L., Rye (1; TUM) N 103 

Setaria parviflora (Poir.) Kerguel., Bristlegrass 
RLM, TLO, TUM, TUO) N 
2213, 2278 


=> 


N; 
197, 285, 623, 1627, 


Sorghastrum nutans (L.) Nash, Yellow Indiangrass 
(N; TUM, TUO) N 1090, 1106, 2435 

Sorghum halepense (L.) Pers., Johnsongrass (I; 
RUM, TUM, TUO) N 108, 1734, 2033 

Sphenopholis interrupta (Buckl.) Lamson-Scribn., 
Wedgescale (N; TUO) N 1689 

Sphenopholis obtusata (Michx.) Lamson-Scribn. 
var. major (Torr) Erdman, Wedgescale (N; 
TLM, TLW,TUO) N 1255, 1306, 15 

Sphenopholis obtusata (Michx.) Lamson-Scribn. 

tusata, Prairie Wedgescale (N; TUO) N 


= 


pact 


1488 

Sporobolus compositus (Poir.) Merr. var. macer 
(Trin.) Kartesz & Gandhi, Dropseed (N; TUM 
TUO) N 1059, 2439 

Sporobolus coromandelianus (Retz.) Kunth, 
Dropseed (N; TUO) N 2174 

Sporobolus cryptandrus (Torr.) A. Gray, Sand 
Dropseed (N; TUM) N 2279 


NEILL AND WILSON, TEXAS 1119 


Sporobolus indicus (L.) R.Br, Smutgrass (N;TUM, Zea mays L., Corn (I; TLO) N 2207 
TUO) N 1330, 1523, 2462 Zizaniopsis miliacea Oaleste) Doell. & Asch., 
Stenotaphrum secundatum (Walter) Kuntze, St. Southern Wildrice (N; RLM, RUM) N 220, 1890 
Augustine Grass (I; RUM, TLO) N 763, 2641 PONTEDERIACEAE 


Tridens flavus (L) A.S. Hitche. uel chapmanil Pontederia cordata L.,Pickerelweed (N; AEM) N 788 
(Small) Shinners, Chapman Tridens (N; RLM, 


TUM) N 783, 804 POTAMOGETONACEAE 

Tridens flavus (L.) A'S. i var. fe Purpletop een oo (Mart.) Solms, Water Hya- 
N;TUM,TUO) N 2301, 2 Murray 34 (TAMU), Griffith 28 (TAMU) 

Tridens strictus (Nutt) qe longs Tridens paren diversifolius Raf. var. diversifolius, 
(N:TUM,TUO) N 976, 1060, Waterthread Pondweed (N; AFL) N 1983, 

at dactyloides (L.) L., sea Gamagrass 481, 2446 
(N; TLM, TLO, TUO) N 150, 645, 2732 SMILACACEAE 

rca Bevan ee (I, TUM) N 102 Smilax bona-nox L.,Cat-briar (N; RLM, TUM, TUW) 

Wright) R.D.Web N 123, 1053, 1127, 2071, 2455 

Broadleaved Signalgrass (N; TUM, TUO) N Smilax glauca Walter, Saw-briar (N; TLW, TUM) N 
2271, 2372, 2679 1769, 1791 

Urochloa texana (S. Buckl.) R.D. Webster, Texas Smilax glauca Walter var. leucophylla Blake, Saw- 
Signalgrass (N; TUM, TUO) N 915, 1124 briar (N;TUM) N 232 

Vulpia myuros (L.) KC. Gmel. var. myuros, Rattail — Smilax smallii Morong, Green-briar (N;TLW,TUM) 
Sixweeks Grass (I; TLO, TLW) N 1470, 1667 N 340, 560, 1768, 2068 

— octoflora (Walter) Rydb. var. nel ora, — Smilax tamnoides L., Hellfetter (N; RLM, TLW) N 

on Sixweeks Grass (N;TLM, TUM, TUQ) 873 2115. 2404 


ae 1507, 1546, 2587 
Vulpia sciurea (Nutt.) Henr., Squirrel Sixweeks 
Grass (N; TUO) N 2563 


TYPHACEAE 
Typha domingensis Pers., Cattail (N; AEM 
590, 1037, 1628, 1982 


N 97, 


a 


ACKNOWLEDGMENTS 


We thank Stephan Hatch and Dale Kruse (TAES), Monique Reed (TAMU), Stan- 
ley Jones (BRCH), Ted Barkley (BRIT), and Richard Rabeler (MICH) for their 
assistance in the identification of several taxa. Jim Yantis and Baron Rector are 
thanked for their knowledgable advice. Valuable help and companionship in 
the field were given by Toni Wilson, John Janovec, William and Charlotte Neill, 
Ed McWilliams, Carolyn Rose, John McEachran, Eddy Dawson, Dave Cairns, 
and students of Botany 620 and Geography 624 at Texas AWM. Guy Nesom, 
Larry E. Brown, and Barney Lipscomb provided thorough reviews and excel- 
lent advice. This project would not have been a success without the generous 
permission of the landowners and leaseholders of Madison County—too nu- 
merous to mention here—and we thank them for access to their property and 
hospitality. 


REFERENCES 


Atten, H.G. 1974. Woody vegetation of the lower Navasota River watershed. M.S. Thesis, 
Texas A&M University, College Station. 


1120 BRIT.ORG/SIDA 19(4) 


Awvrep, B.W. and H.C. MitcuHett. 1955. Major plant types of Arkansas, Louisiana, Oklahoma, 
and Texas and their relation to climate and soils. Texas J. Sci. 7:1-19. 

Biair, W.F. 1950. The biotic provinces of Texas. Texas J. Sci. 2:93-117. 

Bray, W.L. 1906. Distribution and adaptation of the vegetation of Texas. University of Texas 
Bull. 82. Austin. 

Bryant, V.M. Jr. 1977.A 16,000 year pollen record of vegetational change in central Texas. 
Palynology 1:143-156. 

Corrett, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner 

Cory, V.L.and H.B.Parks. 1937. Catalogue of the flora of the state of Texas. Texas Agric. Exp. 
Sta. Bull. 550. 

Cronauist, A. 1982. Map of the floristic provinces of North America. Brittonia. 34:144-145. 

Cronauist, A. 1988. The evolution and classification of flowering plants, ed.2. The New York 
Botanical Garden, Bronx. 

Diamonp, D.D. 1980. Remnant plant communities of the Fayette Prairie, Texas. M.S. Thesis, 
Texas A&M University, College Station. 

Dicas, G.M. Jr., B.L. Liescome, and R.J.O’Kennon. 1999, Shinners & Mahler's illustrated flora of 
north central Texas. Sida, Bot. Misc. 16. 


FLORA OF Texas Consortium. 2000a. Herbarium specimen browser website: http:// 
www.csdl.tamu.edu/FLORA/ftc/ftchsb.htm. 

Flora OF Texas Consortium. 2000b. Vascular plants endemic to Texas website: http:// 
www.csdl.tamu.edu/FLORA/endemics/endemic1.htm. 

Goutp, FW. 1975a. Texas plants: a checklist and ecological summary. Texas Agric. Exp. Sta. 
Misc. Pub. 585/revised. 

Goutp, FW. 1975b. The grasses of Texas. Texas A&M University Press, College Station. 

Goutp, F.W.,G.O. Horrman, and C.A. RECHENTHIN. 1960.Vegetational areas of Texas. Texas Agric. 
Exp. Sta. Bull. 1070. 

Hatch, S.L., K.N.GaNnbHi, and L.E. Brown. 1990, Checklist of the vascular plants of Texas. Texas 
Agric. Exp. Sta. Misc. Pub, 1655:1-158. 

HicHTower, T.P.1987.Vegetation dynamics of a native haymeadow, a grazed meadow, and 
an old field within the San Antonio Prairie of Texas. M.S. Thesis, Texas A&M University, 
College Station. 

Hipp, L. 2000. Bedias reservoir proposed. Bryan-College Station Eagle, Bryan, TX. 4 Oct 
2000:A1,A3. 

Jones, S.D.s.d. Key to Carex species of Texas. Unpublished manuscript. College Station. 

Jones, S.D., J.K. Wierr, and PM, Montcomery. 1997. Vascular plants of Texas: A comprehensive 
checklist including synonymy, bibliography, and index. University of Texas Press, 
Austin. 

LAUNCHBAUGH, J.L., JR. 1952. Vegetational changes in the San Antonio Prairie caused by 
mowing, grazing, and abandonment. Ph.D. Dissertation, Texas A&M University, College 
Station. 


NEILL AND WILSON, TEXAS 1121 


Levey, K. 1998. Botanist looking for access: student collecting plants for thesis. Bryan-Col- 
lege Station Eagle, Bryan, Texas. 5 May 1998:A9,A12. 

LuNoewt, C.L. (ed). 1942. Flora of Texas. Vol. 3, Pt. 1. University Press, Dallas, Texas. 

LUNbELL, C.L. (ed). 1943. Flora of Texas. Vol.3, Pt. 2. University Press, Dallas, Texas. 

Mapison County Historical Commission. 1984. A history of Madison County, Texas. Taylor Pub- 
lishing Co., Dallas, TX. 

McCates, J.E. 1954. An ecological and range vegetation analysis of the upland sites of the 
southern extension of the Oak-Hickory forest region in Texas. Ph.D. Dissertation, Texas 
A&M University, College Station. 

Nei, A.K. 1999. Vicia lutea (Fabaceae) new to Texas. Sida 18:1265-1 266. 

Nett, A.K.2000. The vascular flora of Madison County, Texas. M.S. Thesis, Texas A&M Univer- 
sity, College Station. 

Neitsch, C.L. 1994. Soil survey of Madison County, Texas. U.S.D.A., Soil Conservation Service, 
Washington, D.C 

Ramos, M.G. (ed.). 1997. Texas almanac 1998-1999. Dallas Morning News, Dallas. 

Reep, M.D. 1997. Manual of the dicot flora of Brazos and surrounding counties. M.S. Thesis, 
Texas A&M University, College Station. 

RICHARDSON, T.C. 1940. East Texas: its history and its makers. Vol. 3. Lewis Historical Publish- 
ing Co., New York, NY. 

SPEARING, D. 1991. Roadside geology of Texas. Mountain Press Publishing Co., Missoula, MT. 

STarBuck, T.J. 1984. The vascular flora of Robertson County, Texas. M.S. Thesis, Texas A&M 
University, College Station. 

Texas Aumanac. 1951.A.H. Belo Corporation, Dallas, TX. 

Texas Parks & Witouire DEPARTMENT. 1991. Natural Heritage Program special plant list. Austin, 
Texas. (mimeo). 

THarp, B.C. 1926. Structure of Texas vegetation east of the 98th meridian. University of 
Texas Bull. 2606, Austin. 

U.S. DEPARTMENT OF THE INTERIOR. 1998. National historic trail feasibility study and environmen- 
tal assessment: El Camino Real de los Tejas. U.S. Goverment Printing Office, Washing- 
ton, D.C. 

U.S. GEoLocical Survey. 1962-1989. 7.5 minute topographic map series (14 maps) for Madi- 
son County, Texas. Washington, D.C 

Wison, H.D. 1972. The vascular plants of Holmes County, Ohio. M.A. Thesis, Kent State Uni- 
versity, Kent, OH. 


1122 BRIT.ORG/SIDA 19(4) 
Book NOTICE 


Graby L. Wessrer and Conran J. Bane (editors). 2001. Changing Plant Life of La 
Frontera. (ISBN 0-8263-2239-5, hbk.). University of New Mexico Press, 1720 
Lomas Boulevard NE, Albuquerque, NM, 87131-1591, U.S.A. $60.00, hbk., 272 
pages, 41 halftones, 6 maps, 81/2" x 11", 


Table of Contents.—List of Figures; List of Tables; Foreword. Frederick Gehlbach 


1) Introduction 

2) Reconnaissance of the Flora and Vegetation of La Frontera. Grady L. Webster 

3) Climate of La Frontera. M_L. Shelton 

4) Deep History and Biogeography of La Frontera. Thomas R. Van Devend 

5) Historic Vegetation Change in La Frontera West of the Rio Grande. Conrad J. Bahre and 
Charles FE Hutchinson 

6) Climate and Herbivory in Structuring the Vegetation of the Malpai Borderlands. Charles G 
Curtin and James H. Brown 

7) Forest Fire Histories of the Sky Islands of La Frontera. Thomas W. Swetnameta 

8) Fire and Elevational Zonation of Chaparral and Conifer Forests in the Peninsular Ranges of 


La Frontera. Richard A. Minnich 
9) Land Use, Climate Soils, and Forest Structure in the Animas Mountains and the Sierra de los 
jos. Guy R. McPherson and Jose Villanueva-Diaz 
10) Grazing Impacts on Mountain Meadows of the Peninsular Ranges in La Frontera. Joaquin 
Sosa-Ramirez and Ernesto Franco- Vizcaino 
11) Human | nee Vegetation Changes in the Tamaulipan Scrub of La Frontera. Ti mothy E 
Fulbrig} 
12) Oaks of A Frontera. Richard Spellenberg 
13) Coastal Strand Vegetation of La F a Hleana Espejel et al 


Epilogue 

Appendix: Taxonomic Overview of the Oaks of La Frontera 

Glossary of Latin and Vernacular Plant Names 

Literature Cited and Index 

This book isa result of the Changing Vegetation and Flora of the U.S./Mexico Boundar 'y Region sym- 
cats a at the pipeucan Institute or ae uaa to honor the late Forrest Shreve, a pio- 


| biologists, environmentalists, and 


neer C 
jee fem both sides of La Frontera to present a new agenda for the study of the strikingly 
diverse shrub and grassland ecosystems of the U.S./Mexico border. C sae cross border changes in 
Ne eaaas oe from in eispatale land) use a in the region, fire histories, pollen stud- 
botani¢ has fa biogeograp hy, and paleoecol- 
ye U.S./Mexico 


1es 
a oO C o x 
| ] 1 | : Cc . q 
geta ition of t 


this texti 
border should have this bok on his or her shelf.—Kevin D.Janni, ee Research Institute of Texas, 


Fort Worth, TX 76102-4060, U.S.A, kianni@brit.org 


SIDA 19(4): 1122. 2001 


CONSIDERACIONES SOBRE EL 
ORIGEN DELA FLORA ARVENSE Y RUDERAL 
DEL ESTADO DE QUERETARO, MEXICO 


José Aurelio Colmenero Robles Concepcidn Rodriguez Jiménez 


ene ee eee and Fernandez Nava Rafael 
bias lit, Nar] | ~ Vlawal tra de RatAnica 

artado Postal /5- Escuela Nacional de Ciencias Bioldgicas 
Col i 07300, ME a DF Instituto P lité i Nac ql 


Col. Santo Tomas, Apartado Postal 17-564 
11410 MEXICO, DF 


RESUMEN 


Se efectuo un estudio del origen de la flora arvense y ruderal presente en el estado de Querétaro, 

México. Una de las principal de la distribuci6n es la ubicacion geografica del estado, paso 

obligado hacia ne regiones norte y sur del pais. Otro factor implicado fueron las actividades 

agropecuarias practicadas por los grupos nativos  Mehicghimecas y arenes); mocificatas 
con | 


males. 
Se epistracene a fe esr 211 géneros y 47 familias de las regi gricolas del do; d 

I Compositae (81 spp p.), G i (6] p.), Euphorbi (22 spp.) 
Leguminosae (17 | Sol PE ), isc (12 spp ) Gaiciheras (12 spp ) y Convolvulaceae 
(10 spp.) de las que el 66% fueron lectadas en cone y el 14% en la vegetacion rud- 
eral. Se listan 125 espe nsideradas nativas, la mayoria de las cuales pertenecen a las familias: 
Compositae, Gramineae, Solanaceae, a CER Ne eien y Cucurbitaceae; ademas de 
las 57 de origen norteamericano, hay 57 de América tropical, 29 europeas, 23 sudamericanas, 20 


euroasiaticas, 16 asiaticas, 8 caribenas, 8 africanas, ; - la region de Oceania y 2 de Centroamérica. 


ABSTRACT 


The origin of weeds and ruderal species from Queretaro, Mexico was studied. One of the principal 
causes for the distribution of weeds of diverse origin is the location of the state, through which much 
north-south ves was focused. Another factor was the agricultural practices of native people 
(chichimecas y ), later modified by European immigrants, with the introduction of new crops 
and diverse Mania as 


ciates. 

ed a species, 211 genera, and 47 ami The families with largest number of taxa 

are eee (81 spp), Gramineae (61 spp.), Euph 22 spp.), Leg (17 spp.), Solan- 
aceae (16 spp.), Malvaceae (12 spp.), Cruciferae a2 spp.) and Convolvulaceae (10 spp.) of which 66% 
were collected in field crops and 14% in ruderal vegetation. There are 125 species considered native, 
most of theta belong to the families: Compositae, Gramineae, Solanaceae, Euphorbiaceae, 
€ ucurbitaceae. In addition to 57 from North America, there were 57 from Tropi- 
cal Aeriek: 29 from Europe, 23 from South America, 23 from European-Asiatic, 16 from Asia, 8 
from Caribbean, 8 from Africa, 2 from Oceania, and 2 from Central America. 


SIDA 19(4): 1123 — 1145. 2001 


1124 BRIT.ORG/SIDA 19(4) 


INTRODUCCION 


El estudio de las especies de maleza en México se ha incrementado 
considerablemente en los ultimos anos, llamando la atencion el aumento de 
trabajos registrados en este campo de la Biologia. Estas plantas presentan dos 
propiedades fundamentales: resistencia a situaciones adversas y agresividad, 
que les permiten causar serios danos a los cultivos agricolas: incrementando el 
costo de produccion y disminuyendo los rendimientos. Cumplen sin embargo 
funciones ecoldgicas importantes al ser consideradas pioneras 0 colonizadoras 
en areas con vegetacion alterada o en proceso de sucesion, donde con sus 
sistemas radiculares, a veces muy extensos, retienen el suelo y evitan el riesgo 
de erosion, proveen de néctar y polen a insectos meliferos, sirven como 
fertilizantes y contribuyen a la formacion del suelo (Villarreal 1983). 

Caracterizar a las plantas arvenses o ruderales no es facil, sin embargo, se 
considera que todas presentan: L.-Alta produccién de semillas con elevada tasa 
de germinacion y desarrollo en distintas condiciones ambientales. 2.- 
Crecimiento acelerado desde la etapa juvenil hasta la floracién. 3.- 
Autocompatibilidad 0 autofertilizacion, pero no siempre son autégamas o 
apomicticas. 4.-Diversos mecanismos de dispersion a distancias cortas y/o 
largas mediante estructuras morfoldgicas presentes en sus frutos y semillas. 5.- 
Alto grado de te proauccian vegetativa (Baker 1974; Espinosa y Sarukhan 1997). 

Reciben estas plantas dix bres comunes, tales como: malas hierbas, 
plantas indeseables, yuyos, plantas arvenses o agrestales y en nahuatl son 
reconocidas como acahuales, jehuites 0 jihuites (Villegas 1970). 

Desde el punto de vista ecologico se reconocen dos grandes grupos, las 
plantas arvenses, que se encuentran ligadasa los cultivos y las plantas ruderales, 
que son propias de poblados, lotes baldios y multiples vias de comunicacién 
(Rzedowski y Rzedowski 1957; Rzedowski 1978). 


ANTECEDENTES 


Las especies de arvenses y ruderales de la Reptblica Mexicana varian de una 
region a otra, en funcion de los distintos factores ambientales de cada sitio, 
quedando su abundancia sujeta a las actividades antropégenas (Rzedowski 
1978) 


Por lo que se refiere al origen de las arvenses mexicanas diversos autores 
han hecho mencioén de la procedencia de un buen numero de ellas en estudios 
realizados para vari g del pais (Rodriguez 1967; Villegas 1970; Agundis 
y Rodriguez 1978; Villarreal 1983; Nava 1991; Rzedowski 1993; Rzedowski y 
Rzedowski 1993; Vibrans 1998a); lo mismo que para las plantas ruderales 
(Rapoport et al. 1983; Vibrans 1996, 1998b). El analisis de los datos recabados ha 
llevado a concluir que en este grupo de plantas prevalece cuantitativamente la 
maleza autoctona (Rzedowski 1978, 1991; Bye 1993). El arribo de plantas 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 1125 


introducidas de origen europeo y asiatico se realiza en los siglos del XVI al XVIII, 
con motivo de la colonizacién espahola y el intercambi 
de manera inmediata. Entre | 


] que se efectuod 


al 


eneste periodo 
destacan: Asphodelus, Brassica, Conium, Eruca, Fumaria, Melilotus, Sisymbrium, 
Sonchus, Taraxacum y Verbascum. En cuanto al componente de origen africano 
se registran a la fecha alrededor de 42 taxa para el pais, sobresaliendo los géneros 
Bryophyllum, Chloris, Cynodon, Eleusine y Eragrostis (Rzedowski & Rzedowski 
1990). En la actualidad se cuenta con nuevos registros de exoticas introducidas 
en nuestro pais, algunos ejemplos de ellos son Amaranthus muricatus (Moq.) 
Gillies ex Hicken, Bellis perennis L., Chenopodium giganteum D. Don, Digitaria 
wallichiana (Wight & Arnold) Stapf, Guizotia abyssinica (L.f.) Cass. y 
Hypochoeris radiate L. (Vibrans 1998b). 

presencia deal Sticas esta estrechamente relacionada 
con laintroduccion del la ganaderia a nuestro pais. Los primeros hatos ganaderos, 
provenientes de las Islas de Cuba y la Espanola a cargo de Nuno Guzman 
llegaron en 1527 a Panuco, Veracruz. Este hecho promovi6 la formacion de tres 
regiones ganaderas en la Nueva Espana: la de Panuco, la de Llanos de Almeria 
(hoy las tierras bajas de Misantla, Vega de Alatorre y Nautla) y lade Papaloapan. 
Este acontecimiento favorecié también el arribo de diversas especies forrajeras 
para la alimentacion del ganado y a su vez la paulatina transformacion de los 
ecosistemas del tr6pico hamedo. Alrededor de 1870 se registra por primera vez 
el pasto guinea 0 privilegio (Panicum maximum Jacq.) en las tierras del norte 
de México y se piensa que éste fue utilizado como cama para los esclavos 
africanos tt TanSpo} tados masivamente hacia América (Challenger 1998). 

Las pri forrajeras de origen africano presentes en México 
son: el zacate Rhodes (Chloris gayana Kunth), del sur y este de ese continente; el 
pasto estrella gigante (Cynodon plectostachyus (K. Schum.) Pilg,), del oriente; 
Dichanthium annulatum (Forssk.) Stapf., del norte; el zacate pangola (Digitaria 
decumbens Stent), del sur; el zacate kikuyu (Pennisetum clandestinum Hochst. 
ex Chiov.), de Africa central y oriental. Otras especies como el zacate elefante 
(Pennisetum purpureum Schumach.), el pasto natal o pasto rosado 
(Rynchelytrum repens (Willd.) C.E. Hubb ) y el zacate Johnson (Sorghum 
halepense (L.) Pers.) este Ultimo nativo del area mediterranea y posiblemente 
de ascendencia africana (Rzedowski & Rzedowski 1990). Al parecer muchas de 
estas especies se cultivan en nuestro territorio desde el primer tercio del siglo 

X, como forraje para el ganado cebu (Barrera 1996). De acuerdo con Davila y 
Sanchez (1996), existen alrededor de 49 géneros de gramineas introducidos a la 
fecha, la mayoria de sus representantes se consideran malezoides. 

Otros factores que favorecieron el desarrollo de especies exoticas en el Alti- 
plano durante épocas pasadas fueron la tala de los bosques para su utilizacién 
como madera y combustible y la introduccion de ganado cuyo sobrepastoreo 


1126 BRIT.ORG/SIDA 19(4) 


sobre la vegetacion nativa, promovi6 la distribucion de especies como Urtica 
dioica L., algunas otras de Plantago y del trébol blanco, Trifolium repens L., 
plantas que se extendieron muy rapidamente, de tal manera que los aztecas 
llamaron a esta ultima “ocoxichtli de castilla,” nuevamente de acuerdo a la 
apreciacion de Challenger (1998). 

Los estudios sobre el inventario y el conocimiento de las plantas arvenses y 
ruderales para el estado de Querétaro son relativamente escasos, se inician en 
los anos de 1983-1984 con Agundis, O. y colaboradores quienes elaboran el 
primer inventario floristico de arvenses para la zona centro del estado, 
registrando un total de 234 especies en 14 cultivos principales (inédito). 

De acuerdo con Fernandez et al. (1993), existe ademas de la zona agricola, 
un area ganadera de gran importancia para el estado, que no registraba ningtn 
trabajo que permitiera conocer las plantas malezoides que crecian en ella, por 
lo que se abocaron a realizar este inventario para el noreste de Querétaro (Ar- 
royo Seco, Jalpan, Landa de Matamoros y Pinal de Amoles). Los resultados 
establecieron que existia una cantidad considerable de arvenses, alrededor de 
172 especies, como consecuencia del intensivo uso del suelo en esta region. 

Zepeda (1996), emprende el estudio de las especies de malezas presentes en 
los cultivos de maiz y de sorgo en el Centro-Sur de Querétaro. Sus resultados 
indican la presencia de 21 familias y 53 taxa, destacando las familias Gramineae 
(16 spp.) y Compositae (10 spp.). 

Suarez y Serrano (1997), en su publicacion “Principales malezas (arvenses) 
en el campo queretano,” citan 14 especies su respectiva descripcion, imagen 
fotografica e indicando los cultivos donde prosperan. Finalmente, Villasenor y 
Espinosa (1998), enlistan un numero considerable de taxa para Querétaro. 


AREA DE ESTUDIO 


Querétaro se ubica en la Region Central de la Republica Mexicana entre los 
paralelos 20°01’16" y 21°35°38" de latitud norte y los meridianos 99°00°46" y 
100°35°46", esta limitado por 5 estados: San Luis Potosi al Norte y Noroeste, 
Hidalgo al Este, Guanajuato al Oeste y Suroeste, Estado de México y Michoacan 
al Sureste y Sur respectivamente (INEGI 1986). 

Querétaro ha sido por siglos un estado agricola, aunque en la actualidad la 
industria, ha cobrado mayor importancia, pero a pesar de esto, la agricultura y 
la ganaderia forman parte de la columna vertebral de su economia; existen mas 
de 30 cultivos, destacando por la magnitud de la superficie sembrada maiz, trigo, 
alfalfa, frijol, sorgo, jitomate, lenteja, chile verde, cebada, cebolla y ajo entre otros 
(Ramirez 

La distribucion de las areas agricolas se establece de acuerdo al criterio de 
Holt (1970) en 3 Distritos: Norte, Centro y Sur, incluyendo algunos datos de las 
caracteristicas fisicas del ambiente tomados de INEGI (1986), asi como los tipos 
de vegetacion considerados por Zamudio et al. (1992) (Fig. 1). 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 1127 


Distrito Norte 

Queda comprendido en la Provincia Sierra Madre Oriental, de forma particular 
dentro de la Subprovincia Carso Carstico; el clima que predomina corresponde 
a los calidos (Aw) y templados (A)C(wo-w2), razon por lo cual se presenta de 
norte a sur un gradiente térmico de calido a frio, con precipitaciones de 770- 
1050 mm de lluvia promedio anual y temperaturas promedio anual de 18-20 
C; la conformacion geoldgica esta constituida por rocas sedimentarias, calizas 
y lutitas; los suelos predominantes son de los tipos litosoles, luvisoles cr6micos, 
luvisoles 6rticos y rendzinas; la actividad agricola tiende a ser de temporal con 
cultivos de ciclo anual y Poe areas dedicadasa la fruticultura; los tipos de 
vegetacion que se presentan en ranas son: Bosque de Pinus, Bosque 
de Abies, Bosque Meséfilo de Montana, Bosque Tropical Caducifolio y Matorral 
Submontano. Comprende los municipios de Pinal de Amoles, Arroyo Seco, 
Jalpan y Landa de Matamoros. 


Distrito Centro 

Esta inserto en la provincia Mesa del Centro, de forma particular en las 
subprovincias de las Sierras y Llanuras del Norte de Guanajuato y en una seccion 
de la Provincia del Eje Neovolcanico. Los tipos de clima corresponde a los 
semisecos (BS}) y los secos (BS), la cantidad de lluvia es alrededor de los 370- 
630 mm de precipitacion promedio anual y la temperatura oscila entre los 20- 
25° C; geologicamente ‘Prccommnat rocas igneas, es y otras de caracter 
sedimentario; los ti les, luvisoles crémicos, 
regosoles etttricos, ore haplico y feozem crémico; en cuantoa la agricultura 
tiende a ser variada, se caracteriza por ser la region del cultivo de vid, aguacate, 
mango y pequenias areas de cultivos de ciclo anual; los tipos de vegetacion que 
sobresalen entre las canadas, laderas y elevaciones orograficas de mediano 
tamano son: Bosque de Quercus, Bosque de Juniperus, Encinar Arbustivo, 
Matorral Rosetofilo y Matorral Microfilo. Su extension incluye los municipios 
de Cadereyta, Ezequiel Montes, Penamiller, San Joaquin y Toliman. 


Distrito Sur 

Se ubica en lo que se conoce como Provincia del Eje Neovolcanico y en una 
seccion pequena de la Subprovincia Mil Cumbres. Los tipos de climas 
principales son los secos (BSg), semisecos (BS)) y templados (Cw), la cantidad 
de lluvia promedio anual se sittia entre los eacicutcs de i 400- 860 mm, con 
temperaturas promedio anual de 18-25'C; las 

basaltos, tobas, calizas, brechas volcanicas y aluviones; entre Jos tipos de suelos 
sobresalen principalmente el vertisol pélico (muy ricos en materia organica) y 
el feozem haplico; destaca la agricultura de riego de dos ciclos (primavera e 
invierno); los tipos de vegetacion para esta seccién son el Bosque de Pinus 
cembroides, el Bosque de Quercus, el Matorral Crasicaule y el Pastizal. Esta 
conformado por los municipios de Amealco, Colon, Villa Corregidora, 


1128 BRIT.ORG/SIDA 19(4) 


DISTRITOS AGRICOLAS DEL ESTADO DE QUERETARO 


EDO. SAN LUIS POTOSI 


EDO. 
GUANAJUATO 


DISTRITO NORTE 


001 ARROYO SECO 

002 JALPAN 

003 LANDA DE MATAMOROS 
004 PINAL DE AMOLES 


005 PENAMILLER 
ye 
EDO*e we 006 SAN JOAQU 
MICHOACAN 007 TOLIMAN 


008 CADEREYTA 
009 EZEQUIEL MONTES 
DISTRITO SUR 


010 COLON 
011 TEQUISQUIAPAN 
012 ELM 


018 AMEALCO 


Fic. 1. Ubicacion del area de estudio. 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 1129 


Huimilpan, El Marqués, Pedro Escobedo, Querétaro, San Juan del Rio y 
Tequisquiapan. 


METODOLOGIA 


Para llevar a cabo la presente investigacion se realizé trabajo de campo en dos 
etapas (1982 a 1983 y de 1992 a 1993) colectando material en las diferentes 
regiones y areas agricolas de Querétaro, el que se encuentra depositado en el 
Herbario ENCB; de manera complementaria se revisaron ejemplares de maleza 
del Instituto Nacional de Investigaciones Agricolas, SARH, actualmente Insti- 
tuto Nacional de Investigaciones Forestales y Agropecuarias (INIFAP) y se 
consultaron distintas fuentes bibliograficas relacionadas con la presencia de 
maleza en este estado (Zepeda, A. op. cit; Suarez y Serrano op. cit.) y flora re- 
gional (Arguelles et al. 1991). Para la designacion del lugar de procedencia de 
las especies, fue necesario consultar diversas obras (Marzocca 1976; Rzedowski 
1978; Holm et al. 1977; Boelcke 1986), trabajos floristicos o grupos especificos 
(Rodriguez op. cit; Rollins 1981; Villarreal op. cit; Fryxell 1988; Nava op. cit. 
McDonald 1991; Rzedowski 1993; Lira et al. 1998), labor que result6 compleja, 
debido a los diferentes criterios y multiples confusiones que se aprecian entre 
los autores. Los conceptos designados en cuanto al origen se especifican al 
principio de listado floristico (Tabla 1). 


RESULTADOS Y DISCUSION 


Se registraron para el estado 347 especies que se comportan como arvenses 0 
ruderales comprendidas en 211 géneros y 47 familias, un centenar menos que 
las especies de arvenses reconocidas para el Valle de México (Espinosa y 
Sarukhan 1997) y un numero inferior al reconocido para Querétaro por 
Villasenor y Espinosa (1998). El criterio de estos ultimos autores es muy amplio 
ya que consideran dentro del grupo a las arvenses, ruderales y las especies 
secundarias que dan origen a las “acahualeras”(Rzedowski 1978). 

Destacan por el numero de taxa registrados las familias Compositae (81), 
Gramineae (61), Euphorbiaceae (22), Leguminosae (17), Solanaceae (16), 
Malvaceae (12), Cruciferae (12) y Convolvulaceae (10), que en conjunto 
representan el 66% de las especies colectadas en terrenos de cultivo y areas 
circunvecinas, el resto (116 especies) queda comprendido en 39 familias (Fig. 2). 

Al igual que en otros trabajos relacionados con el estudio de la flora arvense 
(Rodriguez op. cit; Rzedowski op. cit., Zepeda op. cit; Suarez y Serrano op. cit), 
las compuestas y las gramineas ocupan el primer lugar en importancia, seguidas 
por las euforbiaceas y leguminosas que también se perfilan como grupos 
importantes en el estado. 

De los grupos de plantas cuya presencia se ve favorecida por la actividad 
del hombre, las ruderales estrictas en Querétaro ocupan solo el 15%, quedando 


1130 BRIT.ORG/SIDA 19(4) 


Tasta 1. Listado floristico de las plantas arvenses y ruderales del estado de Querétaro, México 
indicando lugar de origen. 


CONCEPTOS 


NATIVAS (NT)= endemicas al territorio mexicano; NORTEAMERICANAS (NA)= tienen su origen 
en la parte que comprende los paises de Canada, Estados Unidos y la regidn Norte y centro de 
México; CENTROAMERICANAS (CA)= implica el area desde el Sur y Sureste de México hasta el 
Norte y NW de Colombia; CARIBENAS (CB)= se considera la porcion Sur de Texas y la Peninsula de 
La Florida, parte de la Costa del Golfo de México, Islas del Caribe, parte de sn atlantica costera 
de Centro ae hasta el delta del Rio Orinoco de los paises de Colombia y Venezuela; 
SUDAMERICANAS (SA)= al Norte y NW de Colombia hasta la Argentina y ee RTE 
iit re comprende las parciones principalmente bajas calido-humedas de México, 

tro ae Sudanne érica; PEUROEER? ae incluye a: Peninsula Ibérica, slain Peninsula de 


abarca el area central entre Europa y ia ASIATICAS (AS)= se inicia a OesCe la Peninsula de en 
hasta el Caucaso y Siberia, asi como la seccion Sur de Asia China, Vietnam); AFRICANAS 


(AF)= se situan en la porcidn Norte de Africa, Egipto hasta el Sur del Cabo de Buena Esperanza, 
también la Isla Socotora y el Golfo de Guinea; OCEANIA (OC)=_ es el area de Australia, Nueva 
Guinea, Nueva Zelanda y Tasmania, asi como la gran cantidad de Islas en la Polinesia. 


ESPECIES 0 R I G E N 
NT NA CA CB SA AT EU EA AS AF OC 


EQUISETACEAE 
Equisetum hyemale L. 
subsp. affine (Engelm.) Calder & R.L. Taylor X 
nica 


(L.) var. pubescens Underw, X 


AIZOACEA 
Mollugo iia L. X 
ACANTHACEAE 
tne ia tpl (Vahl) Pers. X 
AMARA ACEAE 
Amaranthus hybridus L. 
Amoaranthryic nalr ic Watson 4 
Gomphrena deeimrben s Jacq. X 
Gomphrena pringlei Coult.& Fisher X 
petra AGEN 


~< 


Va 


Sarcostema geno des Decne. 
subsp W. Holm 
BORAGINA ACEA 

Heliotropium engoseu Murray X 
ede 

atea micrantha (Kunth) McVaugh X 
a ne (A. Gray) McVaugh X 
Lobelia fenestralis Cav. X 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 


1131 


ESPECIES 


0 


R 


NT NA CA CB SA AT EU EA AS AF OC 


il diate ie 
leome aculeata L 
ore 
Drymaria arenarioides Willd. 
Drymaria glandulosa Bartl. 
ples ate Benth. 
Spergulaa isl 
Stellaria see : Willd. 
CHENOPODIACEA 
ae asieeta a 
Ch mL. 


dittm ambrosioides | 


Ch 
Chenopodium saat ye 


Pp Mt: 
Phah podit mmm irale L 


COMMELINACE EAE 
Commelina diffusa Burm. f. 


yuan erecta peed) Schitl 


Hisecandia purpurascens aeehauel Handlos 


subsp. purpurascens 
COMPOSITAE 
Achillea millefolium L. 
aan centate ri Llave & Lex. 


— 


Anibmotia psilostac a 
Aphanostephus ramosissimus DC. 
Artemisia ludoviciana Nutt. 

subsp. mexicana (Willd.) Keck 
Aster spinosus Benth. 
Aster subulatus Mic 
Baccharis salicifolia (Ruiz & Pav.) Pers. 
Bidens aurea (Aiton) Sherff 
Bidens bicolor Greenm. 
Bidens bigelovii A. Gray 


Bidens serrulata (Poir) Desf. 

on Leela alis Less. 

is (Hemsl.) Petr. 

ae bonariensis (L.) Cronquist 

Conyza canadensis (L.) Cronquist 
ronopifolia Kunth 


TTY ZuU 


(A. Gray) W.W. Payne 


~< 


x< 


x< 


=< 


~< 


1132 BRIT.ORG/SIDA 19(4 


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Conyza sophiifolia Kunth X 
Cosmos bipinnatus Cav. X 
tula australis (Sieb.) Hook. f. X 

Dugesia j A. Gray X 
Dyssodia papposa ee Hitche. X 

Dyssodia pinnata (Cav.) B.L. Rob. var. pinnata X 
Dyssodia porophyllum (Cav.) Cav. X 
Eclipta postrata (L) L X 
Erechtites hieracifolia ie Raf. 


acaloides ae ie 


Frygeron karvinek 
xy 


X 
X 
dad X 
X 
X 


a 
ere trinervia Soci C. Mohr 
Florestina pedata Cass. 
Florestina tripteris DC. X 
Galinsoga parviflora Cav. 
Galit quadriradiata Ruiz & Pav. 
j ill. 


io 
Gn apt 


Helenium mexicanum Kunth 
Heliopsis annua Hemsl. 
Heterosperma pinnatum Cav. X 
Heterotheca inuloides Cass. var. inuloides X 
Jaegeria hirta (Lag.) Less. X 
Lactuca graminifolia Siew X 
Lactuca serriola L. X 
SotaOn MIGROS (Vill.) Mérat xX 
n S.Watson 
Melampodium divaricatum (Rich.) DC. 
Melampodium perfoliatum (Cav.) Kunth 
Melampodium sericeum Lag 
Parth INIIM Aininnogatifa, (Ort.) Rollins 
Picris echioides L. X 
pinalopappusr roseus (Less.) Less. var. roseus 

V. 


X 

X 

X 

Grindeli ia subdecurrens DC. X 
Xx 

X 


lamp 


<x <x KK OX 


Piqueria trinervia Ca X 
orophyllum linaria ( 
Pseudelephantopus spicatus ane ) Rohr X 
Psilac tis A a Sch. Bip. X 
Saba is (Kunth) Cass. 4 
Santo aprocumbers Lam. X 
nnata (Lam.) Kuntze 
var. virgata (La Llave ex Lex.) Heiser X 
adie se Poepp. & Endl. X 
(Cav.) Pers. X 


Sinsia joer de (Cav.) S.F. Blake var. foetida xX 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 1133 


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~< 


Sonchus asper (L.) Hill 
Sonchus oleraceus L. x 
Spilanthes oppositifolia (Lam.) D'Arcy X 

Stevia serrata 


SJ 
Ka) 
a 
ag 
ny 
nA 
> 
Ss) 
2 = 
S 
OE ses 
Ba*s 
<x KKK OK 


Tagetes lunulata Ort. 
Tagetes micrantha Cav. X 
Taraxacum officinale Weber x 
Tithonia tubiformis (Jacq.) Cass. X 
Tridax coronopifolia nee Hemsl. xX 
Midax ena (Cav .) Hers| 
i, 


~< 


es v.) Benth. & Hook. xX 
Viguiete eee (Cav.) on xX 
Xanthium strumarium 
ar. canadense (Mill. Toit & A. Gray x 
Za ae triloba (Ort.) Pers. Xx 
Zinnia peruviana (L.) L. X 
so ed VUES CEAE 


pane sericea Sie 

Ipomoea hederifolia L. 

mee indica ( Burm.) Merr. 

Ipomoea neei (Spreng.) O'Donell 

Ipomoea bani (L.) Roth 

omnis a stans Cav. X 

moea trifida (Kunth) G.Don X 

ees triloba L 

Merremia ae ata (L.) Hallier f. X 
CRASSULACEA 
Bryophyllum ue (Lam.) Kurz xX 


x KK OK OK 


x 


Brassica nite .) Koch X 
Capsella bursa-pastoris (L.) Medik. X 
Descurainia pinnata (Walter) Britton xX 

Descurainia virletii (Fourn.) O.E. Schulz xX 

pce sativa Mill. X 


m (Ny itt.) 
UINUTE. 


fa Cc 

Lepidium virginicum L. var. pubescens (Greene 
Lesquerella argentea (Schauer) S.Watson 
Raphanus raphanistrum L. xX 
Rapistrum rugosum (L.) All. 
Sisymobrium irio L. xX 


hae 


X 
Hitchc. X 
X 


~< 


1134 BRIT.ORG/SIDA 19(4) 


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CUCURBITACEAE 
Cucurbita foetidissi ima Kunth 
GC ta (Torr. & A. Gray) Arn. X 
Oxlante ribiflora (Schltdl.) Cogn. X 

Echinopepon milleflorus Naudin X 

AA ali bh +p | : XxX 


~< 


x< 


Sicyos deppei G.Don 

Sicyos laciniatus L. 

Sicyos parviflorus Willd, X 
CYPERACEAE 

Bulbostylis funckil (Steud.) C.B. Clarke 

Bulbostylis juncoides (Vahl) Kuk. X 

Cyperus esculentus L. X 

Cyperus manimae Kunth 


x< 


~x< 


x xX 


yperu us L. 
Cyperus seslerioides Kunth xX 
Cyperus surinamensis Rottb. X 
Eleocharis acicularis (L.) Roem. & Schult. X 
Eleocharis geniculata (L.) Roem. & Schult. X 
EUPHORBIACEAE 
Acalypha indica L. 
var. ora (Mull.Arg.) Pax & K. Hoffm. xX 
A infect” Po & Endl. X 
pei ostryifolia Rida, X 
Acalypha setosa A. Rich. 
Alchornea ee Sw. X 
Croton dioicus Cav 
Fup Ho abomarginot Torr.& A. Gray X 
Eu ichx. 


~< 


~x< 


~< 


fuphorola iaeceuna Bois X 
ens rbia glomerifera mils ) Wheeler 

Euphorbia oe 

Euphorbia 

Euphorbia eae fal. 

Eupborbia bs Lag. 

Euphorbia peplus L. X 
Euphorbia ae Aiton X 
Euphorbia villifera Scheele X 
Euphorbia serpens Kunth X 
Euphorbia sti ictospora Engelm. 


x KX KK 


x< 


SL. 
Tragia nepetifolia Cav. X 
GERANIACEAE 
Er m tarlum (L.) UHér. X 
Geranium seemannii Peyr. X 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 1135 


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GRAMINEAE 

AON seimivemiciiar seen C.Chr. X 

Kunth xX 
Aristicta ndstensionis L. X 
Avena fatua L. X 
Routsioud barbata Lag. 
dula (Michx.) Torr. X 

Bouteloua hirsuta Lag. X 
Bothriochloa saccharoides Sw 

var. torreyan aie Gould xX 
Brachiaria meziana Hitche. 
Brachiaria plantag! ae Hitchc. X 
Bromus catharticus Vah| 4 
Buchloé ues ee sie ) Engelm. X 

enchrus echinatus L. 

ie nchrus incertus M.A ue X 
Cenchrus myosur Kunth 


~< 


x< 


x< 


Cenchrus Sane Kunth 
Chloris rufescens a 
Chloris virgata 

Cynodon dacty as Pers. X 
Digitaria ciliaris (Retz.) Koeler X 
Digi itaria insularis (L.) Fedde X 

Digitaria ternata (A. Rich. ) Stapf xX 
Echinochloa colonum (L.) Link 
nalaall ne crus-galli P. Beauv. xX 


x KM 


x< 


Echinochloa opli (Kunth) chase xX 

Eleusine indica (L.) Gaertn. X 

Eleusine multiflora Hochst. X 

Eragrostis barrelieri Daveau X 

Eragrostis cilianensis (All.) Vignolo X 

Eragrostis glomerata it L.H. Dewey 

Eragrostis intermedia Hitchc. X 

Eragrostis lugens Nees X 

Eragrostis mexicana (Hornem.) Link Xx 

Eragrostis pectinacea (Michx.) Nees x 

page plea (L.) P. Beauv. X 
mina ta (C. Presl.) Kunth X 


x 


Hilari id eenennel ides Kunth X 
Lasiacis ruscifolia (Kunth) Hitchc. 4 
nine hloa dubia (Kunth) Nees X 


Lolittm mu/stifloriym | Lam X 


x 


one phleoides Kunth var. phleoides 
Muhlenbergia rigida (Kunth) Trin. X 


1136 BRIT.ORG/SIDA 19(4) 


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Oplisment mpositus (L.) P. Beauv. xX 
Panicum decolorans arin X 

Pani. -rum fascicitlatiim Sw. xX 

P WicimM miliacelywmy I XxX 
Pani bt Kunth X 

Panicum recone Scribn. > 4 


Bae setuM purpureum peanumnad X 
aannua L 
Polypogon monspeliensis (L.) Desf. X 
Rynchelytrum repens (Willd.) C.E. Hubb. X 
Setaria geniculata (Lam.) P. Beauv. X 
Setaria grisebachii E. Fourn. X 
Setaria lutescens (Weigel) C.E. Hubb 
iq verticillata (L.) P Beauv. Xx 
Sorghum b asus (L.) Moench 
ense (L 


x< 


x 


1, 
fae 


~< 


GUTTIFERAE 
Hypericum philonotis Cham. & Schltdl X 
HYDROPHYLLACEAE 
Nama dichotomum (Ruiz & Pav.) Choisy var. dichotomum X 
IRIDACEAE 
Sisyrinchium schaffneri S.Watson X 
LABIATAE 
Marrubium vulgare L. xX 
Molucella laevis L. X 
Salvia hirsuta Jacq. 
Salvia reflexa Hornem, 
Salvia tiliifolia Vah| 
tachys agraria Cham. & Schitdl. 
LEGUMINOSAE 
Astragalus nuttallianus OC. X 
Crotalaria pumila 
mae cifortana wil X 
I iton) Bullock xX 
Dal ed ee Ort. X 
Desmodium procumbens (Mill.) Hitch 
esmodium sericophyllum Schitdl. X 
eee glauca (Ort.) Eifert X 
L. 


wn 
x x KK 


x< 


<< 


Medicago pobmorpha. var. vulgaris (Benth.) Shinners X 

Melilotus albus D 

as eee indicus i ‘ All. xX 
olus heterophyllus Willd. X 

Sia uniflora (Mill) Irwin & Barneby x 


~< 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 


1137 


ESPECIES 


— 


NA 


R I G E N 
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Trifolium locarpum Lojac 
Trifolium repens L 


Vicia americana Muhl. ex Willd. var. americana 


LILIACEAE 

Allium glandulosum Link & Otto 
Alara gracile (Aiton) Stearn 
LYTHRACEA 

ee aeuipetl Cav. 
Cup is Jacq.) Macbr. 


— 


ee ne Kunth 
MALVACEAE 

Anoda cristata (L.) Schltdl. 

Anoda thurberi A. Gray 

Herissantia crispa (L.) Brizicky 

Malva parviflora L. 

Malvastrum americanum (L.) Torr. 


At ne aa coromandelianum (L.) Garcke 
Modiol lini (L.) G.Don 


Sida abutifolia Mill. 
Sida rhombifolia L. 
Sida spinosa L. 
5 aan ee ene G.Don 
pidiu se (L) K 
MARTYNIA CEAE 
Proboscidea louisianica (Mill) Thell. 
subsp. fragrans (Lindl. Bretting 
NYCTAGINACEAE 
al ionia incarnata L. 


q Nill 
MH, 


> 


eb L 

ONAGRACEA 

cour coccinea Pursh 
Cav 


eee octovalvi sUaeer ) Raven 
Oenothera pubescens Willd. 
Oenothera rosea L'Her 


L 
Oenothera tetrapterd Cav. 


OXALIDACEAE 


~< 


x< 


X 


Oxalis corniculata is su ea albicans (Kunth) Lourteig 


Oxalis decaphylla 
Oxalis latifolia ae 
PAPAVERACE ag 


Argemone fii Icand L. 


Argemo Sweet subsp. ochroleuca 


X 


Xx 


~< 


1138 


BRIT.ORG/SIDA 19(4) 


ESPECIES 


0 R | G E 
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PLANTAGINACEAE 
Plantago lanceolata L. 


Plantago linearis var. mexicana (Link) Pilg. 


Plantago major L. 
POLYGONACEAE 


Dolvanniim avicilar, L 


Polygonum mexicanum Small 

Polygonum punctatum Ell. 

Polygonum segetum Kunth 

Rumex acetosella L. 

mex conglomeratum Murr. 

Rumex Crispus 

Rumex obtusifolius L. 
eee 
C 


PRIMULACEAE 
Annnantllic ary 


RESEDACEAE 
Reseda luteola L. 
ROSACEAE 
Alchemilla aphanoides 
var, subal, is (Rose) L.M. Perry 


RUBIACEAE 
Bouvardia ternifolia (Cav.) Schltdl. 
Crusea diversifolia (Kunth) W.R. Anderson 


Spe CHIOACUCE tenulor Ly 
SCROPHULARIACEAE 

Bacopa procumbens (Mill.) Greenm. 

Castilleja arvensis Cham. & Schitdl. 

Veronica peregrina L. subsp. xalar 
SOLANACEAE 

Datura discolor Bernh. 

Datura innoxia Mil 

Datura stram 

Jaltomata pees ea (Cav.) J.L. Gentry 

Nicotiana glauca Graham 

Nicotiana trigonophylla Dunal 

Petunia parviflora Juss. 

Physalis aequata Jacq. 


Physalis phi ladelphi ca | 


Physalis solanaceous (Schitdl.) Axelius 
Physalis viscosa 

Solanum americanum Mill. 

Solanum elaeagnifolium Cav. 
Solanum mozinianum Dunal 


~< 


~< 


x< 


Xx 
is (Kunth) Pennell xX 


~< 


<< 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 1139 


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Solanum nigrescens M. Martens & Galeotti X 
C lant mm roctratiyim Dunal XxX 
STERCULIACEAE 
Melo fi ee idata L. X 
OEE 
Anium le jee phyllum ae F.Muell. X 
Eryngium comosum F. Delaroche X 
egw serratum a X 
tee iadium rellieense (Kunth) Matthias X 
VERBENACEAE 
ee prismatica (L.) Kuntze X 
Lippia nodiflora (L.) Michx. X 
Verbena bipinnatifida Nutt. X 
Verbena gracilis Desf. X 
Verbena menthaefolia Benth. X 
VIOLACEAE 
Hybanthus attenuatus (Humb. & Bonpl.) Schulze-Menz X 
ZYGOPHYLLACEAE 
Kalstroemia rosei Rydb. X 
Tribulus terrestris L. X 


el 85% restante en la categoria de arvenses, existiendo algunas como: Anoda 
cristata, Aster subulatus, Bouteloua hirsuta, Euphorbia heterophylla, Galinsoga 
parviflora, Jaltomata procumbens, Lobelia fenestralis, Malva parviflora, 
Malvastrum coromandelianum, Modiola caroliniana, Sida spinosa, 
Melampodium divaricatum, Physalis viscosa, Tithonia tubiformis, Trifolium 
repens y Zinnia peruviana que se ubican tanto en zonas perturbadas como en 
terrenos de cultivo. 

Por otra parte, existe una aparente regionalizacion de un grupo de 61 taxa, 
para cada uno de los 3 Distritos. En el Norte se encuentran 26 taxa entre los que 
destacan: Achillea millefolium, Alchornea latifolia, Andropogon condensatum, 
Bryophyllum pinnatum, Convolvulus arvensis, Cotula australis, Croton dioicus, 
Digitaria insularis, Dichondra sericea, Elytraria imbricata, Erysimum asperum, 
Euphorbia hirta, Heliotropium angiospermum, Hybanthus attenuatus, Ipomoea 
hederifolia Jaegeria hirta, Lasciasis ruscifolia, Merremia umbellata, Oplismenus 
compositus, Pennisetum purpureum, Pseudelephantopus spicatus, Pteridium 
aquilinum var. pubescens, Rhodosciadium toluscense, Setaria geniculata, 
Stellaria cuspidata y Solanum nigrescens. 

En el Centro solo se localizan 11 taxa: Artemisia ludoviciana ssp. mexicana, 
Datura discolor, Eragrostis barrelieri, Eragrostis lugens, Eruca sativa, Euphorbia 


1140 BRIT.ORG/SIDA 19(4) 


thymifolia, Lactuca graminifolia, Mollugo verticilata, Nothoscordum gracile, 
Sabazia humilis y Tagetes micrantha. 

Los 24 taxa del Sur son: Agrostis semiverticillata, Boerhavia coccinea, 
Cenchrus myosuroides, Cenchrus pilosus, Cyperus odoratus, Chenopodium 
mexicanum, Chenopodium murale, Descurainia pinnata, Diastatea micrantha, 
Eleocharis acicularis, Erechtites hieracifolia var. cacaloides, Euphorbia 
albomarginata, Euphorbia glomerifera, Gnaphalium americanum, Ipomoea in- 
dica, Lactuca serriola, Malvastrum americanum, Oenothera pubescens, 
Oenothera tetraptera, Petunia parviflora, Polygonum mexicanum, Polygonum 
monspeliensis, Rumex acetosella y Simsia foetida var. foetida. 

En la tabla 2, se pueden observar los datos cuantitativos de los taxa con 
distribuci6n restringida por Distrito agricola: en el Norte, estos corresponden 
al 7.49% del total, en el Centro al 3.17% y en el Sur al 6.91%. Solo cerca del 18% de 
los taxa registrados para el estado presentan cierta exclusividad regional. Fl 82% 
(286 taxa) restante, se encuentra distribuido en estos 3 Distritos de manera 
indistinta. 

Por lo que se refiere al lugar de procedencia de las especies, el mayor numero 
de taxa son nativos (125) y entre los introducidos se registraron en el estado los 
siguientes: 5/7 norteamericanos, 57 de América tropical, 29 europeos, 23 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 1141 


Tasta 2..Numero y porcentaje de especies arvenses y ruderales con distribucion restringida por 
Distritos agricolas y su relacidn con el numero total de especies malezoides para el estado de 
Querétaro. 


Distrito Agricola Relacién de numero taxa por Distrito Porcentaje 
Agricola/numero de taxa totales 


Norte 26/347 7.49 

Centro 11/347 Ble 
24/347 6.91 

Total 61/347 17.57 


sudamericanos, 20 euroasiaticos, 16 asiaticos, 8 caribenos, 8 africanos, 2 de la 
region de Oceania y 2 centroamericanos (Fig. 3). 

El porcentaje de especies de nativas es importante (36%), aunque la suma 
de las especies introducidas es mayor (ca. 64%), lo que contrasta con el hecho 
que la maleza autdéctona es la que prevalece en el pais (Rzedowski 1978, 1991) 

Por la ubicacion geografica del estado de Querétaro, una pequena 
proporcion de su superficie se encuentra dentro de la region arida y templada 
de la llamada Altiplanicie Mexicana 0 Mesa del Centro, razon por la cual se 
favorece la distribucién de elementos de origen norteamericano y la presencia 
de endemismos, algunos de ellos en la categoria tanto de arvense como ruderal. 

Las principales familias con taxa endémicos de México y presentes para 
este estado son: Compositae (45 spp.), Gramineae (14 spp.), Solanaceae (7 spp.), 
Euphorbiaceae (5 spp.), Caryophyllaceae (4 spp.), Cucurbitaceae (4 spp.); 
Labiatae (4 spp.), Leguminosae (4 spp.), Onagraceae (4 spp.), Chenopodiaceae 
(3 spp.), Umbelliferae (3 spp.) y Commelinaceae @2 spp.) (Tabla 1). En funcion 
del numero de elementos floristicos endémicos de la familia Compositae para 
México, Turner y Nesom (1998) han senalado que tienen su origen in situ dentro 
de la Altiplanicie Mexicana y el Eje Neovolcanico Transversal. 

El mayor numero de taxa introducidos registrados para Queretaro, quiza 
se explica por la ubicacion geografica del estado, paso obligado hacia las regiones 
norte y sur, asi como del Pacifico al centro del pais, su presencia probablemente 
se puede remontar al intercambio comercial de granos y especias con Asia, 
iniciado en 1565, por la ruta naviera entre los Puertos de Acapulco (estado de 
Guerrero) y Manila, Filipinas (King 1966). Otro factor que pudo influir en estos 
eventos, se relaciona con la intensa actividad minera existente durante el 
Virreinato espanol (siglos XVII y XVIII), ya que favorecio los asentamientos 
humanos, previamente iniciados por los chichimecas y otomies y como 
consecuencia de esta nueva actividad aumentaron los espacios para el cultivo, 
para la actividad ganadera y para las diversas actividades antropicas que en el 
estado se han realizado a lo largo de algunos cientos de anos, por lo que varios 


1142 BRIT.ORG/SIDA 19(4) 


Fic.3.R tacion del nu d ies de la fl {eral iond Jenci para el estado de 
f 


Querétaro. 


taxa de la familia Gramineae pudieron ingresar y adquirir la posicion de 
forrajeras sil vestres (Cynodon dactylon, Lycurus phleoides) y mas tarde el caracter 
de cultivadas como Sorghum bicolor y Sorghum halepense (ésta tltima 
convertida en una agresiva arvense), motivo por el que esta familia, contiene un 
buen porcentaje de elementos introducidos. Por otra parte, la apertura de la 
carretera panamericana en el periodo postrevolucionario, asi como la 
construccion de caminos en los Valles Centrales de la entidad, permitierona su 
vez el tratico continuo y la dispersion de varias especies hacia las regiones norte 
y occidente, asi como en sentido inverso. Este pudiera ser el caso para las familias 
Cyperaceae, Convolvulaceae, Malvaceae y Polygonaceae, que aportan un 
numero de taxa de otras regiones. A su vez se favorecio la llegada de elementos 
tropicales y caribenos, con posible via de la Planicie Costera del Golfo de México, 
hacia la parte norte del estado (municipios de Landa de Matamoros y Jalpan). 
El éxito de las especies malezoides provenientes de otras areas como la 
curoasiatica,en comparacion con las del Nuevo Mundo, se explica por la estrecha 
similitud ecologica y agricola, existente entre ambas regiones (tierras de clima 
templado y sin barreras orograficas notables). Esto favorecié la introduccién y 
naturalizacion de una buena cantidad de especies con caracteristicas 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 1143 


particulares que les permitieran adaptarse perfect te al arribar a las zonas 
templadas de América, donde ya existia una agricultura t d 

Otro de los multiples factores que favorecieron el desarrollo de especies 
exOticas en nuestro pais, fue el manejo particular del suelo agricola con tipicas 
practicas del Viejo Mundo, permitiendo el establecimiento de plantas ruderales 
de esa region, en los distintos nichos ecolégicos donde se implant6 un nuevo 
cultivo (sin competencia alguna), dando lugar a que alg ruderales exoticas 
presentaran ciclos de vida sincronicos con las actividades agricolas, 
comport tandose entonces como arvenses. 

sQue se prevee en el futuro? El crecimiento y la industrializacion de las zo- 
nas agricolas centro y sur del estado, pueden modificar la presencia de las 
especies actualmente consideradas como malezas y favorecer la introduccion 
de otras que tengan las caracteristicas adecuadas para adaptarse al trafico y 
perturbacion antropica, circunstancias que en la region norte nose ha reflejado 
y donde seguramente la representacion de este tipo de plantas seguira sin cambio 
considerable o bien muchas de estas ruderales podrian empezar a comportarse 
como especies arvenses. 


AGRADECIMIENTOS 


Los autores desean agradecer a Ma. Luisa Juarez, Ma. del Pilar Marcial y a Nor- 
man Regalado por el apoyo técnico brindado para la realizacion de esta 
investigacion. 


REFERENCIAS 


Acunois, M.O. y C. Ropricuez J. 1978. Maleza del algodonero en la Comarca Lagunera 
(Descripcion y Distribucidn). Folleto miscelaneo No. 40. INIA. SARH. México. 

Arcueltes, E., R. FernAndez N.y S. ZAmupio. 1991. Listado floristico preliminar del estado de 
Querétaro. Flora del Bajfo y de regiones adyacentes. Fasciculo complementario II. Insti- 
tuto de Ecologia, A.C. Centro Regional del Bajfo, Patzcuaro, Michoacan, México. 

Baker, H.G. 1974. The evolution of weeds. Ann. Rev. Ecol. Syst. 5:1-24. 

Barrera, B.N. 1996. Los origenes de la ganaderia en México. Ciencias. Rev. Difusion. Fac. Ci. 
Univ. Nac. Auton. Mexico 44:14—-27. 

Boecke, O. 1986. Plantas vasculares de la Argentina. Edit. Hemisferio Sur. Buenos Aires, Ar- 
gentina. 

Bye,R.1993.The role of humans in the diversification of plants in Mexico.In:Ramamoorthy, 
T.P, R. Bye, A. Lot y J. Fa, eds. Biological diversity of Mexico. Origins and distribution. 
Oxford University Press. New York. Pp. 707-731. 

CHALLENGER, A. 1998. Utilizacidn y conservacion de los Ecosistemas Terrestres de México. 
Pasado, Presente y Futuro. Comision para el Conocimiento y Uso de la Biodiversidad 
(CONABIO), Instituto de Biologia, Universidad Nacional Auténoma de México (UNAM) 
y Agrupacion Sierra Madre, S.C., México, D.F. Pp. 119-156. 


1144 BRIT.ORG/SIDA 19(4) 


Davita, PA. y J. SANCHEZ K. 1996. La importancia de las gramineas como forraje en México. 
Ciencias. Rev. Difusion, Fac. Ci. Univ. Nac. Auton. México 44:32-34. 

Espinosa, G.F.J. y J. SARUKHAN. 1997. Manual de malezas del Valle de México. Claves, 
descripciones e ilustraciones. Universidad Nacional Autonoma de México (UNAM) y 
Fondo de Cultura Econdmica (FCE), México, D.F. 

FERNANDEZ, N.R., C. Ropricuez J. y J.A. COLMENERO R. 1993. Malezas del NE del Estado Querétaro. 
Resumenes. XIl Congreso Mexicano de Botanica. 3-8 de Octubre, 1993. Mérida, 
Yuc.Mexico. 

Fryxett, PA. 1988. Malvaceae of Mexico. Syst. Bot. Monogr. 25:1-522. 

Hom, G.L., D.L. Pluckwett, J.V. PANCHO y J.P. Hersercer. 1977. The word’s worst weeds. Distribu- 
tion and Biology. University Press of Hawaii, Honolulu. 

Hout, E.B. 1970. Desarrollo General, Agropecuario y Forestal del estado de Querétaro en el 
periodo 1930-1960. Bol. Inst. Geografia. Univ. Nac. Auton. México 3:134-193. 

INEGI, 1986. Sintesis Geografica de estado de Querétaro, Nomenclator y Anexo Cartogrdfico. 
S:P.P. México. D.F 

Kina, L.J. 1966.Weeds of the world. Intersciense Publ. Inc. New York. 

Lira, R.,C. Rooricuez J., J.L. Atvarano, |. Ropricuez, J. CASTREJON y A. Domincuez M. 1998. Diversidad 
e importancia de la Familia Cucurbitaceae en México. Acta Bot. Mexicana. 42:43-77. 

Marzocca, A. 1976. Manual de malezas. Edit. Hemisferio Sur, Buenos Aires, Argentina. 

McDonato, A. 1991. Origin and diversity of Mexican Convolvulaceae. Anales Inst. Biol. Univ. 
Nac. Auton. México. Ser. Bot. 62:65-8? 

Nava, T.M.M. 1991. Estudio ecolégico de las plantas arvenses a lo largo de la carretera 
Naucalpan-Jiquipilco (México).Tésis de Bidlogo. Escuela Nacional de Ciencias Bioldgicas, 
Instituto Politénico Nacional (IPN), México, D.F. 

Ramirez, V.B.R. 1995. La regién en su diferencia: Los Valles Centrales de Querétaro 1940- 
1990. Edit. Red Nacional de Investigacién Urbana, Universidad Autonoma 
Metropolitana-Unidad Xochimilco (UAM-X) y Universidad Auténoma de Querétaro 
(UAQ), México. 

Rapoport, E.H.,M.E. Diaz B.y ILR. Lopez M. 1983. Aspectos de la ecologia urbana en la Ciudad 
de Mexico. Flora de calles y baldios. Publ. Il, Inst. de Ecologia. Edit. Limusa, México, D.F. 

Ropricuez, J.C. 1967. Estudio ecoldgico de las malas hierbas del Valle de Toluca, México. 
Tesis de Licenciatura. Facultad de Ciencias, Universidad Nacional Autonoma de México 
(UNAM), México, D-F. 

Rotuns, R.C. 1981.Weeds of the Cruciferae (Brassicaceae) in North America. J. Arnold Ar- 
bor.62:517-540. 

Rzepowski, J. 1978. Vegetacion de México. Edit. Limusa, México. Pp.67-71. 

Rzevowski, J. 1991. Diversidad y origen de la flora fanerogamica de México. Acta Bot. Mexi- 
cana 14:3-21. 

Rzepowsk\, J. 1993. El papel de la familia Compositae en la flora sinantrdpica de México. 
Fragm. Florist. Geobot. Suppl. 2(1):123-138. 

RzeDowski, J. y G.C. be Rzebowski. 1957. Notas sobre la flora y la vegetacion del estado de San 


— 


COLMENERO ET AL., MALEZAS DE QUERETARO, MEXICO 1145 


Luis Potosi.V.La vegetacion a lo largo de la carretera San Luis Potosi-Rio Verde. Acta Ci. 
Potos. 1:/-68 

Rzevowski, J.y G.C. pe RzeDowsk. 1990. Nota sobre el elemento africano de la Flora adventi- 
cia de México. Acta Bot. Mexicana 12:21-24. 

Rzeoowski, J.y G.C. pe Rzepowski. 1993. Datos sobre la dinamica de la flora fanerogamica del 
Valle de México con énfasis en especies nativas raras, en peligro de extincion y 
aparentemente extintas. Acta Bot. Mexicana 25:81-104. 

SuArez, R.G. y V. Serrano C. 1997. Principales malezas (arvenses) en el campo queretano. 
Herbario Querétaro No.4. Serie Cientifica. Instituto Tecnoldgico de Estudios Superiores 
de Monterry (ITESM) y Universidad Auténoma de Querétaro. Mexico 

Turner, B.L.y G.L. Nesom. 1998. Biogeografia, diversidad y situacion de peligro o amenaza 
de Asteraceae de México. En: Diversidad Biolégica de México. Origenes y Distribucion. 
Comp.T.P. Ramamoorthy, R.Bye, A. Lot y J.Fa. Instituto de Biologia, Universidad Nacional 
Autonoma de México (UNAM), México, D.F. Pp. 545-561. 

Vigrans, H. 1996. Notes on neophytes 2. New records for Asteraceae from the center of 
Mexico. Phytologia 81:369-381. 

Vigrans, H. 1998a. Native maize field weed communities in south-central Mexico. Weed 
Res. 38:153-166. 

Visrans, H. 1998b. Urban weeds of Mexico city. Floristic composition and important fami- 
lies. Anales Inst. Biol. Univ. Nac. Auton. México, Ser. Bot. 69:37-69. 

VILLARREAL, O.J.A. 1983. Malezas de Buenavista Coahuila. Universidad Autonoma Agraria, 
Antonio Narro. Buenavista, Saltillo, Coahuila, México. 

VILLASENOR, R.J.L.y F.J.Esrinosa G. 1998. Catalogo de malezas de México. Universidad Nacional 
Autonoma de México (UNAM), Consejo Nacional Consultivo Fitosanitario (CNCF) y 
Fondo de Cultura Econémica (CFE), México, D-F. 

ViLLEGAS, DE G.M. 1970. Estudio floristico y ecoldgico de las plantas arvenses de la parte 
meridional de la Cuenca de México. Anales. Esc. Nac. Ci. Biol. 18(1-4):1 7-89. 

Zamupio, S.R.,J. RZEDOwskI, E. CARRANZA G.y G.C. De RzEDowski. 1992. La vegetacion en el estado 
de Querétaro, CONCYTEQ. Instituto de Ecologia, Centro Regional del Bajio, Patzcuaro, 
Michoacan, México. 

Zereva, AS. 1996. Especies de malezas en maiz y sorgo en el centro-sur de Queretaro. 
INIFAP CONCYTEQ y Monsanto Comercial S.A. de CV., Querétaro, Queretaro, México 


1146 BRIT.ORG/SIDA 19(4) 


Book Notices 
Piers Vitessky. 2001. Shamanism. (ISBN 0-8061-3328-7, pbk.). University of Oklahoma 
Press, 4100 28th Avenue N.W, Norman, OK 73069-8218, U.S.A. (Orders: 
wwwou.edu/oupress, 405-364-5978 fax). $12.95, pbk, 128 pp., 250 color and 
b&w illustrations, 25 maps, 51/2" x 8" 


Table of Contents. —Introduction 
The Shamanic Worldview 
2) Regional Traditions 
3) Becoming a Shaman 
4) Shamans and Clients 
5) neue: Shamans 
) Documentary Refer 
siBhicersphy, ae iat ais and Acknowledgements 


letailed by region sur- 


This isa fine introductory text on Shamanism worldwide. Inch 2 
vey of shamanism with full color sete cover ane beth spina and psychological ae and the 
relevance of shamanism to West mallangles 
and ee ines. From the back cover: “The shaman occupies a key role as a healer, mediating be- 
the spirits, and is a potent figure in alterne medi- 


J 


tween the world of the living and the world of 
cine. Shamanism, a richly illustrated guide, looks at both historic and present- day manifestations.” | 


Id recommend this book for anthropology classes in Religion and/or Shamanism. It is reason- 
well-written, and Se —Kevin D.Janni, Botanical Research Institute of Texas, 
hy 


WOU 

ably priced, 

Fort Worth, TX 76102-4060, U.S.A, kj 

B. LERoy Davipson. 2001. Lewisias. 2000.(ISBN 0-88192-447-4, hbk.). Timber Press, 
133 SW Second Ave., Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: 
www.timberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). 
$34.95, hbk. 238 pp., 68 color photos, 20 color plates, 13 line icine 


RA 


This isa book a Hower S and prctiye onesat that - s me genus | 
Lewis (1774-18 Portulacaceae.“A genus of succulent asta 
numbering appr vosimae Ean speties Lewisia has been adopted asa favorite by rock gardener 

ant to have an impressive collection of lewisia photos in habitats — 


throughout the world.” If you w 
gardens an nd text to 80 along with it then this is book. “Lewisias is that rare book that seamlessly 
e a holistic picture of a botanical treasure: it is 


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artist Micheal Moshier are included.—Barney Lipscomb, Botanical Research i ome of Texas, Fort 


Worth, TX 76102-4060, U.S.A, barney@britorg. 


SIDA 19(4): 1146. 2001 


VINES OF A TEMPERATE STATE: STILL UNDERCOLLECTED? 


Alexander Krings 
Herbarium, Department of Bota 
North Carolina State ree 
Raleigh, NC 27695-7612, U.S.A. 
Alexander_Krings@ncsu.edu 


ABSTRACT 


Over the Bast decades, there has been a steady effort to contribute to our gow cage ol herbaceous 
an dw oody vines (lianas) a habit previoushy largely neg pected nm pote 


] i pee ] a] 


ies. This study sought to evaluate to 

ing of lianas and vines in a state hosting the highest manic a eietiay Cees per squar 

mile in the southeastern United States—North Carolina ly r floristic ae 

standing, baseline and updated county species lists of ane and vines were penee based on 

literature review and herbarium survey. To compare liana and vine species richness by ecophysi- 
ological province, an index of species similarity was calculated. The pleemone and Coastal Plain 

host the largest absolute number of vine taxa in Nee Carolina, as we gest mean one 

a 


L: 


per county. The higher Piedmont mean richne 
lapping distributions from adjacent provinces. ee reasons for the poaineulery high species 
richness of individual counties and the significant variation in county to county species richness 
remain unclear. Macro-climate, as well as diversity in topography and associated microclimate may 
pe factors. However, the chee most Species men counties also happen to host the three largest her- 

f historically larger eee a Other 
non-biological factors potentially involved include distance to site and accessibility 


RESUMEN 


En las décadas  pasadas, ha habia un mesttier29 constante para contribute a nuestro Sonocimiente o de 


] 
lenosas 


Logi Este aa peti evaluar e en que medida se ha Digetesade! en mice conoci- 
ees 

por mill drada del de Estados l inidos Carolina del Norte. Para analie ar el estado de nuestro 
to floristi desarrollaron li por condado, basicos y puestos al dia, 

de lianas basados e en revisiones ioe y de herbarios, pare aa la anaes de especies de 
lianas por p ecofisioldgica, se cal ie de monte y la 
llanura costera tienen el aaer numero soso 5 ue de eons del Norte, asi como la mayor 
riqueza media por condado la posicién 


poe ence ya las distribuciones solapadas ee = provincias adyacentes sin embargo, er razones 
ara 


P 
de condado a condado no o quedan ¢ claras. El macroclima, asi como ate diverscad en aa y el 
Sin been Date l I 


g ars : pe een (errr M4 y eee Sy | - yee ee Foam | 


también 
de colecta historicamente mas amplios. Otros Betores no biolégicos potencialmente implicados 
incluyen la distancia al lugar y la accesibilidad 


An important aspect of recent studies of vines has been the exploration of the 
underlying causes determining and limiting the distribution of species and 


SIDA 19(4): 1147 — 1155. 2001 


1148 BRIT.ORG/ SIDA 19(4) 


individuals (Bell et al. 1988; Molina-Freaner & Tinoco-Ojanguren 1997; 
Castellanos et al. 1999). Understanding factors controlling vine distributions is 
of interest in understanding broader patterns of vine species richness. Several 
factors, such as soil moisture (Bell et al. 1988; Collins & Wein 1993), availability 
of small diameter supports (Putz & Chai 1987), distribution and spatial arrange- 
ment of supports (Putz & Chai 1997), as well as preferences for light microenvi- 
ronments within host canopies (Castellanos et al. 1999), have been found im- 
portant, but further study is warranted before a synthesis can be obtained. 
Although lianas and vines have been previously largely neglected in both bo- 
tanical and ecological studies Jacobs 1976; Putz 1984; Gentry 1991; Collins & 
Wein 1993), there has been a steady effort over the past decades to contribute to 
our floristic knowledge base of this important group. Gentry (1991, 1995) re- 
viewed the floristics of lianas as determined froma series of 0.1 ha plots mostly 
located in Neotropical lowlands or Andean montane sites. Work by Grubb et 
al. (1963), Putz (1983, 1984), Collins and Wein (1993), Burnham (1997), and Krings 
(2000a, 2000b, 2001), among others (see Putz & Mooney 1991), has also con- 
tributed to our understanding of New World lianas and vines. This study sought 
to evaluate to what extent progress has been made in our floristic understand- 
ing of lianas and vines in temperate North Carolina. With the highest number 
of herbarium specimens per square mile in the southeastern United States (Funk 
& Morin 2000), North Carolina presents an interesting situation for evaluating 
the status of collections of this historically undercollected group. 


METHODS 


To analyze the state of our floristic understanding, baseline county species lists 
of herbaceous and woody vines [hereafter collectively “vines”] were developed 
based on Radford et al. (1968). Separate, updated species lists were then devel- 
oped based on literature review (Kessler 1956; Gupton 1960; Morgan 1962; Sears 
1966; Blair 1967; Tucker 1967; Bruton 1968; Hartshorn 1968; Sawyer 1968; Michael 
1969; Pittillo et al. 1969; Pultorak 1969; Wells 1970; Jones 1971; Pittillo et al. 1972; 
Jones 1973; Taggart 1973; Taylor 1974; Racine & Hardin 1975; Smith 1977; Lacey 
1979; Corda 1982; Skean 1982; Rohrer 1983; Sieren 1983; Mayes 1984; Bradshaw 
1987; Pittillo & Brown 1988; Matthews & Mellichamp 1989; Palmer 1990: Ingle 
1993; Floyd 1997; Strickland 2000), as well as herbarium searches at NCSC and 
NCU. All climbing, woody and herbaceous taxa known from North Carolina 
are included. Prostrate, mat-forming taxa, such as Mitchella repens L. 
(Rubiaceae), are excluded. 

To compare vine species richness by ecophysiological province, an index 
of species similarity was calculated based ona modification of Sorensen (1948). 
As area has been shown to significantly influence species richness (White et 
al. 1984; Kohn & Walsh 1994), the Sorensen index is most precise when two 
communities of the exact same size are compared (e.g., using fixed area 


KRINGS, VINES OF A TEMPERATE STATE 1149 


transects). When communities of different sizes are compared a modified in- 

dex can be calculated: 

(Eq. 1) Index of similarity (area-weighted) = C/log area 4 + C/log area p * 100, 
A/log area a + B/log area g 


where C is the number of species shared between two respective sites, and A 
and B are the numbers of species present at each respective site. Although not 
as precise, non-modified Sorensen indices have been used between communi- 
ties of different sizes (see Sklena¥ & Jorgensen 1999) but should be interpreted 
cautiously for revealing broad trends in similarity. 


RESULTS 


Including escaped and persisting taxa, the North Carolina vine flora currently 
comprises 155 species in 31 families. 

Over the period of 1968-2001, the largest percentage of new North Caro- 
lina county vine species presence records came from the Mountain and transi- 
tional Mountain-Piedmont counties (Table 1). Except for Wilkes County (no 
change), all Mountain counties showed an increase in the number of vines 
known from 1968 to 2001. County records of Piedmont, Piedmont-Coastal Plain 
transition, and Coastal Plain counties, increased only a third or less (by per- 
cent) of Mountain record increases (Table 1). Nine of the 31 Piedmont counties 
showed negative changes in the number of vines known over the period. Nine 
Coastal Plain counties also showed negative change over the period. Negative 
change resulted from changes in taxonomy or resolution of formerly mis-ap- 
plied names. No change in the number of species known occurred in 20 of the 
100 counties of North Carolina. Thus, 38% of all counties displayed either a nega- 
tive change or no change in the number of species of vines known in 2001 vs. 
1968. 

Based on the updated 2001 data, Piedmont counties host a significantly 
higher mean number of species (x = 40.7) than do Mountain counties (t = 5.140, 
p< 0.0005) or Coastal Plain counties (t = 4.283, p < 0.0005). Coastal Plain coun- 
ties also host a significantly higher mean number of species (x = 37.) than 
Mountain counties (x = 35.5; t = 1.785, p < 0.025). The Piedmont and Coastal 
Plain host the largest absolute number of vine taxa in North Carolina (Table 2), 
as well as the largest mean richness per county. 


DISCUSSION 


The higher Piedmont mean vine species richness may be due in part to geo- 
graphic position. Piedmont counties host a larger percentage of vines overlap- 
ping in distribution from adjacent provinces than Mountain or Coastal Plain 
counties. However, reasons for the particularly high richness of several, scat- 
tered counties remain unclear (Fig. 1). 


1150 BRIT.ORG/ SIDA 19(4) 


Taste 1. Changes in liana and vine species totals known per county in North Carolina 1968-2001. 


Avg.no.ofspp. Avg.no.ofspp.per Changeinspp. Percent change in 


per county county (2001 nos. 1968- spp. nos. 1968- 
(1968)* update) 2001 2001 
Mountain counties — 30.7 35.5 +48 + 15.6 
Mountain-Piedmont 
transition counties (N=5)** 29.2 35.0 +58 19.9 
Piedmont counties (N=31) 38.6 40.7 +21] +54 
Piedmont-Coastal Plain 
transition counties(N=3)*** 43.0 45.0 + 2.0 +47 
Coastal Plain counties (N=41) 35.8 37.1 + 1.3 + 3.7 


Based on Radford et al. (1968). 
** Alexander, Burke, Polk, Rutherford. and Surry counties. 
™* Lee Moore, and Richmond counties. 


Taste 2. Matrix of Sorensen's indices of similarity (area-weighted) for the vine flora of North Carolina 
nd vine species and, in p 


> physiographic province. Diagonals indicate total number of liana an 
ntheses, number and percentage of included, introduced liana and vine species. 


Mountains Piedmont Coastal Plain 
Mountains 89 (20; 22.5%) - 
Piedmont 75.16 111 (35; 31.5%) - 
Coastal Plain 65.03 82.13 118 (36; 30.5%) 


Note:Transitional counties of overlapping physiographic provinces are not considered in the table. 
These counties are Alexander, Burke, Lee, Moore, Polk, Richmond, Rutherford, and Surry. 


Itis possible that richness in some counties, particularly in the eastern Pied- 
mont, may be linked to macro-climate, as well as diversity in topography and 
associated microclimate. For example, Hardin and Cooper (1967) suggested that 
eastern Piedmont communities (especially in Durham, Orange,and Wake coun- 
ties) are particularly rich in plant taxa (of all habits) due toa significant moun- 
tain and mountain disjunct element in their flora. Citing growing season and 
precipitation data, Hardin and Cooper (1967) provide indirect support for their 
hypothesis that this component of the flora is largely remnant from Pleistocene 
times and persists in cooler microsites. Topographically more highly dissected 
than the western Piedmont, the eastern Piedmont may provide a higher num- 
ber of sites amenable to the occurrence of montane elements (Hardin & Coo- 
per 1967), although further studies are needed. 

Although Hardin and Cooper (1967) discount montane disjunction pat- 
terns as an artifact of sampling, the same may not yet be defendable for the dis- 
tribution of vine taxa. As Connor and Simberloff (1978) have explored, in some 


KRINGS, VINES OF A TEMPERATE STATE 1151 


P may / r J 
NCSC and NCU). Counti ith 50 i haded 


- 
> 
> 
> 
im 

al 


instances collection effort may be a greater indicator of species richness and 
similarity among communities than biological factors. Based on our present 
knowledge, county to county vine species richness vari generously within 
North Carolina provinces (Fig. 1) that an analysis of variance (ANOVA) found 
no significant difference by province in county species richness (F 0.05: 2, 91 = 
1.355 <F crit 3.098; p > 0.26). Such variation is unexpected based on distinct dif- 
ferences in the climates and soils of the state’s three provinces (Robinson 1979) 
and suggests non-biological causal factors. Potentially the result of historically 
larger collecting programs, the three most species rich counties also happen to 
host the three largest herbaria in the state (DUKE, NCU, and NCSC) (Fig. 1). 
Distance to the study site from the residence of researchers could likely influ- 
ence the number of visits and hence the completeness of inventories. Accessi- 
bility and topography could also play a limiting role. Combined with less than 
5.5% growth in county records for 75% of the potentially most speciose North 
Carolina counties (e., Piedmont and Coastal Plain counties) over the past thirty- 
three years (Table 1), the encountered county to county species richness varia- 
tion indicates that, at least geographically, vines still remain largely 
undercollected for much of North Carolina. 


ACKNOWLEDGMENTS 


I thank the ILL department of the North Carolina State University libraries for 
help with obtaining materials not owned by the University and NCU for access 
to its collections. 


REFERENCES 


Bett, DJ., |.N.Forseth, and A.H. Teramura. 1988. Field water relations of three temperate vines. 
Oecologia 74:537-545. 


1152 BRIT.ORG/ SIDA 19(4) 


Biair, A.E. 1967. Vascular flora of Beaufort County, North Carolina. M.S. Thesis, North Caro- 
lina State University, Raleigh 

BrapsHaw, TL. 1987. Floristic survey and vegetational analysis of Lost Cove, Avery County, 
North Carolina. M.S. Thesis, North Carolina State University, Raleigh. 

Bruton, C.V. 1968. Floristic survey and vegetational analysis of Cliffs of the Neuse State 
Park in North Carolina. M.S. Thesis, North Carolina State University, Raleigh 

BurNHam, R.J.1997.Stand characteristics and leaf litter species composition of a dry forest 
hectare in Santa Rosa National Park, Costa Rica. Biotropica 29:384-395. 

CASTELLANOS, A.E.,C. TINocO-OQJANGUREN, aNd F. MOLINA-FREANER. 1999. Microenvironmental hetero- 
geneity and space utilization by desert vines within their host trees. Ann. Bot. 84: 
145-153 

Cotuns, B.S. and G.R. Wein. 1993. Understory vines: Distribution and relation to environ- 
ment on a southern mixed hardwood site. Bull. Torrey Bot. Club 120:38-44. 

Connor, E.F. and D. Simpertorr. 1978. Species number and compositional similarity of the 
Galapagos flora and avifauna. Ecol. Monogr. 48:219-248. 

Corba, S.L. 1982. The flora and community types of Goose Creek State Park, North Caro- 
lina. M.S. Thesis, North Carolina State University, Raleigh 

Duncan, W.H. 1975.Woody vines of the southeastern United States. The University of Geor- 
gia Press, Athens. 

FLoyp, J.W. 1997.Vascular flora of Medoc Mountain State Park, Halifax County, North Caro- 
lina. M.S. Thesis, North Carolina State University, Raleigh 

Funk, V.A.and N. Morin. 2000. A survey of the herbaria of the southeast United States. Sida, 
Bot. Misc. 18:35-52. 

Gentry, A.H. 1991. The distribution and evolution of climbing plants. In: FE. Putz and H.A. 
Mooney, eds. The biology of vines, Cambridge University Press, New York. Pp. 3-42. 
GentRY, A.H. 1995. Patterns of diversity and floristic Eom Osho: in eet les montane 

forests. In:S.P. Churchill, H. Balslev, E. Forero, and J.L.Luteyn, eds. Biodiversity and conserva- 


tion of neotropical montane forests, New York Botanical See York. Pp. 103-126. 

Gruss, A.J.,J.R. Ltoyo, T.D. PENNINGTON, and T.C. WHitmore. 1963.A comparison of montane and 
lowland rain forest in Ecuador, |: The forest structure, physiognomy, and floristics. J. 
Ecology 51:567-601. 

GupTon, O.W. 1960. A vascular flora of the North Carolina Sandhills Wildlife Management 
Area. M.A. Thesis, University of North Carolina, Chapel Hill. 

Haron, J.W. and A.W. Cooper. 1967. Mountain disjuncts in the eastern Piedmont of North 
Carolina. J. Elisha Mitchell Sci. Soc. 83:139-150. 

HarTSHORN, G.S. 1968. Vegetation patterns in southern Beaufort County, North Carolina. 
M.S. Thesis, North Carolina State University, Raleigh 

Hecarty, E.E. and G. Casaite. 1991. Distribution of vine communities. In: FE. Putz and H.A. 
Mooney, eds. The biology of vines, Cambridge University Press, New York. Pp. 313-335. 

INGLE, R.F. 1993. Floristic survey and biogeographical analysis of the granite outcrops of 
eastern North Carolina. M.S. Thesis, North Carolina State University, Raleigh 

Jacoss, M. 1976.The study of lianas. Flora Malesiana Bull. 29:2610-2618. 


KRINGS, VINES OF A TEMPERATE STATE 1153 


Jones, 5.1. 1971. Floristic survey and vegetational analysis of Yates Pond Biological Area in 
Wake County, North Carolina. M.S. Thesis, North Carolina State University, Raleigh 

Jones, T.H. 1973. Raven Rock State Park, Harnett County, North Carolina. M.S. Thesis, North 
Carolina State University, Raleigh 

Kartesz, J.T. 1994.A synonymized checklist of the vascular flora of the United States, Canada, 
and Greenland. Timber Press, Portland. 

Kesster, P-A. 1956. A floristic study of the Triassic sediments, Deep River Coal Field, North 
Carolina. M.A. Thesis, University of North Carolina at Chapel Hill. 

KoHN, D.D and D.M.Watsu. 1994. Plant species richness — the effect of island size and habi- 
tat diversity. J. Ecology 82:367-377. 

Krines, A. 2000a.Floristics and ecology of Mesoamerican montane climber communities: 
Monteverde, Costa Rica. Selbyana 21:156-164. 

Krincs, A. 2000b. Phytogeographical characterization of the vine floras of the Sonoran 
and Chihuahuan Deserts. J. Biogeogr. 27 (6):1311-1319. 

Krincs, A. 2001. Vines of two Texan semi-arid desert communities: Floristic composition 
and ecological notes. Southw. Naturalist 46:400-404. 

Lacey, V.H. 1979.A floristic survey of Phoenix Mountain, Ashe County, North Carolina. M.A. 
Thesis, Appalachian State University. 

MattHews, J.F.and T.L.MetuicHamp. 1989. Additional records to the vascular flora of the Caro- 
linas and a selected bibliography of floristic studies, 1964-1987. J. Elisha Mitchell Sci. 
Soc. 105 (1):34-53. 

Mayes, C.H. 1984. The flora of Smith Island, Brunswick County, North Carolina M.S. Thesis, 
University of North Carolina at Wilmington. 

Michael, G.L. 1969. Vascular flora of Bullhead Mountain, Alleghany County, North Carolina. 
M.A. Thesis, University of North Carolina, Chapel Hill. 

Motina-FRreaner, F.and C. Tinoco-OJancuren. 1997.Vines of a desert plant community in Cen- 
tral Sonora, Mexico. Biotropica 29:46-56 

Morean, J.D. 1962. A vascular flora of Morrow Mountain State Park, Stanly County, North 
Carolina. M.A. Thesis, University of North Carolina, Chapel Hill. 

Nekota, J.C.and PS. Wuite. 1999. The distance decay of similarity in biogeography and ecol- 
ogy. J. Biogeogr. 26:867-878. 

Parmer, M.W. 1990.Vascular flora of the Duke Forest, North Carolina.Castanea 55 :229-244, 

Pittitto, J.D., J.H. Horton, and K.W. Greene. 1969. Additions to the vascular flora of the Caro- 
linas. |. J. Elisha Mitchell Sci. Soc. 85:18-22. 

Pirtito, J.D., JH. Horton, and K.E. Herman. 1972. Additions to the vascular flora of the Caroli- 
nas. Il. J. Elisha Mitchell Sci. Soc. 88:144—-152 

Pittito, J.D.and A.E. Brown. 1988. Additions to the vascular flora of the Carolinas. Ill. J. Elisha 
Mitchell Sci. Soc. 104:1-18. 

Puctorak, R.W. 1969. Vascular flora of northern Wake County, North Carolina. M.S. Thesis, 
North Carolina State University, Raleigh. 

Putz, FE. 1983. Liana biomass in a'tierra firme’ forest in the Rio Negro basin, Venezuela. 
Biotropica 15:185-189. 


1154 BRIT.ORG/ SIDA 19(4) 


Putz, FE. 1984. The natural history of lianas on Barro Colorado Island, Panama. Ecology 
65:1713-1724. 

Putz, FE. and P. Cual. 1987. Ecological studies of lianas in Lambir National Park, Sarawak, 
Malaysia. Journal of Ecology 75:523-31. 

Putz, FE. and H.A. Mooney. 1991. The biology of vines. Cambridge University Press, New 
York, 

Racine, C.H. and J.W. Haroin. 1975. The vascular flora and vegetation in the Green River 
Gorge, North Carolina. Castanea 40:319-345, 

Raproro, A.E., H.E. AHtes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
University of Chapel Hill Press, Chapel Hill. 

Rosinson, PJ. 1979. The climate of North Carolina. In: H. Lieth and E. Landolt, eds. Beitrage 
zur Kenntnis der Flora und Vegetation der Carolina, Ergebnisse der 16.|Internationalen 
Pflanzengeographischen Exkursion (IPE), 1978, durch die stiddstlichen Vereinigten 
Staaten, Verdff. Geobot. Inst. ETH, Stiftung Rubel Ztirich 68:5 1-60, 

Rourer, J.R. 1983. Vegetation pattern and rock type in the flora of Hanging Rock Area, 
North Carolina. Castanea 48:189-205. 

Sawyer, G.P. 1968. The vascular flora of William B. Umstead State Park, Wake County, North 
Carolina. M.A. Thesis, University of North Carolina, Chapel Hill. 

Sears, M.N. 1966. Floristic study of the limestone along Island Creek in Jones County, North 
Carolina. M.A. Thesis, University of North Carolina, Chapel Hill. 

SecHrest, C.G. and A.W. Cooper. 1970. An analysis of the vegetation and soils of upland 
hardwood stands in the Piedmont and Coastal Plain of Moore County, North Carolina. 
Castanea 35:26-57. 

SierREN, D.J. 1983. New records of vascular plants from Pender County, North Carolina. J. 
Elisha Mitchell Sci. Soc. 99:13-16. 

SKeAN, J.D, Jr. 1982. The vascular flora and plant community types of Durant Nature Park, 
Wake County, North Carolina. M.S. Thesis, North Carolina State University, Raleigh 

SKLENAR, P.and P.M. Jarcensen. 1999. Distribution patterns of paramo plants in Ecuador. J. 
Biogeography 26:681-691. 

Smith, E.L. 1977. Floristic survey and vegetational analysis of the South Mountains State 
Park, Burke County, North Carolina. M.S. Thesis, North Carolina State University, Raleigh 

Sorensen, T.1948.A method of establishing groups of equal amplitude in plant sociology 
based on similarity of species content and its application to analyses of the vegeta- 
tion on Danish commons. Det Kongel. Danske Vidensk. Selk. 5:1—34. 

STRICKLAND, E.L. 2000. A vegetational study of Ev-Henwood. M.S. Thesis, University of North 
Carolina at Wilmington. 

Swas, E.C. 1990. The flora and vegetation of White Pines Natural Area, Chatham County, 
North Carolina. M.S. Thesis, North Carolina State University, Raleigh 

Tacoart, J.B. 1973. Floristic survey and vegetational analysis of Stone Mountain State Park 
in North Carolina. M.S. Thesis, North Carolina State University, Raleigh 

Tayior, T.D.1974. A check list of vascular flora of Tablerock Mountain, Burke County, North 
Carolina. M.A. Thesis, Appalachian State University, Boone, NC. 


KRINGS, VINES OF A TEMPERATE STATE 1155 


Tucker, G.E. 1967. The vascular flora of Bluff Mountain, Ashe County, North Carolina. M.A. 
Thesis, University of North Carolina, Chapel Hill. 

Weuts, E.F. 1970.A vascular flora of the Uwharrie Wildlife Management Area, Montgomery 
County, North Carolina. M.A. Thesis, University of North Carolina, Chapel Hill. 

Wuire, PS., R.I. Miter, and G.S. Ramseur. 1984. The species-area relationship of the southern 
Appalachian high peaks:Vascular plant richness and rare plant distributions. Castanea 
49:47-61. 


1156 BRIT.ORG/SIDA 19(4) 


Book REVIEW 


Jit Nokes. 2001. How to Grow Native Plants in Texas and the Southwest. Second 
Edition. (ISBN 0-292-75573-2, hbk.). University of Texas Press, PO. Box 7819, 
Austin, TX 78713-7819, U.S.A. (Orders: http://www.utexas.edu/utpress, 512- 
471-4032).$60.00 hbk., 566 pp., 30 watercolor plates, line drawings by Kathryn 
Miller Brown, 6" x 9" 

This isa bible when it comes to propagating the native flora in the Southwest. This second edition is 

a beautiful ple of an author's passion for, knowledge of, and expertise in native plants. The first 

edition that was ublished in 1986 has acted as the foremost guide for nurserymen, scientists and 

laymen trying to grow species of plants that are native in the temperate climate of Texas and the 


Southwest. 


he first edition Was ] i | } | d Texas AGM. 
It contains an interesting analysis of he benefits for using native plants i in the | landscape verses the 
typical, exotic, ae maintenance nursery stock. When the first edition was published the native 


plant movement was just beginning to be po pues however, at that time there was little supporting 

literature on the ae ect. inthe 15 years past a very healthy amount of supporting literature has been 
J years pe y| y 

publis 


In the second edition, the author addresses the common notions that the general public has 
about native plants. Those being that they are shabby, ill behaved and second rate compared to the 
typical, traditional landscapes. She talks about the obliteration of the native flora in this country 
due to unknowing land developers. The author also describes how people think that a native land- 
scape is a “zero-scape.” This misconception has led to apathy in using native plants since people do 
not realize that a native plant garden can be lush, evergreen, and formal. A very good point made by 
the author is how important the basic design is to a more natural and less formal garden. Another 
idea offered is the importance of educating the public in choosing adapted, resource-efficient plants. 
She brings up the water concerns that have come about recently and how public places can be key 
factors in making the public aware about using water-efficient plants. 

e second edition has been significantly expanded as far as the number of species addressed, 
approximately 75 new along with the original 350 trees, shrubs and woody vines. Some of these spe- 
cies are from the subtropical southern part of Texas; a vegion apparently left out of the pi alae 
Other revised sections of the book are those that give the user precise 
storing seed, seed gecmnation, planting, aut propagation and transplanting. The bulk a the 
book consists of the propagation techniques for the genera and species. Both editions are well illus- 
trated but the new edition includes photographs showing working examples of nursery methods 
that are described in the text. One of the best ane of the work is the group of beautiful watercolor 


orn 


pate a Hustle the Snatactensace i 30 pe 

numerous experts some of whom 
ave past away since the first edition. The author says aie has acted as a bridge between the younger 

generation of horticulturists and the elder atone by Bringing rogetner cet MEOW IStge in bls 

publication. can techniques in horticulture that | 

availabil _— these plants from the nursery trade. 

- is not only a guide for greenhouse and nursery workers but also a field guide for 
anyone wanting to identify a plant that has been found in the wild that could be considered for orna- 
mental applications. This publication sets high standards for horticulturalists for many years to 
come.—Justin Allison, FNA oe Assistant, Botanical Research Institute of Texas, 509 Pecan 
Street, Fort Worth, TX 76102-4060, U 


SIDA 19(4): 1156. 2001 


A NEW SYNONYM FOR ERAGROSTIS PILGERI 
(POACEAE: ERAGROSTIDEAE) 


J. José Alegrfa Olivera and Arturo Granda Paucar 


Departamento de Biologia, Herbario (MOL) 
Universidad Nacional Agraria La Molina 
Apartado 456, Lima, PERU 
gmn@lycos.com and a.granda@lycos.com 


ABSTRACT 


The identity of Eragrostis ancashensis PM. Peterson, Refulio & Tovar, a Peruvian endemic, is discussed; 
the name is considered to be asynonym of Eragrostis pilgeri Fedde. Lectotypes are also designated for 
the names of some synonyms of E. pilgeri, whose holotypes at B were destroyed in 1943. Further 


te 
marks about the identity of two paratypes of E. ancashensis (Ferreyra 14577 and 14594 luded. 


RESUMEN 


de Peru. Fl b id inonimo de Erag pilgeri Fedde. Se desig r 
para los nombres de algunos sinonimos de E. pilgeri, cuyos holotipos depositados en B fueron 
destruidos en 1943. Se hace un comentario adicional sobre la identidad de dos paratipos de E. 
ancashensis (Ferreyra 14577 y 14594). 


| laidentidad de E ti hensis PM. Peterson, Refulio & Tovar, una especie endémica 
is pi i S i bién lectoti 


1pos 


Eragrostis Wolf (Chloridoideae: Eragrostideae) is a genus with about 350 species 
distributed in the tropical and subtropical regions throughout the world (Clay- 
ton & Renvoize 1986). In Peru, the genus is represented by 22 species according 
to Tovar’s (1993) treatment, although Davidse and Brako (1993) recognize the 
existence of 29 entities, which grow mainly in open fields, in dry and sandy 
soils of the Andes at altitudes between 0 and 4000 meters. Far from being well 
understood, Peruvian species of Eragrostis require a modern treatment making 
use of more reliable taxonomic characters such as degree of development of 
secondary branches, disarticulation of the spikelet, length of glumes in rela- 
tion to the contiguous lemma, and the number of stamens. Using these reliable 
characters in any future revision of the genus will reflect a clearer picture of 
new taxa remaining to be discovered, new records, and nomenclatural changes. 
Likewise, anatomical characters have been demonstrated to be taxonomically 
significant for infrageneric levels in Eragrostis (Van den Borre & Watson 1994). 

Eragrostis ancashensis PM. Peterson, Refulio & Tovar, a Peruvian endemic 
known only from the Andes of the Cordillera Blanca (Department of Ancash), 
was recently described by Peterson et al. (2000) on the basis of collections with 
diagnostic characteristics such as dark-green and plumbeous-spotted spikelets 
with a ciliate, flattened rachilla. When discussing the relationships of their new 
entity, the authors pointed out, on the one hand, the possibly close relationship 


SIDA 19(4):1157 — 1161. 2001 


1158 BRIT.ORG/SIDA 19(4) 


with Eragrostis magna Hitchcock, and on the other, a certain similarity with 
Eragrostis macrothyrsa Hack. Curiously, a detailed examination of the holotype, 
isoparatypes and the original illustration of E. ancashensis does not reveal its 
genuine alliance with E. magna as Peterson et al. (2000) mentioned, since these 
specimens as well as their figure 1, exhibit a particular set of features, viz. inflo- 
rescence with spreading or patent branches, reflexed pedicels, ovate spikelets 
3.0-6.1 mm long, glumes equalling or exceeding the contiguous lemma, tena- 
cious rachilla with hairs up to 1.5 mm long, 3-8 florets per spikelet, and ovate 
lemmas, which clearly distinguish these plants from Hitchcock's entity and 
which, on the contrary, show an undeniably closer similarity to another well 
known member of the genus also endemic to Peru, Eragrostis pilgeri Fedde. In- 
deed, when examining the lectotype and additional collections of E. pilgeriand 
comparing them with the material on which E. ancashensis was founded, it is 
impossible in our opinion to find the slightest difference in any way that al- 
lows any specific separation between the two species. Glands—which have not 
been previously reported for E. pilgeri—are usually present on the sheaths, 
pedicels, glumes, lemmas, and paleas of E. pilgeri; they are very conspicuous in 
Weberbauer 2746 (type of E. andicola var. robustior). Moreover, it is important 
to point out the remarkable overlap in distribution of both species, mainly con- 
fined to the Department of Ancash (northern Peru), which reinforces our argu- 
ment. The similarities of E. pilgeri were not taken into account by Peterson et 
al. in the discussion of their new species, E. ancashensis 

The preceding evidence—in our opinion—leaves no doubt on the 
conspecificity of E. ancashensis PM. Peterson, Refulio & Tovar and E. pilgeri 
Fedde and the consequent synonymy of the two names. The following list pro- 
vides complete and updated synonymy for Eragrostis pilgeri. 


Eragrostis pilgeri Fedde, Just’s Bot. Jahresber. 34:18, pl. 3. 1908. Based on Eragrostis 
andicola Pilger, Bot. Jahrb. Engler 37:377. 1906. Non E. andicola Fries, 1905. Type: PERU: DEPT. 
ANCA ACHS: inter Samanco et hae ama Hacienda Cajabamba; in formatione + aperta, 
3000-3500 ms.m., Weberbauer sek florens 
mense aren 1903 (HOLOTYPE: B- . stroyed LECTOTYPE, here designated: MOLI, Fig. 1; 
ISOLECTOTYPES: fragments at BAAI, l 


Eragrostis andicola forma humilior Pilger, Bot. Jahrb. Engler 37:377. 1906. Type: PERU: DEPT 
CACHS: inter Samanco et Caraz, infra Hacienda Cajabamba; in formatione aperta, 3700 
ms.m., Weberbauer 3036, florens mense Majo 1903 (HOLOTYPE: B-destroyed; LECTOTYPE, here 
designated: MOL). 

Eragrostis andicola var. robustior Pilger, Bot. Jahrb. Engler 37:377. 1906. Type: PERU: DEPT. ANCACHS: 
in prov. ale infra Ocros, in formatione aperta, fruticibus et plantis herbaceis mixta, 
3000-3200 ms.m., Weberbauer 2746 (HOLOTYPE: B-destroyed; LECTOTYPE, here designated: 
MOL ISOLECTOTYPE: fragment at US 

Eragrostis carazensis Pilger, Bot. Jahrb. ile 56:Beibl. 123:27. 1920. Typr: PERU: Dept. ANCACHS: 
Caraz, offene Formation bei 2200-2500 m t. M., Weberbauer 2999, Mai 1903 (HOLOTYPE: B- 
destroyed; LECTOTYPE, here designated: MOL! ISOLECTOTYPES: fragments at BAAI, US). 

Eragrostis pilgeri A. Hitchcock, Contr. US. Natl. Herb. 24:342. 1927. Based on E. andicola Pilger, 


AICCDIA AND CRANDA 


ba 
8 ie Gee Fear 


for de las forest Oo 


len y dom . 


erie 


DE AGRICULTURA 
Lime -  rexu 

RBARIO NACIONAL 
te Breast, 


OME sncreemcee ov + 


TLE, 


PPT AS 


Localidad LA yl {} 
Calldad de} sitio ia al Meeps ss 
Tap he Gprnnintnd Cor artatlpe hae 


Itura sobre ol nivel del mar : Jeno 5500 w mt Fecha 
re vulgar: Q 


ome aT 
* es eee 


Coleotor : ey Weis ; 


Tadigcd. 
riba, ontie Carag oy Sacerance,. Bar Hucaylas. 
 Pritan tad OAS “tcntni 


corr ondearted eoharcetes 


Ce aL 


> EB. pilferi Fedde . 
es P t Ta. 
__ 


1159 


3114, MOL). 


1160 BRIT.ORG/SIDA 19(4) 


; ti } l {. Peterson, Refulio & Tovar, Sida 19:66. 2000. TyPE: PERU: DEPT. ANCASH. 
Pomnee Recuay: Cordillera Blanca, approximately 20 km E of Raquia on Route 02-014 on 
roads towards Huaraz, along roadside and steep rocky slopes with shrubby Asteraceae, 
Lupinus, Vicia and Commelina, 3000 m, 20 Mar 1997, PM. Peterson & N. Refulio Rodriguez 
13793 (HOLOTYPE: USM, ISOTYPES: K, MO, NY, RSA, TAES, UC, US, WIS). 


Lastly, we refer to the paratypes of E. ancashensis ees 14577 and 14594, 
which we had the opportunity to check. Here, the characteristics so dissimilar 
in appearance to E. pilgeri and consequently to E. manera sensu Peterson et 
al—namely taller culms (70-95 cm tall), longer and wider inflorescence (20- 
37 cm long x 12-16 cm wide) with longer branches (up to 14 cm long), longer 
spikelets (8-15 mm long) which are oblong to narrowly oblong rather than ovate, 
shorter glumes which are shorter than the contiguous lemma, rachilla with 
shorter and fewer hairs, more florets per spikelet [(8-)12-20], etc.—could well 
belong to another member of the genus with completely different relationships. 

Thus, a remarkable affinity with E. bahiensis Schrader ex Schultes, a spe- 
cies not known from Peru (Davidse & Brako 1993) could rather be established. 
This species shares some features with the collections of Ferreyra, e.g, plant 
size, inflorescence size, spikelet size and shape, length of glumes in relation to 
the contiguous lemma, tenacity of rachilla, number of florets per spikelet, and 
lemma shape (ovate). Nevertheless, E. bahiensis differs by possessing shorter 
glumes (1.0-15 vs 2.0-2.5 mm long), shorter lemmas (1.5-2.0 vs 25—3.0 mm 
long), and a glabrous rachilla (Hitchcock 1951; Renvoize 1998). 

We agree with Peterson et al. (2000) that the collections of Ferreyra could 
have some relationship with Eragrostis magna. With this species, they share 
some characteristics such as spikelet shape and rachilla pilosity. However, E. 
magna differs by having ascending panicle branches, shorter spikelets (7-10 
mm long), a deciduous rachilla disarticulating between the florets, and fewer 
florets per spikelet (7-10) (Hitchcock 1927). 

On the other hand, it is possible that such peculiar collections represent 
quite old stages of E. pilgeri because the features they possess have been ob- 
served in the latter species with relative frequency by one of us (J.A.) despite 
not having been reported by previous authors (Hitchcock 1927; Standley 1936; 
Tovar 1993). In this respect, it is worth remarking the high index of variability 
noticed in different populations of E. pilgeri for characters suchas size of plant, 
degree of pubescence on blades and sheaths, presence of glands on sheaths and 
spikelets (pedicels, glumes, lemmas, and paleas), degree of panicle development, 
size and shape of the spikelet, degree of pubescence of rachilla, number of flo- 
rets per spikelet, which could have compelled Tovar (1993) to place E. carazensis 
into the synonymy of E. pilgeri. 

Finally, it is possible that these two paratypes belong to a species remain- 
ing to be named; however, in our opinion, only a careful examination of a greater 
number of future collections of these plants will make it possible to determine 
the consistency and taxonomic validity of the characteristics mentioned. Be- 


AL EGRIA AND GRANDA 1161 


fore such time, any decision that seeks to clarify the situation of the specimens 
Ferreyra 14577 and 14594 would be questionable to us. 


Additional material examined: Eragrostis pilgeri: PERU. Dept. Ancash. Provincia Bolognesi: entre 
Chasquitambo y Conococha, 3000-3200 m, 24 May 1962, Ferreyra 14476 (USM); Mishahuayunca, 
cerro al frente de Ocros, 3400 m, 8 Jul 1974, Cerrate 6190 (USM); Rumpuquio cerro al S de Ocros 
3500 m, 27 Abr 1977, Cerrate et al.6510 (USM); camino a Bellavista, 3200 m, 29 Abr 1977, Cerrate 6580 
(USM). Provincia Huaraz: 10 km by road from Cachabamba (77°51'W, 9°27'S), 2870 m, 6-8 Jun 
1985, Smith & Buddensiek 10872 (USM). Provincia Huaylas: encima de Huaylas, 2700-2800 m, 3 Jun 
i a 14606 (USM); Huascaran National Park, Auquispuquio area of ruins (77°58'W, 8°50’S), 
3800-3900 m, 7 Apr 1986, Smith et al. 11959 (USM). 
pee ancashensis: PERU. Dept. Ancash. Provincia Corongo: 7 km NW of Yupan on road 
to Bambas, 3220 m, 26 Mar 1997, Peterson & Refulio 13915 (USM, isoparatype); 7 km NW of Bambas, 
2710 m, 26 Mar 1997, Peterson & Refulio 13919 (USM, isoparatype). Provincia Huaylas: entre Caraz y 
Huallanca, 2200-2300 m, 2 Jun 1962, Ferreyra 14577 (USM, paratype); entre Huaylas y el Callejon, 
400-2500 m, 3 Jun 1962, Ferreyra 14594 (USM, paratype). 


ACKNOWLEDGMENTS 


The authors wish to express their gratefulness to Zulma Rugolo (SI) for review- 
ing the first draft of the manuscript and mailing Pilger’s protologues; Gabriel 
Rua for sending images of the types at BAA; Werner Greuter and Mia Ehn Salter 
for providing information about the types housed at B and S, respectively; and 
Nancy Refulio (USM) for supplying part of the literature. Appreciation is ex- 
tended to the staff of USM and MOL for allowing access to the collections. We 
are also indebted to Kelly Allred and Paul Peterson for their reviews of the manu- 
script, and Barney Lipscomb without whose support the present paper could 
not have been published. 


REFERENCES 


Cayton, L.and S.A. Renvoize. 1986. Genera graminum. Grasses of the World. Kew Bull. Add. 
Ser. 13:1-389 

Daviose, G.and L. Brako. 1993. Eragrostis, in L. Brako and J.L. Zarucchi, eds. Catalogue of the 
flowering plants and gymnosperms of Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 
45:945-947 

HitcHcock, A.S. 1927.The grasses of Ecuador, Peru and Bolivia. Contr. U.S. Natl. Herb. 24:291—256. 

HircHcock, A.S. 1951. Manual of the grasses of the United States. Ed. 2. U.S.D.A. Bull. Misc. 
Publ. 200:1-1051. 

Peterson, PM.,N. Rerutio, and O. Tovar. 2000. Eragrostis ancashensis (Poaceae: Chloridoideae), 
a new species from Ancash, Peru. Sida 19:65-70. 

Renvoize, S.A. 1998. Gramineas de Bolivia. Royal Botanic Gardens, Kew. 

STANDLEY, PC. 1936. Gramineae. In J.F. Macbride, ed. Flora of Peru. Field Mus. Nat. Hist., Bot. 
Ser. 13:96-261. 

Tovar, O. 1993.Las gramineas (Poaceae) del Peru. Ruizia 13:1-480. 

VAN DEN Borre, A.and L.Wartson. 1994.The infrageneric classification of Eragrostis (Poaceae). 
Taxon 43:383-422. 


1162 BRIT.ORG/SIDA 19(4) 


Book REVIEW 


WILLIAM Dupin. 200L Soils. (ISBN 0-565-09148-4, pbk.). lowa State University Press, 
2121 South State Street, Ames, IA 50014, U.S.A. (Orders: 800-862-6657, 515- 
292-015—distributed for the Natural History Museum, London). $19.95, 110 
pp, 97 color photos, 19 b/w photos, 10 drawings, | color map, 81/4" x 91/4". 


Contents.—Preface, The author, Soils—the nuts and bolts; Soil taxonomy and geography; Soil biology; 
Soil fertility; Soil use and misuse; What lies ahead?. pues bine alice information. 


As stated in the preface, “The soils that cover the Earth’ -e determine, to a large extent, 
the prosperity of those who depend on them for the prin oe food and fibre. .. coe recent ex- 
amples of soils degradation emphasize the strong lin Ww ity and human welfare. Sus 


tained prosperity therefore depends on a thorough seers of oi coupled with sound man- 
apoTient PrEcHees, 


= 


ssmall volume is a good beginning for gai I d li | iation for 
this “indispensable natural resource.” The book oronde: a brief, yet iiderstandable and well-illus- 
trated saeaet toa complex subject. It does not attempt to go into great depth about the details 
f soil science. Rather, it gives a basic presentation of the field that is accessible, informative, and 
interesting to - intelligent lay person. For example, the introductory chapter, “Soils—the nuts and 
olts,” covers a variety of important basics such as soil formation, pei materials, soil minerals, 
mineral weathering, soil texture, eeu matter, porosity; and water in soils. 

The chapter on “Soil taxon phy” is particularly helpful. Currently, there are two 
main hier ars c nee oe of the sorts soils. One system (referred to as UNESCO- 
FAO) was developed by the United Nations Educational, Scientific and Cultural Organization 
(UNESCO) et with the Food and Agriculture Organization (FAO), while the other, called Soil 
Taxonomy, was developed by “a team of international scientists and the U.S. Department of Agricul- 
ture.” Clear descriptions and photographs of the 12 orders of the Soil Taxonomy system, and a table 
comparing this system with the UNESCO-FAO classification, bri ing satisfying clarity to what can 

ometimes be a confusing array of names. The numerous excellent color photographs are particu- 
larly helpful in visualizing the various soils being discussed in the text. 

As someone living on the Blackland Prairie of Texas, whose soils have been devastated by years 
of misuse and now increasing development, it was very lee: toseea wa. chapter, “Soil use and 
misuse,” devoted to issues of soil conservation. Concise but i discussions of erosion, the 
buildup of salts in soils, and soil pollution oo. important issues in soil management and con- 
servation. Also, such special focus topics as “Salinity ‘down under’ (Australia),” “Bioremediation— 
nature’s chimney-sweep,” and “Famine in Africa” Aes real world examples and a human face to soil 
conservation. In addition, conservation is a theme incorporated throughout the book—for example, 
there isa special section on eutrophication (‘Too much of a good thing—eutrophication”) in the chapter 
on i fertility.” 


roar) 


In several sections I had the feeling | wanted more—additional information and details. That's 
probably a positive thing. As many teachers will attest, if you can interest a student so they want to 
learn more on their own, you've won half the sae Indeed, the book only intends to be an under- 

tandable introduction to a large and comple line, and it clearly succeeds in that goal. It is a 
2 yet helpful overview of the field and should aes cobs understanding to a wide audi- 
ence. yee ie ae more in- peep — wa appreciate a page at the end of the book 
with a list of 


is anumber of web site addresses.—George 
M. ne Dept. of Biology, Austin College Sherman, TX 75090, and Botanical Research Institute of 
Texas, Fort Worth, 1X 76102, ediges@austinc.edu. 


SIDA 19(4): 1162. 2001 


SCLERIA LACUSTRIS (CYPERACEAE), AN AQUATIC AND 
WETLAND SEDGE INTRODUCED TO FLORIDA 


Colette C. Jacono 
United States Geological Survey (USGS) 
7920 NW 71° St. 
Gainesville FL 32653, U.S.A. 
ABSTRACT 


A non-native species of Scleria, S. lacustris is reported from six counties and three major hydrologic 
regions in Florida. Biogeography and habitat in Florida are addressed. A description, key features 
and illustration are presented. 


RESUMEN 


Se presenta aqui | iad ica del género Scleria, S. lacustris, de seis condados y 


de tres regiones hidrol dgicas de Florida. Se hacen anotaciones sobre su biogeografia y habitat en 


Florida. Se presenta una descripcion con caracteristicas clave junto con una ilustracion. 


A robust, annual member of its genus, Scleria lacustris C. Wright is being found 
with increasing frequency in freshwater marshes of peninsular Florida. This 
noteworthy species is native to scattered regions of the Neotropics and Africa 
(Core 1936; Hennessy 1985). Adventive populations have been documented in 
six counties located in the headwaters of three major USGS hydrologic regions: 
the Upper St. Johns River, the Kissimmee River, and the Big Cypress Swamp 
drainages. 

Scleria lacustris was first reported in 1998 based on a 1989 specimen from 
a Lee County wetland in the upper reaches of the Big Cypress Swamp drainage 
(Wunderlin 1998; Tobe et al. 1998). In 1990 it was documented at a wetland eight 
km east and noted as the dominant plant (M. Roessler, pers. comm.) Not recog- 
nized at the time was a 1988 specimen from Jane Green Swamp, a conservation 
marsh in the St. John’s Water Management District, Brevard County in eastern 
Florida. The Brevard County specimen had been misidentified as Scleria 
vaginata Steudel. In 1999 S. lacustris was documented in the Kissimmee River 
drainage of central Florida in Polk County at the Disney Wilderness Preserve 
(DWP) and at London Creek, two km west of DWP boundaries. The following 
year additional stations were found within DWP (Polk and Osceola Counties), 
and east of DWP on South Florida Water Management District conservation 
lands. Since 2000, collections have been made from St. Johns River Water Man- 
agement District marshes in Indian River County and from the lower Kissimmee 
drainage in Okeechobee County. These latest stations lie south of the earliest 
locale (Brevard County) where recollection has also been made (Fig. D). 


SIDA 19(4): 1163 — 1170. 2001 


1164 BRIT.ORG/SIDA 19(4) 


80° 


Atlantic 


28° of 


Mexico 


N 
26° l \ 


100 0 100 Kilometers 


Fic. 1.6 hir-a HT + a a gy ee | fe im Clarida Callarez + peer Pee wees 1 TICCC hudenl 
. Yi} f 


hagdad t ddal; Juateth hala tt C lat * eb rt (ee £¢} Din f. 


i ¥ J f r J J IF 


g 
Swamp, the Kissi and the Upper St. Johns River 


Scleria stands apart from other genera in the Cyperaceae for its unisexual 
flowers and prominent, white, bone-like achenes (Kessler 1987). The genus is 
distributed primarily in tropical and warm-temperate regions. Kessler (1987) 
recognized and described ten North American species of Scleria, placing em- 
phasis on the sculpturing of achenes and pubescence on the achenes, scales, 
bracts, leaves, and culms. Camelbeke and Goetghebeur (1999) used the ligule as 
a diagnostic character in Scleria and presented a key to the five ligulated Neo- 
tropical species so far known. In the Americas, the Neotropical S. lacustris was 
most recently treated by Core (1936). Hennessy (1985) most recently detailed 
the species from Africa. 

Scleria lacustris lacks a genuine ligule, but does have a sheath-top exten- 
sion called a contraligule (Camelbeke & Goetghebeur 1999; Pic. 2e) 1 he 
contraligule for this species has a membranous margin, which Core (1936) de- 


— 


JACONO, SCLERIA LACUSTRIS IN FLORIDA 1165 


scribed but failed to accurately account for in his key. All Florida specimens 
exhibit this margin; a scarious, tongue-shaped appendage, 2-5 mm long, at the 
top of the contraligule. It is also apparent in a digital image, Wright s.n. (NY) 
(NYBG 2001). 

Core (1936) believed Scleria lacustris to be endemic to the type locality of 
Cuba; only three specimens from two locales were available on which to base 
his description. Fairey (1972) discovered that the original notes for Core (1936) 
described leaves as “6 mm to 16cm wide.” This differs from “1-6 mm wide,” an 
error that appeared in the published monograph (Core 1936). Florida plants, 61 
to about 230 cm in height, stand two to three times taller than specimens from 
Cuba. Also the leaf width is broader, 11-25 mm wide. Consistently larger mate- 
rial was collected in the St. Johns drainage where plants grew emergent from 
water. These plants were taller, with solitary culms, long panicles (to 21 cm) 
and adventitious roots up to 44. cm above the culm base. In these respects plants 
from the St. Johns drainage bore a greater resemblance to S. lacustris described 
from aquatic habitats in Africa (Hennessy 1985; Robinson 1966, Nelmes 1955). 
African plants originally described as S. aquatica Cherm. are considered con- 
specific with S. lacustris (Nelmes 1955, Hennessy 1985; Fairey 1972). The Afri- 
canentity asa whole is infrequently distributed in six countries across tropical 
Africa and in Madagascar (Hennessy 1985). 

Fairey (1972) examined both Neotropical and African speci of Scleria 
lacustris and found African plants taller and more robust, with larger, better- 
developed panicles. He noted leaves on African specimens as acute and those 
on Neotropical plants as obtuse. Core (1936) described Cuban plants with ob- 
tuse leaves. In Florida, the leaves of S. lacustris are acute; they gradually taper to 
a point. Fairey (1972) attributed differences between Neotropical and African 
representatives of S. lacustris to their wide geographical separation. 

In the Neotropics, Scleria lacustris is now known to range far beyond Cuba. 
Camelbeke confirmed specimens from Costa Rica, Cuba, Jamaica, Guyana, 
Suriname, French Guiana, Brazil and Paraguay, but within this wide distribu- 
tion still regards S. lacustrisas “a very rare species” (K. Camelbeke, pers. comm.). 

In Florida, Scleria lacustris exists as an annual species, recurring by seed 
in locally abundant populations. Seedlings emerge in late spring. Flowering 
occurs August to September and achenes remain on standing plants Septem- 
ber through December. By February plants in the St. Johns drainage were col- 
lapsed with bases and roots withered. Persisting structures are absent in Florida, 
although S. lacustris may perenniate in Africa (Nelmes 1955; Fairey 1972, 
Hennessy 1985). 

Throughout its Florida range, Scleria lacustris resides in seasonal marshes 
in or near former wet cattle pasture. The mechanisms of introduction and dis- 
persal are not known. According to Hennessy (1985), the achenes sink in water, 
yet, one hundred achenes collected from the ground in May remained buoyant 


1166 BRIT.ORG/SIDA 19(4) 


in the laboratory for several days. Achenes floating through ditches and washed 
along lakeshores may explain spread in some regions. However, water flow can- 
not account for widely scattered populations in more isolated sites, suggesting 
dispersal by birds or other animals as an additional mechanism of introduction. 

In marshes of the Upper St. Johns River drainage, Scleria lacustris is en- 
countered as scattered patches approximately five m2 to several hectares in size 
on slight rises in peat substrate. Plants are observed in drier years and may re- 
quire recession of standing water for seedling recruitment. However, mature 
specimens collected in late season are commonly found growing in water to 
one m (L. & J. Harrison, pers. comm.). Drier conditions existed at DWP in the 
Kissimmee River drainage, where S. lacustris was observed during 2000 at ap- 
proximately 20 stations (C. Campbell, pers.comm.). At that time colonies reach- 
ing L8 ha were found at marshes, ditches, and lakefronts remaining dry after 
several years of extreme drought. Hennessy (1985) suggested that the annual 
habit, uncommon in the subgenus Scleria, serves as a drought-escape mecha- 
nism in tropical regions that experience seasonal drought. The increasing oc- 
currence of S. lacustris in Florida marshes could be related to changing hydro- 
logic conditions. 

Scleria lacustris can be recognized in the field by its large size, thick three- 
sided culms and broadly linear, plicate leaves. Terminal panicles may extend 
over I-2 meter in height and are open and conspicuous when achenes mature. 
Plants are extremely scabrous, especially leaf margins, midribs, rachis and 
bracts. The harsh texture results from minute prickles, developed as barbed 
outgrowths of silica deposited in the epidermal cells (Metcalfe 1971). Seedlings 
are bright green, with purple tinged basal sheaths, red roots and tapering, pli- 
cate leaves. 

The hypogynium, a hardened pad at the base of the achene, is an important 
taxonomic feature in Scleria. Wunderlin’s (1998) key. in addressing Florida, dis- 
tinguishes S. lacustris from North American species and groups S. lacustris with 
S. triglomerata in having a smooth-surfaced achene and a hypogynium with- 
out tubercles. Wunderlin (1998) separated S. lacustris from S. triglomerata by 
noting the broad, crusty hypogynium and the narrower leaves of S. triglomerata 
versus the narrow, smooth hypogynium and the wider leaves of S. lacustris. 

Kessler (1987) noted the achene body of S. triglomerata as ovoid-globose, 
and pointed to the resulting circular base of the achene as a key element. The 
achene body of S. lacustris appears more ovoid-ellipsoid and its achene base 
more triangular. Addition of these elements after line 3 in the key of Kessler 
(1987) is as follows: 

1. Achene body smooth or with longitudinal ridges (2) 
1. Achene body rough, reticulate or with transverse ridges. 
2. Base of achene circular to obtusely triangular, with a hypogynium (3) 
2. Base of achene triangular without a hypogynium. 


— 


JACONO, SCLERIA LACUSTRIS IN FLORIDA 1167 


3. Tubercles present between achene and hypogynium Scleria oligantha 
3. fetes absent between achene and hypogynium (4) 
Hypogynium broad, granular- or papillose- crusty; base of achene circular; 
“leaves less than 1 cm wide Scleria triglomerata 
4. Hypogynium narrow, smooth; base of achene obscurely triangular; leaves 
greater than 1 cm wide cleria lacustris 


The following description of Scleria lacustris in Florida, adapted from Core 
(1936), is based on all specimens reviewed and six unvouchered plants from 
Indian River County. 


Scleria sepia C. Wright in Sauvalle, Anales Acad. Ci. Med. Habana 8:152. 1871. 
(Fig. 2.) Type: CUBA: Wright s.n. (LECTOTYPE: K; designated by E.A. Robinson, Kew Bull. 18:517- 


519. 1966). 


Common names.—Wright’s Nut-rush, Canuella, Little Cane 

Robust, annual herb 0.6-2.3 m tall. Rhizome absent. Roots stout and fibrous, 
at base and submersed nodes of culm, red-brown. Culms erect, solitary to sev- 
eral, often with 1-2 undeveloped basal tillers, 8-24 mm thick near base to 5-13 
mm thick aerially, triquetrous with angles retrorsely scabrous. Leaf blades 41- 
90 cm long, 11-25 mm wide, acute, plicate, transverse section inversely W- 
shaped, glabrous, gi d midribs scabrous; sheaths scarcely winged, loose, 
closed, tinged with purple streaks, sheath hairs strigose to hispid, white and 
sparkling; ligule absent; contraligule rounded, 3-12 mm long, 6-17 mm wide at 
base, the margin with a scarious appendage, appendage triangular to tongue 
shaped, 2-5 mm long. Inflorescence open paniculate, the branches ascending to 
spreading, rigid, antrorsely scabrous, axillary and terminal, terminal inflores- 
cence 19-48 cm long; lower bract of inflorescence foliaceous, 9-30 cm long, 5- 
1.6 mm wide, bractlets to 48 mm long, < 1 mm wide, antrorsely scabrous, awl 
shaped from broadened base, base margined with dark bristles. Spikelets both 
bisexual and male. Staminate spikelets, 4-5 mm long, 0.8-1.2 mm wide, scales 
lanceolate, mucronate, maroon to dark purple, flower with 3 stamens. 
Gynecandrous spikelets common, 4.5-6.5 mm long, pistillate scales 3-4.5 mm 
long, 2.4-4 mm wide, broadly ovate, abruptly acuminate, ciliolate on the mar- 
gin, maroon to dark purple. Androgynous spikelets less common. Achene 3-4 x 
2.3-2.5 mm, ovoid- ellipsoid, faintly three sided, base obscurely triangular, 
smooth, shining, white to mottled gray or tan, equaling or exceeding the scales. 
Hypogynium small, depressed 0.5-<] mm, obtusely triangular to three lobed, 
the margin entire, narrow, 0.1-0.2 mm high, tan to brown. 


Voucher specimens: U.S.A. FLORIDA. Brevard Co.: Rare, growing in water, Jane Green Swamp, W 

of Lake Hell’n Blazes, near W property peed about 14 mi oy of peieO Hine ca.28°02'N, 80°48 Ww 
6 Oct 1988, G.B. Hall s.n. (FLAS!), duplicate sp norig ja vaginata Steudel,; 

Jane Green Swamp, St. eu Water iancecgen District UTM 3099747.5 N,518595.9 W,abundant 
at this station, growing in 14 cm of water, freshwater marsh with Rhynchospora inundata, Cyperus 
articulatus, Pontederia Ae Sagittaria lancifolia, Panicum hemitomon, Polygonum punctatum, 
Leersia hexandra, Cyperus haspan, 21 Oct 2000, J. Harrison & L. Harrison 259 with K. Snyder (FLAS!). 


1168 BRIT.ORG/SIDA 19 


— 


4) 


Fic. 2. Scleria lacustris C. Wright drawn from Harrison & Harrison 259 by Laura Line. A. Achene, profile and base with 
hypogynium. B. Habit. C. Porti f culm showing leaf, sheath and ligule. D. Spikelets 


J J 


Indian River Co.: Blue Cypress Marsh, ca.600 m E of northern section of Blue Cypress Lake, Nov 
2000, G. Nichols s.n. (FLAS!); Blue Cypress Marsh,ca.600 m E of northern section of Blue Cypress Lake, 
27°41 .869'N, 80°41.659'W withered plants with Eleocharis, 10 Feb 2001, CC Jacono 204 with G. Nichols 
FLAS!); Blue Cypress Marsh, 27° 41.869’ N, 80° 41.659’ W, immature plants 150 cm high growing in 
76 cm of water, 16 Aug 2001, G. Nichols s.n. (FLAS!). Lee Co.: ca. 10 mi SE of Ft. Myers, herbaceous 
wetland, 27 Nov 1989, W. Cox s.n. (USF);ca. 10 mi SE of Ft.Myers, hert wetland, 16 Dec 1989, W. 


— 


JACONO, SCLERIA LACUSTRIS IN FLORIDA 1169 


Cox s.n. (USF;FLAS!);wet prairie along Griffin Dr.,0.5 mi N of Gateway Blvd,,in the Gateway Commu- 
nity, Nov 1990, M. Roessler s.n. (FTG, scanned nase seen; USF), eae originally determined as 
Scleria triglomerata Michx. Okeechobee Co.: Approx. 5 mi W of US 441 on SR 724, uae 50m 
from wetland edge, 19 Sep 2000,C. Lane s.n (FLAS). Osceola Co.: Disney Wilderness Preser U3T4, 
along transect in HU, cypress dome community, 40-50 plants, C. Campbell s.n. 16 Oct 2000 an 

Polk Co.: London Creek, Tract A. Oct 1999, C. MacGregor s.n. (USF!); Disney Wilderness Preserve, N 
ce of Lake Hatchineha, ditch of N side lakefront road, water depth 45 cm, colony 9 m diameter, P. 
Royston, 8 Nov 1999 (PIHG)); Disney Wilderness Preserve, HUT, Aug 2000, C Campbell s.n. (dwp!);just 
W of Dead River on Johnson Island, McKinney Tract, 1285 R30E Sec.13, open marsh with Sagittaria 
lancifolia and Sacciolepis striata, 12 Oct 2000, B. Hansen 12,894 with R. P Wunderlin & C. MacGregor 
(USF!); Disney Wilderness Preserve, N side of Lake Hatchineha, S of the T off old marsh front road, 
seedlings 11-13 cm tall with empty pericarp attached, 7 May 2001, C.C. Jacono 266 w/V. Ramey, GC; 
Campbell and K. Yeuell (FLAS!); Disney Wilderness Preserve, N side of Lake Hatchineha, S of the T off 
old marsh front road, mature plant cultivated from seedling collected 7 May 2001, 11 Sep 2001,C.C. 
Jacono 309 (FLAS!). 


ACKNOWLEDGMENTS 


I gratefully acknowledge the curators of the herbaria FLAS, FSU, FTG, PIHG, 
USF and dwp (Disney Wilderness Preserve, herbarium) for granting access to 
their collections and freely providing information on specimens. Kent Perkins 
of FLAS was instrumental in focusing my attention on the initial Florida speci- 
men and his expertise in searching the literature and catalogues was extraor- 
dinarily helpful. Konraed Camelbeke confirmed identity of Harrison and Har- 
rison 259 and shared geographic details of collections compiled for his treatment 
of Scleria in the forthcoming Flora of the Guianas and I thank him for his gen- 
erosity. My sincere appreciation is extended to John Fairey, III for providing sec- 
tions of his unpublished dissertation and original specimen notes, to Amy 
Benson for assistance with the cartography and Laura Line for her superb illus- 
tration. My thanks are also extended to Cindy Campbell, Kay Yeuell and Gary 
Nichols for hosting memorable field tours. Financial support was provided in 
part by the United States Fish and Wildlife Service and the University of Florida 
Center for Aquatic and Invasive Plants. 


REFERENCES 


CAameLBeke, K. and P. GoetcHeseur. 1999. The ligule, a new diagnostic character in Scleria 
(Cyperaceae). Syst. Geogr. PI. 68:73-84. 

Core, E.L. 1936. The American species of Scleria. Brittonia 2:1-105. 

Fairey, J.E., lll. 1972. The genus Scleria in North America. Dissertation, West Virginia Univer- 
sity, Morgantown, WV. 

Hennessy, E.FF. 1985. The genus Scleria in southern Africa. Bothalia 15:505—-530. 

Kesster, J.W. 1987.A treatment of Scleria (Cyperaceae) for North America north of Mexico. 
Sida 12:391-407. 

Mercacre, C.R. 1971. Anatomy of the Monocotyledons. V. Cyperaceae. Oxford University 
Press, London. 


1170 BRIT.ORG/SIDA 19(4) 


Newmes, E. 1955. Notes on Cyperaceae, XXXVIII. Kew Bull. 10:415-453. 

NYBG. 2001. The New York Botanical Garden. Vascular plant type catalog. (http:// 
www.nybg.org/bsci/hcol/vasc/). 

Rosinson, E.A. 1966. A provisional account of the genus Scleria Berg. (Cyperaceae) in the 
‘Flora Zambesiaca’ area. Kew Bull. 18:487-551. 

Tose, J.D., K.-C. Burks, R.W. CANTRELL, M.A. GARLAND, M.E. Sweetey, D.W. Hatt, P WALLACE, G. ANGLIN, G. 
Netson, J.R. Cooper D. Bicker, K. Gitgert, N. AYMOND, K.GReENWwOooD, and N.RayMonb. 1998. Florida 


wetland plants: an identification manual. Florida Department of Environmental Pro- 
tection, Tallahassee. 


WUNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, 
Gainesville. 


SOUTH AMERICAN SKULLCAP 
(SCUTELLARIA RACEMOSA: LAMIACEAE) IN THE 
SOUTHEASTERN UNITED STATES 


Alexander Krings Joseph C. Neal 
Herbarium, Department of Botany Department of Horticultural Science 
North Carolina State University North Carolina State University 
Raleigh, NC 27695-7612, U.S.A. Ralei oo NC 27695-7609, U.S.A. 
Alexander_Krings@ncsu.edu e_Neal@ncsu.edu 
ABSTRACT 


Although previously reported from scattered localities, studies of the habitats and distribution in 
the United States of the South American native Scutellaria racemosa Pers. are lacking, To analyze 
trends in the spread of S. racemosa throughout the southeast we e studied 82 Speci from 16 her- 


baria. Collections taken from roadsides, lawns or golf courses horticultural plantings 
accounted for about 76% of specimens examined. Establishment is facilitated by vegetative repro- 
duction via rhizomes and a capacity to produce flowers and fruits throughout the year. Patterns of 
distribution are likely a result of human activity such as transport of contaminated nursery stock or 
sod, as well as seed spread on right-of-way mowing equipment. 


RESUMEN 


itada de varias localidades, no hay estudios de los habitats y la distribucion en los Estados 


Unid os de la especie suramericana Scutellaria racemosa Pers. Para analizar tendencias en el 
movimiento de S. racemosa en el aac de los ae Unidos, estudiamos 82 especimenes de 16 
erbarios. R 1 0 Campos golf, y viveros 0 plantaciones 


horticulturales, eee el 76% de los especimenes examinados EB establecimient est ral Tad 


ae la ee Nee anlve por name as y I flores 
] 


i t 
] | eR | ] } : ] J ] 


[ 
] t ss ] we 5 Tiss = = ] 


. plantas de por cortacéspedes. 


Native to South and Central America (Leonard 1927; Epling 1942), Scutellaria 
racemosa Pers. (South American skullcap, Lamiaceae) was first reported from 
North America in 1973 (Kral 1973) and subsequently reported from sporadic 
collections in the southeastern United States (e.g., Godfrey & Wooten 1981; Tobe 
et al. 1998). Following observation of encroachment of a population into nurs- 
ery pots and nearby natural areas in Chatham County, North Carolina (Krings 
& Neal 2001), we grew concerned over the potential of this species to become 
invasive in our state. Although previously noted from the southeast primarily 
in floras, checklists, and noteworthy addition articles (e.g., Kral 1973, 1981; 
Godfrey & Wooten 1981; Allen 1983; Jones & Coile 1988; Tobe et al. 1998), scant 
information concerning the invasion of habitats and overall distribution in the 
United States has been published. In this study we sought to more carefully 
examine the distribution and habitat of the species in the southeastern United 


SIDA 19(4): 1171 — 1179. 2001 


1172 BRIT.ORG/SIDA 19(4) 


States. Specifically, our objectives were to: (1) compile an up-to-date atlas of S. 
racemosa in the southeastern United States, (2) compile and analyze habitat 
and phenology data, and (3) analyze any trends in the spread of S. racemosa 
throughout the southeast. 


oa) 


METHODS 


The distribution and habitat of Scutellaria racemosa in the southeastern United 
States was compiled based on study of vouchered herbarium specimens, guided 
by previous literature citations (e.g., Penland 1924; Radford et al. 1968; Kral 1973, 
1981; Godfrey & Wooten 1981; Allen 1983; Jones & Coile 1988; Tobe et al. 1998: 
Wunderlin & Hansen 2000). Eighty-two collections were analyzed—compris- 
ing the totality of S. racemosa holdings in the following herbaria: AUA, DUKE, 
GA, GH, FLAS, FSU, LSU, MO, NCU, NCSC, NO, NY, UNA, USCH, USF TEX. 
Habitat and phenology data were taken from mounted specimens. In addition, 
field surveys were undertaken by Krings in east Texas and southeast South Caro- 
ina. 


RESULTS 


An updated distribution of S. racemosa is provided in Figure 1, d. New county 
records are reported from FL, GA, LA, SC, and TX and herbaria interested in 
updating their records may wish to consult the list of exsiccatae (Appendix A). 
The species remains unreported from Mississippi, although, based on its known 
distribution along Interstate 10 and habitat preference, it is extremely likely to 
occur—especially in the southern counties of Pearl River, Hancock, Stone, Har- 
rison, George, and Jackson. It is also likely occur in other Texas counties, par- 
ticularly Harris, Chambers, Jefferson, Orange. 

Scutellaria racemosa has been collected in the southeastern United States 
in fruit and flower in every month of the year and ina variety of habitats rang- 
ing from wet, roadside ditches to lawns and orange groves (Fig. 2). The largest 
number of collections (40%) were taken from roadside ditches, embankments, 
or right-of-ways. Collections from lawns or golf courses constituted approxi- 
mately 19% of all collections. Collections from nurseries or horticultural 
plantings constituted nearly 18%. The species has also been sporadically col- 
lected from pine forest, hayfields, and disturbed sites. Soils ranged from sandy 
to muck and sites from low to high maintenance (e.g, mowed turf )—suggesting 
a potential to infest varied environs and tolerance of site management activi- 
ties such as mulching and mowing. 


— 


DISCUSSION 
An obligate wetland species (Reed 1988), S. racemosa has already shown itself a 
potential weed of irrigated landscapes, such as lawns and golf courses (Fig, 2). 
Although first reported from Baldwin Co., Alabama (8 Jun 1971, Kral 43084, 


KRINGS AND NEAL, 1173 


a \: n lIni ic 


t+. ih + lt 


Fic. 1 f Scutelfari Pers. (Lami ) in tk t fi 
tion in 1965 (Mobile Co., Alabama) to present: A, 1965-1975; B, 1976-1985; C, 1986-1995; D, 1996-2000. 


AUA, FLAS, GA, NCU, UNA, USF) by Kral (1973), the earliest North American 
collection appears to be from neighboring Mobile Co. (10 Apr 1965, Deramus 
D378, UNA). The method of initial introduction is unknown, although it is likely 
that the species was introduced accidentally with horticultural plants shipped 
from Central or South America. Although the species is known from coastal 
Mexico (Epling 1942), an overland introduction does not seem plausible given 
the complete lack of collections from southeastern, coastal Texas and the lack 
of any Louisiana collections before 1982. In the ten years following the initial 
1965 collection in Alabama, further collections had been made in Florida and 
as far north as South Carolina (Fig. l,a). A comparison of Fig. 1 to roadmaps of 
the southeastern US reveals a close agreement between occurrence records and 
major by-ways (e.g,, 1-10, Hwy 17). Although occurrence patterns may be biased 
somewhat by the ease of roadside collection, the small number of collections 
from relatively isolated wetlands remains surprising - especially given the re- 
cent effort to study and delineate wetlands in the southeastern United States. 
Although unknown, it is not likely that seeds were dispersed so rapidly over 
such distances by natural means (eg., through the highly mobile avifauna). It 
is more likely that dispersal was achieved as a consequence to human activity, 
such as transport in sod, nursery crops, or right-of-way mowing equipment. As 


1174 BRIT.ORG/SIDA 19(4) 


Ditch or road right-of-way 
Lawn/golf course 


Nurseries or horticultural plantings 


a 2 
Creek-, lake- or pond edge or shore 

a 

a 

| 

| 

| 

T 


Disturbed sites 


Railroad embankment 


Pine forest 


Orange grove 


Hayfield 


US ee a | £ tl 4 oc eal 4 dD :. +h #L Dies 2: ic 
Fig. 2 


shown in Fig 2, about 18% of herbaria collections came from nurseries, horti- 
cultural plantings, or root balls of nursery crops. The collections from lawns 
and golf courses (19%), as well as lake or pond shores (13%), also potentially 
originated from populations established either from the root balls of introduced 
landscape plants or through the movement of sod. Dispersal based on trans- 
port of nursery stock has been previously implicated in the movement of other 
annual and perennial, rhizomatous weeds in the eastern United States, includ- 
ing Fatoua villosa Thunb.) Nakai (Massey 1975) and Artemisia vulgaris L.(Uva 
et al. 1997). 

Encroachment into natural landscapes by S. racemosa seems to be limited 
by degree of human movement and soil moisture. However, the apparent ease 
and speed of establishment along wet, roadside ditches is of concern as these 
populations can serve as persistent sources for slower, yet repeated advances 
into more natural settings. Establishment is facilitated by vegetative reproduc- 
tion via rhizomes and a capacity to produce flowers and fruits throughout the 
year. The recently documented North Carolina population (Krings & Neal 2001) 
was clearly spreading into nursery pots from established plants growing along- 
side ground tarpons, by sending rhizomes into pots through holes in the bases 
and tarpons. 

In the thirty-five years since its first collection in Alabama, S. racemosa has 
spread into seven states and is to be expected in an eighth—Mississippi. In tropi- 
cal America, the species is known from Vera Cruz to Valdivia (Epling 1942) and 
reported from grassy hillsides to damp thickets and stream banks (Leonard 


KRINGS AND NEAL, 1175 


1927). Based on its zonobiomic distribution in South America (Leonard 1927: 
Epling 1942; Walter 1985), it should subsist at least as far north as Baltimore, 
Maryland. The potential area of expansion in North America extends from the 
southern tip of Florida to southern Maryland, westward to western and north- 
ern Arkansas, and southward into east Texas. To combat this introduced weed, 
future studies are needed to evaluate (1) the population biology of S. racemosa 
and its effect on native, wetland vegetation, and (2) the extent of nursery crop 
infestation. 


APPENDIX A 


List of exsiccatae of Scutellaria racemosa Pers. in the southeastern US. 


U.S.A. ALABAMA. Baldwin Co.: Sandy gravelly railroad embankment through Hypericum pond 
just N of jct I-10 on Ala. 59 and 6 mi S Stapleton, abundant, 8 Jun 1971, Kral 43084 (AUA, FLAS, GA 
MO, NCU, UNA, USF). Covington Co.: LBW Junior College on US Hwy 84 in Andalusia, growing 
around the edge of the lake on the golf course, full sun, sandy wet soil, T4N, R16E, Sec. 15, 21 Apr 
1994, Diamond 9070 (AUA); Sandy loam of shoulder bypass US 29/84, N side of Andalusia, common 
in shadier sites, 2 Apr 2000, Kral 89387 (USCH). Lee Co.: In sandy soil in filtered sun under an azalea 
plant in a flower bed in the Presidents front lawn on the Auburn University Campus, several plants 
present, all on the root ball of the azalea, 8 Jul 1986, Diamond 2613 (AUA); Auburn University cam- 
pus in the Presidents lawn, in a flower bed with azaleas and spreading into the lawn, common 
weed in the lawn, in full sun and sandy soil, 25 May 1987, Diamond 3245 (GA); Auburn University 
campus in ne Presidents lawn, in a flower bed with azaleas and spreading into the lawn, common 


weed in th in full sun and sandy soil, 25 May 1987, Diamond 3246 (NCU). Mobile Co.: Bienville 
Blvd, near water tower, sunny, grassy, mod. Moist, infreg. herb., 10 Apr 1965, Deramus D378 (UNA); 
Bienville Blvd, near E end, sunny, moist, infrequent, 30 Apr 1967,D D1063 (GH, UNA). Pike Co.: 


Troy State University Campus, Lake Lagoona, edge of the lake, full sun, wet muck soil, Common, 
TION, R21E, Sec. 33,11 May 1993, Diamond 8615 (AUA). Tuscaloosa Co.: Common component of 
lawn, front (W side) of Biology Bldg, adjacent to Hackberry Lane, Univ. Alabama campus, Tuscaloosa, 
T/R/S:T21SR10W24, 12 May 1987, Burckhalter 991 (UNA). 
FLORIDA. Baker Co.: Plants trailing, banks of drainage ditch in nursery plots, 
mi S ei Mactienny on Fla 121, 16 May 1975, Heppner s.n. (FLAS); 2 mi S of US 90 on CR 123,0n left at 
end of CR 123, in hayfield, common, cows will not graze, 28 Apr 1992, Harvey s.n. (FLAS). Calhoun 
Co.: Locally abundant on spoil bank across floodplain, right-of-way paralleling highway bridge over 
Chipola River, S of Blountstown, 21 Jun 1982, Godfrey go NS). Eanes se : Pensacola, com- 
mon, sandy loam soil, low moist weedy grassy place, al Ice wage treatment 
plant, E side of Donelson St.,N of Main St.,4 Apr 1976, Buiehaler 3669 eG: ee S of Cedar 
St., E of Commendencia St., near railroad tracks, soil dark, rich, common in the Pensacola area, par- 
ticularly so in low moist sites, 12 Jul 1981, Burkhalter 7819 (FSU); Frequent, lawn weed, Tom Lane 
Drive, Pensacola, 20 Apr 1989, Dunavin s.n. (FLAS).Gadsden Co.: Frequent among grasses and sedges 
of right of way along highway 90 at Lanier Rd, 3 mi SE of Little River bridge, SW1/4 Sec 30,T2N, R2W, 
13 May 1987,Anderson 10507 (FSU). Gulf Co.: Mat-forming, in grassy lawn-like place, roadside park, 
just by Westarm Creek, N edge of Wewahitchka, 24 Apr 1978, Godfrey 76327 (FSU, GA); On moist 
slope above cypress pond along Rte. 22, ca. 1.5 mi W of Wewahitchka, flowers pink, 4 May 1982, 
Correll & Correll 53923 (NY, USF); N side SR 386, 2.95 mi W from jct with SR 71 (Wewa), damp, peaty 
sands in roadside ditch adjacent to disturbed pine flatwoods, 6 Jun 1987, Gholson with Godfrey & 
Baker 11830 (FLAS, GA); Shallow, broad ditch at edge of fallow field, by Fla. Rd 386, 3 mi from its jct 
with Fla. Rt 71 S of Wewahitchka, slender plants growing extremely densely, 6 Jun 1987, Godfrey 


1176 BRIT.ORG/SIDA 19(4) 


82413 with Gholson & Baker (FSU, GA); Weedy, amongst grasses and sedges in wet roadside depres- 
sion along Rte 22 (N side), 3.4 mi W of Wewahitchka and Rte 71,28 May 1991, Anderson 13425 (FSU). 
Indian River Co.: SW of Feldsmere, orange grove, S7 T31S R37E, 2 Nov 1988, Mears s.n. (FLAS, MO, 
USF). Jefferson Co.: Several weedy patches among flowers planted on W side of Jefferson County 
courthouse, downtown Monticello, 16 Jun 1980, Nelson 1637 (FSU). Leon Co.: In peaty mucky soil, 
small clearing of clay piel shores of Lake Munson at the Public Boat Landing, plants also 
extending into a small law wed area adjacent, 22 April 1974, Godfrey 73437 (FLAS, FSU, NCU, 

USF); Small plant ae Biucseny bush planted at 3237 Sharer Road last summer) actively spread 

ing into lawn,N side of Tallahassee, 26 Mar 1976, Anderson 4129 (FSU, NCU, USCH, USF); Small Sines 
of herbs along hurricane fence behind Florida High School, Tallahassee, 26 Oct 1977, Nelson 949 
(FSU, GA); Plants essentially past fl ring; frequently seen along ditch margin just W of Tully Gym, 
FSU campus, this ditch Buedlowels its banks nearly every time it rains, 1 Aug 1981, Nelson 1940 
(NCU); In enriched soil of vegetable garden, Tallahassee, 29 Apr 1982, Godfrey 79702 (MO); Shrub- 
bery bed on the N side of W Tennessee St., North Florida Education Credit Union, 1 Apr 1983, Le- 
onard 8113 (FSU); Frequent in muddy soil [...] around edge of Lake Ella, just E of N. Monroe St. in 
Tallahassee, 20 May 1985, Anderson 8085 (FSU); Common in lawn of low, shaded area along Pensacola 
St. just W of Dupree St.in Tallahassee, 11 Jun 1986, Anderson 9535 (FSU); Frequent amongst weedy 
growth along Sugar Creek near (edge of shopping mall) near Sharer Rd, just N of Hwy 27 in Talla- 
hassee, 13 Apr 1990, Anderson 12620 (FSU); Lake Jackson in Sunset Park off Old Bainbridge Rd,on a 
wet bank at the edge of a marsh at the lake, 21 Oct 1992, Godfrey 84438 (A, NY); In sand of sparse 
lawn bordering Blountstown Hwy between Tennesee St. and Pensacola St. in Tallahassee, 4 Apr 
1995, Anderson 15412 (FSU, USCH); Elinor Klapp-Phipps Park (W of Meridian Rd, N of Tallahassee, T2N, 
R1W, Sec 35), frequent in disturbed sites bordering open fields of Lake Jackson at edge of hard- 
woods, S of marker 7,23 Apr 1996, Anderson 16264 (FSU). Nassau Co.: White Oak Plantation border- 
ing on the St. Mary's River ca. 10 mi NW of Yulee, near Animal Science Building and the nearby 
ditches, ditch and shady roadbank, 25 Oct 1997, Wilbur 69802 & Moore (DUKE); White Oak Plantation 
bordering on the St. Mary's River ca. 10 mi NW of Yulee, borrow pit along Short-Cut Road near the 
Perimeter Road in the SW corner, highly disturbed area, erect herb, only clump seen, 28 Oct 1997, 
Wilbur 69972 & Moore (DUKE); White Oak Plantation bordering on the St. Mary’s River ca. 10 mi NW 
of Yulee, in the woods behind the Animal Science Building and along a sandy drainage ditch, 25 
Apr 1998, Wilbur 70456 (DUKE, GA);White Oak Plantation bordering on the St. Ne River ca.10 mi 
NW of Yulee, ditch along the E side of the road that encircles the Necroscopy-Biodiversity Building, 
abundant in shallow ditch, 25 Apr 1998, Wilbur 70482 & Wilbur one GA); White ak Plantation 
bordering on the St. Mary’s River ca. 10 mi NW of Yulee, disturbed sites near the Necroscopy Build- 
ing and the slopes W of the road on the western side E of Spare Lake, 25 Apr 1998, Wilbur 70517 
(DUKE); White Oak Plantation bordering on the St. Mary’s River ca. 10 mi NW of Yulee, borrow pit 
near the South Boundary, abundant, 29 Apr 1998, Wilbur 70716 & Wilbur (DUKE, USCH). Orange Co.: 
Soil wet, organic, common, weed in fernery at 2810 Union St., Zellwood, 23 Jun 1990, Boone s.n. 
(FLAS). Santa Rosa Co.: Wayside Park, S side I-10, E side Escambia Bay, near Pensacola, sandy-loam 
soil, which was probably hauled in, 16 Sep 1977, Gholson 6681 (FSU); Milton, between bowling alley 
and Odom Fiberglass (2944 Avalon Blvd) on Santa Monica St.,E at end of road near Avalon Blvd, in 
landscape, frequent, invasive, this plant is vigorous, ae easily, Competes with shrub plantings, 
14 Jun 1994, Hockett s.n. (FLAS). Volusia Co.: Weedy patch within Canna planting, median of Willow 
Run Dr, Port Orange, 27 Nov 1987, Nelson 6273 with ie (USCH). Walton Co.: Among shrubbery, 4 
mi S of Freeport, 25 May 1971, Davis 15823 (FLAS). 

GEORGIA. Chatham Co.: Top of bank on drainage ditch in open low flats of Savannah River, 
just E of Savannah, 22 Apr 1978, Duncan 30489 (GA); Dense colonies in depressions, Hunter pace 
Old Louisville Rd, uncommon, 20 Sep 1979, Duncan 30622 (GA). Colquitt Co.: Moist bank of roa 
side ditch,6 mi S Sac al set US 319, 30 Apr 1978, Godfrey 76344 (FSU). Cook Co.: 1 mi W = 
75 on S.R.37,growin ide between pavement and wet ditch, 14 May 1991, Mears s.n. (USF). 


a 


KRINGS AND NEAL 1177 


Grady Co.: In alluvial mud of ditch, at roadside at Barnett’s Creek, by US Rt 84, E of Cairo, 28 Apr 
1980, Godfrey 77747 (FSU); Roadside embankment, along US #84, ca. 5 mi E of Cairo, 11 Apr 1994, 
Henderson 94-76 (MO); Low roadside, along GA #111,ca.1 mi S of Reno, 20 Aug 1994, Henderson 94- 
850 (MO). Liberty Co.: Ft. Stewart Military Reservation, Training Area D-3, Grid Coor. 385278, 100m 
SE of jct FS 38 and FS 40, loamy soil on bank ] Mill Creek (channelized), locally common, 25 Aug 
1992, Carter 10403 & Lusk (GA). Lowndes Co.: Weedy area along Two Mile Creek in McKey Park be- 
nN. Patterson and N. Oak Streets in a Coastal Plain Province, 15 May 1980, Faircloth 
a (FSU, GA, NCU); US Hwy 84, 2.1 mi E of the Brooks Co. line, on the S side Be road, roadside, 
abundant, moist sandy ditch in full sun, 22 May 1988, Diamond 4801 (AUA). Muscogee Co.: Colum- 
bus, 6219 Windsor Dr, Bermuda grass lawn, full sun, sandy loam soil, 2 Oct 1987, Brantley s.n. en 
Sumter Co.: Locally fairly common on ditchbank on W side of Brady Rd, just N of jct Bone Rd, ca 3 
(air) mi SE of Americus, Sumter Co., elev. ca 350 ft., main soil herecas is Irvington sandy loam (now 
moist to wet), near fencerow of chinaberry, opposite cow pasture and farm pond, Repl. 6 (Pollen 
Sample No. 111), 2 May 1997, Norris 6829 (GA). Thomas Co.: In shallow water of drainage ditch at 
side of road and just outside of fence of pasture, just inside city limits of Thomasville (to W), by US 
Rte 84, 13 Apr 1984, Godfrey 81178 with Gholson (FSU). 
QUISIANA. Allen Parish: Infrequent in pine forest off US 165, ca. 2 mi N of Oberlin, 6 May 
1982, Allen 11947 (FSU). East Baton Rouge Parish: Along Christian St. at Hollydale Town Houses 
and at Perkins Road and I-10 overpass in Baton Rouge, 5 Nov 1993, Thomas 138578 (NY). Lincoln 
Parish: Ditchbank at a nursery on the S side of US Hwy 80, 1.2 mi E of Louisiana Hwy 33, just outside 
the eastern edge of Ruston city limits, sandy loam, common, 28 Apr 1988, Boyd & Boyd 3095 (LSU). 
Livingston Parish: Roadside ditch along eastbound |.H. 12, 2.3 miW of LA 43 exit, W of Hammond, 
3 May 1989, Urbatsch 5445 with Cox (LSU); common along roadside of local rd at jct with I-12, Sat- 
suma, LA, 16 Sep 1996, Montz 8323 (LSU); common in rear of yard at 25088 Hwy 42, Holden, LA, ca. 3 
mi W er unetion of Hwy 42 and Hwy 43, 2 Jul 1998, Montz 8894 (NO). Ouachita Parish: Large 
population between edge of road and end of culvert on first canal N of the entrance to Chennault 
Park E of ienioe 29 May 1997, Thomas 154251 with Gabel (GA, NO, NY, USCH). Saint Charles Parish: 
Bonnet Carre Spillway, rare only 3 plants noted and 2 collected, N of Hwy 61 near E guide levee at 
old Recreation facility, 9 Dec 1995, Montz 8098 (LSU). Saint Tammany Parish: Interstate 12 at the F 
bound rest area near Covington at the Tchefuncte River, 13 May 1999, Alford 1967 with McDaniel 
(DUKE). Tangipahoa Parish: Southeastern Louisiana State University campus, common alon 
lumbus Dr near University Center, Hammond, LA, 9 Apr 1994, Montz 6607 (LSU, NO); Southeastern 
Louisiana University campus, locally common only in this area near Physical Plant & Services Shops 
along N Oak St., Hammond, LA, 28 Jan 1995, Montz 7338 (LSU, NO); Southeastern Louisiana Univer- 
sity campus, infrequent at Horticulture Complex along N Oak St., Hammond, LA, 8 Apr 1997, Montz 
8607 (LSU, NO). 
NORTH CAROLINA. Chatham Co.: Specimen from potted plant collected 4 Dec 1998 from 
2925 NC Hwy 751 and grown in North Carolina State University greenhouse by Joseph Neal, 1 Dec 
2000, Ang: & Neal 30) (NCSC); 2925 NC Hwy 751; From fertilized pots of shrubs kept in enclosed 
lar irrigation,6 Dec 2000, Krings & Neal 366 (BHO, NCSC, USCH : 
SOUTH CAROLINA. Charleston Co.: Jenkins Nursery, Rt.17-S, Charleston, submitted by Joann 
Hubbard for ID (Dept. Horticulture, Clemson Pa alae d weed in boxwood pots and established in 
ditches and nursery, first seen 3 years ago, | me in from Florida, 2 Feb 1989, Hill 20176 (GH, 
MO, NY, USF). Jasper Co.: Ditch near US 1 7A, Savannah Wildlife Refuge, 1 mi NW of SC 170 jct, 20 Apr 
1974, Radford 46268 (FLAS, FSU, GA, GH, NCU, Me USCH, USF). Richland Co.: Moist lawn, 6221 
Monticello Road, Columbia, Aug 1977, Rayner 1013 (USCH); Blossom St. side of Hardees, opposite 
Coliseum, Columbia, fairly abundant, sprawling i intertwined within branches of low shrub- 
bery, 10 Dec 1992, Nelson 13693 (USCH); Commonly matted and tangled together in some abun- 
dance on wet, grassy ground, 1900’WSW of Paton Stadium, 1* Brigade Training Site (Black Lions), ca. 
5 mi E of downtown Columbia, elev. 225’ 11 Jun 1999, Nelson 20597 (USCH). Sumter Co.: Shaw Air 


1178 BRIT.ORG/SIDA 19(4) 


Force Base, Sumter, very abundant and tangled together on wet sandy ground with Cyperus 
ce aie at edge of northernmost of three golf course ponds, along causeway end of the pond, 
July 1994, Nelson 15753 (GH, USCH). 
TEXAS. Liberty Co.: met ae along Hwy 321-105 ca.3 miE or Cleveland 23 Mey 1987, Brown 
11136 (TEX).Mont w Covey, in wet ditch immediately ac to Lake 
Houston State Park, 9 Dec. a os 371 (NCSC, TEX, TAMU). 


ACKNOWLEDGMENTS 


We are grateful to the following herbaria and their staffs for kindly providing 
loans of or access to their collections: AUA, DUKE, GA, GH, FLAS, FSU, LSU, 
MO, NCU, NO, NY, UNA, USCH, USE TEX. In addition, we thank Guy Nesom 
for thoughtful review of the manuscript. 


REFERENCES 


ALLEN, C. 1983. Murdannia keisak (Hassk.) Hand.-Mazz. (Commelinaceae), Bothriochloa 
hybrida (Gould) Gould (Poaceae), and Scutellaria racemosa Pers. (Lamiaceae) new to 
Louisiana. Sida 10:189-190,. 

EpLinG, C. 1942. The American species of Scutellaria. Univ, Calif. Publ. Bot. 20:1-141. 

Goorrey, R.K.and J.W.Wooren. 1981. Aquatic and wetland plants of the southeastern United 
States: Dicotyledons. University of Georgia Press, Athens. 

Jones, 5.B., Jr.and N.C. Coie. 1988. The distribution of the vascular flora of Georgia. Depart- 
ment of Botany, University of Georgia, Athens. 

Krat, R. 1973. Some notes on the flora of the Southern states, particularly Alabama and 
Tennessee. Rhodora 75:366—-410. 

Kral, R.1981.Some distributional reports of weedy or naturalized foreign species of vas- 
cular plants for the southern states, particularly Alabama and middle Tennessee. Cas- 
tanea 46:334-339. 

Krinas, A,and J.C. Neat. 2001.A Scutellaria (Lamiaceae) new to North Carolina and a key to 
the small-flowered Carolina congeners. Sida 19:735-739. 

Leonarb, E.C.1927.The North American species of Scutellaria.Contr.U.S. Natl, Herb. 22:703- 


Massey, J.R. 1975. Fatoua villosa (Moraceae): Additional notes on distribution in the south- 
eastern United States. Sida 6 (2):116. 

PENLAND, C.W. 1924. Notes on North American Scutellarias. Rhodora 26:61—79. 

Raproro, A.E., H.E. AHLes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
The University of North Carolina Press, Chapel Hill. 

Reep, P.B., Jk. 1988. National list of plant species that occur in wetlands: national summary. 
U.S. Fisheries and Wildlife Service Biol. Rep. 88 (24):125. 

Tose, J.D., KC. Burks, R.W. Cantrett, M.A. GARLAND, M.E. Sweeney, D.W Hatt, P Wattace, G. ANGLIN, G. 
Netson, J.R. Cooper, D. BIckNeR, K. Gi-Bert, N. AYMOND, K. GREENWOOD, and N.RaymMonp. 1998. Florida 
wetland plants: an identification manual. Florida Department of Environmental Pro- 
tection, Tallahassee. 


Wa 


KRINGS AND NEAL 1179 


Uva,R.H.,J.C. Neat, and J.M DiTomaso. 1997.Weeds of the Northeast.Cornell University Press, 
Ithaca, NY. 

Wacter, H. 1985. Vegetation of the earth and ecological systems of the geo-biosphere. 
Springer Verlag, Berlin. 

Wunbertin, R.P. and B.F. HANseN. 2000. Atlas of Florida vascular plants (http:// 
www.plantatlas.usfiedu/). [S.M.Landry and K.N. Campbell (application development), 
Florida Center for Community Design and Research.] Institute for Systematic Botany, 
University of South Florida, Tampa. 


1180 BRIT.ORG/SIDA 19(4) 


Book REVIEW 


Mary F Iris. 2000. Gardening in the Desert: A Guide to Plant Selection & Care. 
(ISBN 0-8165-2057-7, pbk.). The University of Arizona Press, 355 S. Euclid 
Ave, Suite 103, Tucson, AZ 85719. (Orders: www.uapress.arizona.edu, 520- 
621-1441, 520-621-8899 fax). $17.95, 210 pp, 20 halftones, 6" x 9". 

Gardening in the Desert is an excellent book for anyone interested in desert gardening. It provides a 

starting point for anyone who has never encountered some of the unique challenges of desert gar- 


dening and saaees ina This book could not be more timely for an area of the United States that has 
h fol] 1| ignificant i ind d | 
g § 
Waterisata premium in] lowd l utilizing pl dapted to dry habi hould 
be near the top of desert inhabitants’ conservation prionaes 
The book is divided into 1] Sa ee begint Be itions of desert gardening. This first 
Snape addresses the seasons of the low deserts of He ona! mae eonienvend texture, as spleen 
annual | ion i 
according to: general growth fone Gece desert ee sane perenmasi acca apices ran- 
nuals, vines, cacti, agaves, other succulents, and distinctive desert plants. Each chapter begins with 


pertinent information on the growth form. The chapter on wildf ae and other annuals, for ex- 
ns with recommendations on propagating annuals, suggests soil amendments and a 
vises on proper seed collection. After a short eae tion, eac : chapter lists a sia of ee 


species to include in your garden. Each plant thi names, a short 
description of the plant ees height, lee color es pheno information on water and soil 
requirements, as well as pruning advice for woody plants and other perennials. In the woody plants 
chapter, Laie sen of tall or vigorous growth looms is ee st 

eral chapters in particular would be useful e desert ee The chapter on cacti 
seems es spesully practical. Within the chapter is information on how to properly transplant cacti, 
how and when to water cacti, and how to avoid frost damage in susceptible species. Numerous ex- 

ples of native Opuntia are mentioned in the book, but sadly the author mentions only Mammill- 

aria and Coryphantha in passing and has no specific suggestions for planting. This is unfortunate 
since members of these two genera are aesthetically pleasing and lack the glochidia which make 
Opuntia gardening poten ally oe a chapter on agaves is MOL iy because a short horti- 


cultural history i ler on their etl | heritage. In the last cha 


_— 


ter, as ae die wicees unusual plants that do not ae fit into a general growth form, These 
esperaloes, and yuccas, among others. A preventative approach to pest and dis- 


a) 


ease contd is stressed throughout the book, thus decreasing the need for toxic ea cides 
though twenty figures provided in this book, they are black and white. The black api white 
photos of different garden settings do little justice to the plants that thrive in desert environments. 
Color figures would have ek enhanced a Book ane wenld have prowided further pee of 
A] oosing 


the natural beauty of these dry I 
garden plants wisely, it cannot < overstated that great care should be taken salient nie exotic 
species. Many non-natives from Africa, Australia, and the Mediterranean, including Senna 
artemesiodes and Dimorphotheca sii mentioned i in the book, can become pests in a matter of a 
few growing seasons and these h habitat for the native flora. There are numerous 
native choices that should be considered first before planting exotics in any desert garden. 

all, ad isa valuable Pook io anyone interested in desert gardening and landscape prac- 


tices. Irish’ ling this book both educational and entertaining. The 
book provides her experience eand ne eats desert eee and these words of wisdom 
ould be most useful to amateur and master gardeners alike. It would make a suitable addition to 


any desert gardener’s library.—Amy Trauth eles Fort Worth, - U.S.A, amy_trauth@yahoo.com 


SIDA 19(4): 1180. 2001 


CAESALPINIA BONDUC (FABACEAE) NEW TO LOUISIANA 


William G.Vermillion 
U.S. Fish and Wildlife Service 
646 Cajundome Blvd., Suite 400 
Lafayette, LA 70506, U.S.A. 


A recent collection of Caesalpinia bonduc (L.) Roxb., a sea bean, from a coastal 
barrier island in Lafourche Parish, southeastern Louisiana, is apparently the 
first record for the state. On 27 April 2001, material of C. bonduc was collected 
on the western end of East Timbalier Island, a barrier island bordering Timbalier 
Bay. The plant was growing adjacent to an interior canal bordered by a low levee 
of rocks (riprap). At the time of collection the plants had mature dehiscing fruits, 
with gray seeds, but no flowers. Identification of the plant was verified by com- 
parison with specimens from Florida at LAF 1 also saw the species at another 
location in lower Lafourche Parish on 5 April 2001, but no specimen collected. 
The plant was ona canal bank approximately 0.6 miles east of Louisiana High- 
way l,ca.1.4 miles south-south east of the La. Hwy. | bridge over Bayou Lafourche 
at Leeville, and ca. 1 mile northeast of the heliport located between La. Hwy. 1 
and Bayou Lafourche, south of the Leeville bridge. The latitude and longitude, 
recorded with a hand-held global positioning unit, is latitude 29°13'36.87, lon- 
gitude 90°12’08.77 (Fig. 1). 

The species is not shown for Louisiana by Isely (1990), Thomas and Allen 
(1998), or the USDA, NRCS database (2001). Prior to its find in Louisiana, it was 
known in North America north of Mexico only from central and southern pen- 
insular Florida (Wunderlin 1998). It ranges widely around the tropics and sub- 
tropics of the world. A voucher specimen has been deposited at LAF, with a 
duplicate at NLU. 

Voucher specimen: LOUISIANA. Lafourche Parish: Sprawling over riprap bordering canal on inte- 
rior of west end of East Timbalier Island, a coastal barrier island, forming a somewhat elongate 
colony ca. 10 m long, 27 Apr 2001, Vermillion s.n. (LAF, NLU). 


ACKNOWLEDGMENTS 


Ithank William D. Reese (LAF), R. Dale Thomas, and Alex A. Lasseigne for sug- 
gestions on the manuscript. 


REFERENCES 


lsery,D. 1990.Vascular flora of the southeastern United States, Volume 3, Part 2, Leguminosae 
(Fabaceae). University of North Carolina Press, Chapel Hill. 
THomas, R.D.and C.M. Atten. 1998. Atlas of the vascular flora of Louisiana, Volume III: Dicoty- 


SIDA 19(4): 1181 — 1182. 2001 


1182 BRIT.ORG/SIDA 19(4) 


East Timbalier acacia 43 
27 April 2001 


4 Miles 


Fic. 1. C Ipinia bonduc | ions in Lafourche Parish, Louisiana 


ledons, Fabaceae—Zygophyllaceae. Louisiana Department of Wildlife and Fisheries, 
Natural Heritage Program, Baton Rouge. 

USDA, NRCS. 2001. The PLANTS database, Version 3.1 (http://plants.usda.gov/plants).Na 
tional Plant Data Center, Baton Rouge, LA 

WUuNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. University of Florida Press, 
Gainesville. 


HOUSTONIA LONGIFOLIA (RUBIACEAE): NEWLY 
DOCUMENTED FOR THE FLORA-OF TEXAS 


Jason R. Singhurst W.C. Holmes 
Wildlife Diversity Program Department of Biology 
Texas Parks and Wildlife Department Baylor University 
ustin, 1X 78704, U.S.A. Waco, Texas 76798-7388, U.S.A. 
Jason.singhurst@tpwa state. tx.us walter_holmes@baylor.edu 
ABSTRACT 


Houstonia longifolia Gaertn. (Rubiaceae) is reported as new to Texas. 


RESUMEN 


Houstonia longifolia Gaertn. (Rubiaceae) se cita como nueva de Texas. 


Houstonia longifolia Gaertn. var. tenuifolia (Nutt.) A. Wood was included in the 
checklist of the vascular plants of Texas by Jones et al. (1997). Its inclusion ap- 
pears based upon the citation by Hatch et al. (1990) of Hedyotis nuttalliana 
Fosberg as occurring in the Pineywoods, Gulf Prairies and Marshes, Cross Tim- 
bers and Prairies, and Edwards Plateau vegetational areas of the state. Ina mono- 
graph of the genus, Terrell (1996) treats both Houstonia longifolia var. tenuifolia 
(Nutt.) A. Wood and Hedyotis nuttalliana as synonyms of Houstonia longifolia. 
He further comments (p.67) that Houstonia longifolia is “quite variable and wide 
ranging, and by far the most difficult species in the genus.” Ultimately he con- 
cludes that the only reasonable taxonomic solution is not to recognize varieties 
or subspecies. Instead, Terrell recognizes “population groups.” The “Ozark- 
Ouachita Group” of Houstonia longifolia is known from southeast Oklahoma 
and southwest Arkansas, barely 20 km north of Texas. The “Tenuifolia Group” 
(formerly H. longifolia var. tenuifolia) is restricted, according to Terrell, to west 
Florida and central Alabama northeastward to Virginia and West Virginia. 

The plant - red to by Jones et al. (1997) and Hatch et al. (1990) is, under 
Terrell’s 1996 tion, Houstonia longifolia, however, this species has not 
previously been found in Texas (Terrell 1996). At this time the identity of the 
plant or plants referred to by Hatch et al. 1990) and Jones et al. (1997) is not 
clear, but is most likely Stenaria nigricans (Lam.) Terrell var. nigricans a species 
widely distributed in Texas. The collection is the first validation of Houstonia 
longifolia from Texas. 


Specimens cited: TEXAS. Red River Co.: W side of Tanyard Creek, 0.6 mi NE of Woodland Cemetery, 
1.2 mi NE of Woodland at jct.of Hwy 195 and Hwy 410, 19 May 1999, Singhurst 7902 (BAYLU, TEX, US). 


The specimens were collected froma thinly wooded hardwood forest dominated 


SIDA 19(4): 1183 — 1184. 2001 


1184 BRIT.ORG/SIDA 19(4) 


by Quercus alba L., Q. shumardii BuckL, Q. muhlenbergii Englem. (Fagaceae), 
Carya tomentosa (Poir.) Nutt. Juglandaceae), and a few scattered trees of Acer 
saccharinum L. (Aceraceae). The site is characterized by an 8-20% slope with a 
sparceness of vegetation. Isolated clumps of Houstonia longifolia were growing 
ona steep bluff overhang above Tanyard Creek and among sandstone outcrops 
on the slope. 

Houstonia longifolia isa perennial, sometimes with a basal rosette during 
flowering, and a height of up to 45 cm. Other than H. purpurea, all east Texas 
Houstonia species are annual, lack basal rosettes at flowering, and are of con- 
siderably shorter height (up to 15cm, but generally less than 10 cm). Houstonia 
purpurea, known in Texas only from Newton Co. (Terrell 1996), while similar 
to H. longifolia, is distinguished by it broader median cauline leaves. A similar 
species is Houstonia ouachitana (E.B.Smith) Terrell of southwest Arkansas and 
southeast Oklahoma. That species is distinguished by the densely pilose hairs 
on the inner surfaces of the corolla. These hairs are 0.5-1.2(-1.8) mm, while the 
inner corolla surfaces of Houstonia longifolia are glabrous or with hairs less 
than 0.5mm long (Terrell 1996). 


ACKNOWLEDGMENTS 


We are indebted to Tom Wendt (TEX-LL) for some assistance and information 
and to Edward E. Terrell (MARY) for confirmation of our determination. Also,a 
special thanks to Mrs. Betty Jo Swindle for access to her Red River County prop- 
erty. John Pruski (US) read and commented on preliminary drafts of the manu- 
script. 
REFERENCES 
Hatcn, 9.L., K.N. GANoHI, and L.E. Brown. 1990. Checklist of the vascular flora of Texas. Texas 
Agric. Exp. Sta. Pub. MP-1655, Texas A&M University, College Station. 


Jones, S.D., J.K. Wiper, and P.M. Montcomery. 1997. Vascular plants of Texas:a comprehensive 
checklist including synonymy, bibliography, and index. University of Texas Press, Aus- 


TerRELL,E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monogr. 48:1- 


NEW RECORDS IN PSEUDOGNAPHALIUM 
(ASTERACEAE: GNAPHALIEAE) FOR THE UNITED STATES 


G.L.Nesom 


Botanical Research Institute of Texas 


509 Pecan Street 
Fort Worth, TX 76102-4060, U.S.A. 


ABSTRACT 
State and namonal distribution records are reported for species of Pseudognaphalium. New for the 
US.A P jali (Arizona, California, Colorado, New Mexico, and Texas) and P. atte (Ha- 
waii). P l hali is fi reported for Texas, and the nearly cosmopolité an weed P. 


luteoalbum is is re — ted for Texas and New Mexico. An early collection for P. stramineum in New York, 
apparently asa wat in Manhatas, is cieg : brief characterization is provided for each species, 


nda P jaliscense. A record of the occurrence of P helleri 
in india is incorrect, based ona allecnon of P. macounii. 


RESUMEN 


Se citan los ceases oe CSneuGO estatal y nacional de las especies de Pseudognaphalium. Son 
nuevas para U.S.A. Pj California, Colorado, Nuevo México, y Texa s) y Patt enuatum 


(Hawaii). Pseud hali izonic por primera vez para Texas, y la mala hierba casi 


cosmopolita P. luteoalum se cita para Texas y pueNe cae Se cita una coleccién previa de P. 


stramineum en Nueva York, aparentemente como abandonada en Manhattan. Se ofrece una breve 
caracterizacion de cada especie, se citan Sane a 7 se escoge un lectotipo para P jaliscense. Una 
cita de la presencia de P hellerien Indiana Sta, y esta basadaen una colecta de P macounii 


In connection with taxonomic studies of Gnaphalieae of Central America, 
Mexico, and the United States, various new distribution records have come to 
light for the United States. First reports for Pseudognaphalium species are docu- 
mented here. 


Pseudognaphalium arizonicum (A. Gray) A. Anderb., Opera Bot. 104:147. 1991. 
Gnaphalium cine A. Gray, Proc. Amer. Acad. Arts 19:3. 1883. TYPE: UNITED STATES. 
ARIZONA. [Cochise Co.], Huachuca Mts., “near Fort Huachuca, along exsiccated beds of streams,” 
[1882,] J.G. Lemmon ae type: GH)). 


Plants annual to short-lived perennial herbs from a slender, lignescent taproot. 
Stems 2-5 cm tall, loosely but densely woolly-tomentose, the hairs from thick, 
viscid-appearing, persistent bases, commonly with reddish or purplish cross- 
walls or segments, eglandular. Leaves ascending, linear-oblanceolate to linear- 
lanceolate, 2-6 cm long, 3-7 mm wide, prominently decurrent, not basally 
ampliate, tomentose above and beneath, weakly or not at all bicolored, vestiture 
like the stems. Capitulescence diffuse-corymboid or of only a few tight clus- 
ters. Capitula 4+.5-5 mm high; phyllaries usually brownish to tawny, rarely 


SIDA 19(4): 1185 — 1190. 2001 


1186 BRIT.ORG/SIDA 19(4) 


slightly rosy, shiny, narrowly triangular-acute, subequal. Pistillate florets 25-) 
30-49(-67 in southwestern Mexico). Bisexual florets (1-)3-6(-10 in southwest- 
ern Mexico). Cypselae longitudinally ridged, densely and minutely papillate. 

Specimens examined. UNITED STATES. TEXAS. Brewster Co.: Chisos Mts.: Boot Creek area, 8 Sep 
1933, Wolff 4609 (BRIT); Emory Peak, 14 Aug 1931, Mueller 32020 (GH, TEX); Boot Spring, 22 Aug 1931, 
Mueller 8208-09 (GH); Big Bend Natl. Park, N branch of Cattail Canyon, igneous soil, 6500 ft, 7 Oct 
1967, Warnock 21281 (SRSC); along trail to South Rim above Boot Springs, 6500 ft, sparse in igneous 
soil, 10 Sep 1950, Warnock 9788 (SRSC). Jeff Davis Co.: Davis Mts.:Mt. Livermore, 29 Sep 1935, Hinckley 
s.n. (GH, TEX); 6.8 mi N on road to Davis Mountain Resort on Tomahawk Trail, which starts at Hwy 
166, 11.0 miW of Hwy 17,15 Sep 1991, Sherman, Noyes, and Brant 194 (MO); upper tale in Madera 
Sales on Mt. Livermore, 7500 ft, 11 Sep 1947, Warnock and Hinckley 7408 (SRSC).No county speci- 
fied: “expedition from western Texas to El Paso, New Mexico,” May-Oct 1849, Wright sas (GH) and 
Wright 395| it (GH). 


Pseudognaphalium arizonicum is widespread in Mexico (from Chiapas north- 
ward through Veracruz and Michoacan to Nuevo Leon and Chihuahua) and 
has been previously known in the U.S.A. only from Arizona (Cochise, Pima, and 
Pinal counties). It has been collected in a wide range of habitats, from agricul- 
tural land to oak and pine woodlands above 1600 meters elevation. It is superfi- 
cially similar in its narrow, non-bicolorous leaves to Pseudognaphalium 
stramineum (Kunth) A. Anderb., but the latter has mostly non-decurrent leaves, 
light yellowish, ovate to obovate phyllaries, and (75-)160-200 pistillate florets 
and (9-)16-28 bisexual florets. 


—_— 
— 


Pseudognaphalium attenuatum (DC.) A. Anderb., Opera Bot. 104:147. 1991. 
Gnaphalium attenuatum DC,, Prodr. 6:228. 1838. Type: MEXICO: “In Mexico circa Tampico de 
Tamaulipas,” Berlandier 70 (G-DC fiche!). 

Plants annual herbs, taprooted. Stems 4-15 dm tall, persistently and closely 

white-tomentose, eglandular. Leaves narrowly elliptic to lanceolate, (3-)4-9(- 

12) cm long, 5-10(-13) mm wide, usually long-tapering on both ends, sessile, 

not at all clasping or decurrent, strongly bicolored, glabrescent above and often 

shiny, eglandular but often appearing roughened with persistent, thick, gland- 
like trichome bases. Capitulescences compactly to diffusely corymboid to 
rounded-paniculate. Capitula 5-6 mm high; phyllaries tawny-whitish, very 
rarely pinkish. Pistillate florets 35-41 (Michoacan westward) or 49-59 (Edo. 

Mexico, Veracruz, and southward). Bisexual florets 2-4 (Michoacan westward) 

or 5-8 (Edo. Mexico, Veracruz, and southward). Cypselae with 4-8 shallow, lon- 

gitudinal ridges, not papillate. 

Specimens examined. UNITED STATES. HAWAII. [Hawaii Co.:] near Kohala Blvd., makai of Belt Road, 

Kahuku, Kau, ubiquitous at 1500 ft in small, sterile kipuka, 10 Jan 1981,0.&/. Degener 35,082 (GH, 

SMU, TEX). 

San onankeliaa attenuatum is widespread in the southern half of Mexico 

and through most of Central America; it also occurs in the West Indies (at least 

in Cuba, Puerto Rico, and Jamaica). Over this range, it occurs at elevations of 


= 


NESOM, 1187 


60-2200 meters in fields, secondary vegetation, woodlands of oak, oak-pine, 
and pine, and sometimes in subtropical forests. Pseudognaphalium in Hawaii 
has previously been reported only as P sand wicensium (Gaud.) A. Anderb. (Wag- 
ner et al. 1990; Wagner et al. 1997). 


Pseudognaphalium jaliscense (Greenm.) A. Anderb., Opera Bot. 104:147. 1991. 
Gnaphalium jaliscense Greenm., Proc. Amer. Acad. Arts 39:96. 1903. TYPE: MEXICO. JALISCO: 
Gravelly soil near Guadalajara, 9 Aug 1902, C.G. Pringle 8628 (LECTOTYPE, designated here: 
GHl. ISOLECTOTYPES: CAS, Fl, LL!, MO!, NY, PH!). Greenman also cited Palmer 256 (GH!, US!) 
from near Guadalajara, Jalisco. 


Plants annual or biennial herbs, taprooted. Stems 3-7 dm tall, unbranched un- 
til the capitulescence, densely and persistently loosely woolly-tomentose-seri- 
ceous, eglandular to sparsely glandular. Leaves narrowly lanceolate to nearly 
linear, 3-10 cm x 3-6 mm, not auriculate-clasping, decurrent 4-8 mm, 
concolorous, closely sericeous-tomentose on both surfaces, the hairs arising from 
a thickened, gland-like base, sessile-glandular but the glandularity obscured 
by the dense vestiture. Capitulescences diffuse-corymboid or of a few tight clus- 
ters. Capitula 5-6 mm high; phyllaries opaque-white, strongly graduated, the 
apex keeled-apiculate. Pistillate florets (80- in U.S.A.)(L00-)138-180. Bisexual 
florets (9- in US.A.)02-)14-22(-30). Cypselae weakly 4(-6)-ridged, minutely 
but prominently papillate. 


Specimens examined. UNITED STATES. ARIZONA. Cochise Co.: Huachuca Mts., plain near Ft. 
Huachuca, 23 Oct 1926, Peebles et al. 34484 [or 3484 fide ARIZ sheet] (ARIZ, LL); Chiricahua Mts., Sep 
1882, Lemmon s.n. (US,WIS). Gila Co.: Pine, 7000 ft, 15 Aug 1937, Darrow s.n. (ARIZ- 2 sheets).Graham 
Co.: eastern foothills of the Galiuro Mts., ca 3.5 mi from end of High Springs Rd, and on foot into 
aa oak-pine-juniper woodland with gra and patches of manzanita-dominated a 
Sep 1989, Keil 21259 (ASU). Navajo ci NAW Co., Alona Hwy 60 at Forestdale, 16 A 66 
eee 66262 (ARIZ); immediately W of Forestdale, pine-juniper-oak woods, 6200 ft, 29 ts 
Bohrer 15366 (ARIZ); 1 mi NE of Forestdale Trading Post,6000 ft, ponderosa zone, 19 Sep 1975, Pinkava, 
Keil, and Lehto L19023 (ASU). Pima Co.: Santa Catalina Mts., 14 Apr 1881, Pringle 317 (GH); Rozemont, 
Santa Rita Mountains, 2 Sep 1905, Thornber s.n. (ARIZ). Santa Cruz Co.: Sonoita, 6 Sep 1940, Benson 
10422 (LL, MO); The Bog Hole, Meadow Valley, headwaters of the Santa Cruz River, 1530-1540 m, 26 
Aug 1976, Van Devender and McCarten s.n. (ARIZ). CALIFORNIA. Del Norte Co.: Hazelview Summit 
in Crescent City, Grants Pass Road, 24 Jul 1928, Kildale 5873 (LL).Marin Co.: Tomales Point, 5 mi from 
Inverness, grassy coastal slopes, 300 ft, 23 Jul 1939, Gould 852 (SMU). Mendocino Co.: near Point 
Arena Light, coastal prairie, 27 Jul 1949, Whitehouse 21713 (SMU). COLORADO. Jefferson Co.: Clear 
Creek Junction, dry open soil, 1590 m, 11 Sep 1920, Clokey 3943 (US). NEW MEXICO. Grant Co.: hills 
around the copper mines, [near Santa Rita, ca. 1-5 Aug], 1851, Wright 1283 (GH); Pinos Altos Mts., 8 
Sep 1880, Greene s.n. (US); Santa Rita del Cobre, 24 Sep 1880, Greene 460 (GH). Lincoln Co.: Lincoln 
Natl. Forest, Smokey Bear R.D., FR 120, grama-juniper flat, 6900 ft, 10 Sep 1990, Pase 3575 (NMC), 
White Mts., 25 Aug 1907, Wooton and Standley s.n. (NMC);White Mts., Gilmore Ranch, 11 Aug 1897, 
Wooton s.n. (NMC).Mora Co.:Watrous, 27 Aug 1926, Arsene and Benedict 17488 (LL); Rio de las Casas, 
28 Jul 1927, Arsene 19159 (LL); Canyon Colorado Equid Sanctuary, 24 mi NE of Wagon Mound, open 
plain, 6020 ft, 27 Sep 1990, Smith 158 (NMC). San Miguel Co.: Vicinity of Las Vegas (8 mi SE), 1950 m, 
2 Sep 1926, Arsene 17373 (MO); Las Vegas, 2 Oct 1926, Arsene 17875 (LL). No county specified: N. 
Mex., 1851-1852, Wright 1283 (GH) and Wright 1284 (GH), mounted on the same sheet. TEXAS. 


1188 BRIT.ORG/SIDA 19(4) 


Brewster Co.: Glass Mts., ron Mountain, 20 Aug 1940, Warnock W219 (ARIZ, SRSC, TEX); Glass Mts., 
abundant locally in upper Green Valley, 18 Sep 1940, Warnock W218 (GH). Jeff Davis Co.: Davis Mts., 
Chitense Spring, 7 Sep 1918, Young 5.n. (GH, MO). 


Pseudognaphalium jaliscense is relatively common from southwestern to north- 
ern Mexico (Nayarit, Jalisco, Sonora, Chihuahua, Coahuila, and Nuevo Leon) 
into localities in the southwestern United States. In Mexico, it has been collected 
along roadsides or in disturbed sites, meadows or openings in pine or oak woods. 
It is recognized by its annual duration, long, narrow leaves with non-clasping, 
short-decurrent bases and barely bicolored, persistently tomentose-floccose 
surfaces, large capitula with relatively thick, white, dull-opaque phyllaries, and 
large numbers of pistillate and bisexual florets. The slightly atypical lower lim- 
its for pistillate and bisexual florets reflect counts from several of the U.S.A. 
collections, but in other features, these plants are correctly identified as P. 
jaliscense. 


a luteoalbum (L.) Hilliard & Burtt, J. E ue Soc., Bot. 82:206. 
1981. Gnaphalium luteoalbum L., Sp. PL 2:851. 1753. LECTOYPE (I rd & Burtt 1981) With- 
out data, Herb. van Royen s.n. (LECTOTYPE: L 900.286-294). 


Plants annual, taprooted or fibrous-rooted. Stems erect to basally decumbent- 
ascending, 15-40 cm, simple or with several branches from the base, loosely 
but persistently white-tomentose, eglandular. Leaves basal and cauline, basal 
narrowly obovate to subspatulate, 1-3(-6) cm x 2-8 mm, becoming smaller 
upwards and oblanceolate to narrowly oblong or linear, basally rounded to 
abruptly tapered, subclasping, short-decurrent or not, densely gray-tomentose, 
often glabrescent adaxially, eglandular. Capitulescences usually one to a few 
tight terminal clusters 1-2.5 cm wide. Capitula broadly campanulate, 3-4 mm; 
phyllaries ovate, silver-gray to yellowish, translucent, glabrous. Pistillate flo- 
rets 140-160. Bisexual florets 5-10, red-tipped. Cypselae weakly ridged, epider- 
mal surfaces minutely papillate; pappus bristles weakly coherent in an easily 
fragmented basal ring. 2n = 14 

Specimens examined. UNITED STATES. NEW MEXICO. Hidalgo Co.: Little Hatchet Mts., Playas, S 
side of town near city buildings in area where water from lawns flows into desert, 4450 ft, alluvial 
fan of mts., watered area, 31 Aug 1997, pS aay ee (SRSC— identified » Worthington as 
bio ca sh ium cf.luteoalbum’and confirmed a species by annotation by Turner in 1998) 
TEXAS. Kimble Co.: ca. 2 mi N of IH 10 along Hie a le perennial creek just N of Johnson 
Draw, 7 Apr 1 ae ie 99-83 (SRSC). El sae a Franklin 1.2 air mi NE from the top of North 
Franklin Mtn.,0.4 mi NW from Indian Spring, 4900 ft., canyon fe E, granite rock, along flow from 
spring (seep areas), 23 Apr 1995, Raia oe (SRSC). Parker Co.: Weatherford, Stuart's be Irs 
ery on Hwy 180,in pots, spreading to nursery, 24 Jun 2001, O’Kennon 14991 (BAYLU, BRIT, OKL, TEX). 


_ 


ad 


Pseudognaphalium luteoalbum is previously known in the U.S.A. from Florida, 
New York, and several western states (California, Nevada, Arizona, Utah, Oregon, 
and Washington) (Kartesz 1999). It apparently is native to Eurasia and isa nearly 
cosmopolitan weed, known from Europe, North America (including Mexico), 


NESOM 1189 


Africa, central and southeast Asia, Australia, New Zealand, and other Pacific 
regions. It is similar to P stramineum in its annual duration, loosely tomentose 
vestiture, concolorous, subclasping leaves, and capitula in small tight glomer- 
ules mostly 1-2 cm wide at the stem tips but differs in the following contrasts. 
1. Florets red-tipped, these tips visible through the translucent phyllaries; capitula 3— 

4 mm high; bisexual florets 5-10; leaves subclasping, usually slightly rounded at 

the base, not decurrent P. luteoalbum 
. Florets evenly yellowish, not red-tipped; capitula 4-6 mm high; bisexual florets 8- 

20 in western U.S.A.(19-28 in Mexico);leaves subclasping but not basally rounded, 

sometimes decurrent 1-2 mm P. stramineum 


= 


P i (Kunth) A. Anderberg, Opera Bot. 104:148. 1991 
TES) non (Kunth) W.A. Weber sie 0June). ona ian stramineum 
Kunth, Nov, Gen. & Sp. 4 [folio]:66. 1818. Type: MEXIC 1 Omitlan, 

{no date], Humboldt and Bonpland s.n. (HOLOTYPE: P es 


Gnaphalium chilense Sprengel, Syst. 3:480. 1826. 


Plants annual or biennial, taprooted. Stems 3-6(-8) dm, simple or several from 
the base, erect to basally ascending, loosely tomentose, eglandular. Leaves lin- 
ear to narrowly lanceolate, oblanceolate, or oblong, 2-8(-9.5) cm x 2-5(-10) mm, 
gradually reduced upward, subclasping but not basally ampliate, sometimes 
slightly decurrrent, concolorous, loosely but persistently gray-tomentose on 
both surfaces, eglandular. Capitulescences 1-few tight terminal clusters 1.5-3 
(-4) cm wide. Capitula subglobose, 4-6 mm; phyllaries ovate to obovate, hyaline 
and somewhat translucent, commonly distinctly yellowish with age. Pistillate 
florets 160-200. Bisexual florets 8-28, yellowish. Cypselae weakly i minutely 
papillate; pappus bristles separate, not basally coherent. 2n = 

Specimen examined. UNITED STATES. NEW YORK: [Manhattan,] ballast filling, at northern termi- 
nus of 8th Avenue, N.Y. Jul 1879, Addison Brown s.n 


Pseudognaphalium stramineum perhaps is broadly native from South America 
through Central America and Mexico to many states of the western United 
States and British Columbia. In the eastern USS., it apparently is adventive on 
the Atlantic coastal plain of Virginia, North Carolina, and South Carolina, where 
it occurs in sandy fields, flowering May through August. The collection from 
New York represents a waif, as P stramineum has never been reported by any 
floristic account of the northeastern United States. 


Pseudognaphalium helleri in Indiana 

Pseudognaphalium helleri Britt.) A. Anderb. was reported/ mapped to occur in 
Indiana by Mahler (1975). This apparently was based on the identification of a 
GH specimen, annotated by C.A. Weatherford and Wim. F Mahler as that species: 


UNITED STATES. INDIANA. Lagrange Co.: 2 Sep 1923, Deam 52927 (GH). 


This plant, however, has distinctly decurrent leaves and is Pseudognaphalium 


1190 BRIT.ORG/SIDA 19(4) 


macounti (Greene) Kartesz, which is abundant in Indiana. Other collections of 
P. macounii from Lagrange County also are housed at GH. 


ACKNOWLEDGMENTS 


lam grateful for loans from various herbaria (ARIZ, ASU, F GH, NMC, PH, SRSC, 
US, WIS) and to the staffs of GH, MO, NCU, TEX-LL, and US for help during 
recent visits. 


REFERENCES 


HiLuiard, O.M. and B.L. Burtt. 1981.Some generic concepts in Compositae—Gnaphaliinae. J. 
Linn. Soc., Bot. 82:181-232. 

Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the 
vascular flora of the United States, Canada, and Greenland. In: Kartesz, J.T. and CA. 
Meacham. Synthesis of the North American Flora, North Carolina Botanical Garden, 
Chapel Hill, NC. 

Manter, W.F. 1975. Typification and distribution of the varieties of Gnaphalium helleri Britton 
(Compositae-Inuleae). Sida 6:30-32. 

Wacner, W.L., D.R. Hersst, and S.H. Soumer. 1990. Manual of the flowering plants of Hawai'i. 
Univ. of Hawai'i Press: Bishop Museum Press, Honolulu, HI. 

Waaner, W.L., R.K. SHANNON, and D.R. Herest. 1997. Contributions to the flora of Hawai'i. VI. 
Bishop Mus. Occ. Pap. 48:51-65. 


ARTHRAXON (POACEAE: ANDROPOGONEAE) 


NEW TO SOUTH AMERICA 
LJ.Dorr a Nino 
Department of Systematic Biology (Botany), MRC- 166 BioCen 
Nationa ural History, UNELLEZ, Mesa ye Cavaca 
mithsonian Institution Portuguesa, VENEZUELA i 
Washington, DC 20560-0166, U.S.A. smiguel@cantv.net 


dorr.laurence@nmnh.si.edu 


ABSTRACT 


Arthraxon hispidus, native to the Old World and adventive in Hawaii, North America, Central 
America, and the Caribbean, is reported for the first time from South America. The records are from 


the Andes of Venezuela. Information also is provided on the earliest dates that this species is docu- 


mented to have appeared in different countries in the New World. 


RESUMEN 
Arthraxon hispidus, nativa del viejo mundo y adventicia en Hawaii, Norte América, Centro América 
y el Caribe, se cita por primera vez en Sur América. Los registros son de los Andes de Venezuela. 
Ge sy r ee . te er 1] pes 1 ] t 1 Pa a | Lae 
L 


de esta especie en diferentes paises del nuevo mundo. 


One of us (SMN) recently collected Arthraxon hispidus (Thunb.) Makino var. 
hispidus in Guaramacal National Park in the Andes of Venezuela and a search 
for additional material revealed that at least two other collections were made 
nearby in 1986 and 1990 (see voucher specimens). These are the first collections 
of the genus Arthraxon P. Beauv. (Poaceae: Andropogoneae) from South America. 
The species, A. hispidus, which is native to Asia, Africa, and Australia, was pre- 
viously known to be adventive in Hawaii, North America, Central America, 
and the Caribbean (van Welzen 1981). Arthraxon hispidus thus is now known 
to occur on all continents except Europe and Antarctica. 

The earliest record we can find of Arthraxon hispidus in the New Worldisa 
collection made in the Japanese Garden on the Centennial Grounds of Fairmount 
Park, Philadelphia in 1877 (Burk s.n., US). The species next was found on ship's 
ballast in Philadelphia in 1878 (Scribner s.n., US) and on ballast in Portland, 
Oregon about 1901 (Suksdorf 5316, US). The label data on a 1903 collection from 
South Carolina stated that the plant was introduced at Clemson College with seed 
from Japan (Newmans.n., US) and there is a specimen indicating that A. hispidus 
was cultivated by the U.S.D.A. at Arlington Experimental Farm in Virginia by 
1906 (Tidestrom s.n., US). These herbarium records support the inference that 
there were multiple introductions of this species into the United States. Irre- 
spective of the source or sources of these introductions, additional specimens 


SIDA 19(4): 1191 — 1193. 2001 


1192 BRIT.ORG/SIDA 19(4) 


in the United States National Herbarium (US) indicate that A. hispidus was fairly 
widely but locally established in North America by the 1920s. When Kiger (1971) 
evaluated the status of this species in the United States he noted that while the 
distribution of the species showed only aslight change in the previous 20 years, 
A. hispidus was becoming more firmly established within that range. Subse- 
quent reports (Newbold 1979; Cusick 1986; Brown & Schultz 1991; USDA, NRCS, 
2001) reveal that A. hispidus is spreading further west and north in the United 
States and merits concern as an invasive weed. 

In tropical America, Arthraxon hispidus first appeared in the Caribbean. 
The earliest collection we can locate was made in Guadeloupe in 1892 (Duss 
2712, US). The species then was found in Jamaica in 1903 (Underwood 1195, NY). 
Acentury later it does not seem to have spread to any other island in the Carib- 
bean (Nash 1912: Hitchcock 1936; Adams 1972; Fournet 1978; Gould 1979; all 
records cited as either the synonym A. quartinianus (A. Rich.) Nash or A, 
hispidus var. quartinianus (A. Rich.) Roberty). In Central America the first 
record of A. hispidus is a collection (Weatherwax 171, US) made in Guatemala 
in 1932. In 1945 the species was first collected in Mexico, in adjacent Chiapas 
state (Hernandez Xolocotzi X-412, US). Arthraxon hispidus was subsequently 
first recorded in El Salvador by 1960 (Weberling & Lagos cited in Berendsohn 
& Araniva de Gonzalez, 1991). In the 1960s it also first appeared in Honduras 
(Molina R. 18516, US), Costa Rica (Pohl & Davidse 11257, US), and Nicaragua 
(Molina R. 22890, NY). By 1980, the range of A. hispidus had expanded north- 
ward in Mexico to include at least one locality in the state of Michoacan (Car- 
vajal Hernandez, 1980; cited as A. quartinianus) and more recently one in the 
state of Colima (Sanders et al. 10373, MO). 

Another species of Arthraxon, A. castratus (Griff.) V. Naray. ex Bor, was 
collected in Puerto Rico in 1981 (Hatch 1982). This species appears to be less 
aggressive than A. hispidus, not otherwise being known outside its native range 
in Asia. Both van Welzen (1981) and Hatch (1982) provided keys that can be 
used to distinguish the two species of Arthraxon that occur as adventives in 
the New World. 

Voucher specimens. UNITED STATES. Oregon. Multnomah Co.: raised from young plans coe 
lected on ballast at Portland, 22 Jan 1901,W.N. Suksdorf 5316 (US). Pennsylvania. Philad 

Japanese Garden, at Centennial Grounds, Phil’, 1877, | Burk s.n. (US); On ship's ballast, Saami 
1878, FL. Scribner s.n. (US). South Carolina. Pickens Co.: Clemson College, introduced with seed 
rom Japan, Oct 1903, C.L. Newman s.n. (US). Virginia. Arlington Co.: cult. Arlington farm, Aug 1906, 
|. Tidestrom s.n. (US). MEXICO. Chiapas: from Finca Liquidambar to El Triunfo, ca. 1100-1850 m, 9 

1945, F. Hernandez Xolocotzi X-412 (US). Colima: Comala, Rancho El Jabali, 22 km (airline) NNW 

| eas (19°26'54"N, 103°4142"W), 1300 m, 7 Jan 1991, A.C. Sanders et al. 10373 (MO). GUATE- 
A. Quezaltenango: along rr.in mountains near Santa Maria, 25 Mar 1932, P Weatherwax 171 
ne HONDURAS. Morazan: between El Edén and El Rancho, mountain Canta Gallo, 1600 m, 1 No 
1966, A. Molina R. 18516 (NY, US). COSTA RICA. Alajuela: 4.5 km S of Zarcero, 1670 m, 15 Oct (eee 
R.W. Pohl & G. Davidse 11257 (US). NICARAGUA. Matagalpa: vic. El Porvenir,4 km E of Santa Maria de 


— 


DORR AND NINO, ARTHRAXON NEW TO SOUTH AMERICA 1193 


Ostuma, 30 Oct 1968,A. Molina R.22890 (NY). JAMAICA. Cinchona,8 Feb 1903, L.M. Underwood 1195 
(NY). GUADELOUPE. Environs de Bagatelle, 24 Feb 1892, Pére A. Duss 2712 (NY, US). VENEZUELA. 
Portuguesa: Mpio. Guanare. Caserio“La Montana’a 4 km NW de Cordoba (approx. 10°24'N,69°52'W), 

1000 m, 11 Dec 1986, G. Aymard 5109 (PORT). Carretera Chabasquén a Cordoba (9°25'N, 69°55'W), 
1200-1500 m, 19 Sep 1990, G.Aymard et al.8845 (MO,PORT). Trujillo: Mpio. Bocond. Parque Nacional 
Guaramacal, bosque a los arededores de la Laguna de Aguas Negras, cerca de Batatal (9°18,382N, 
70°10,515W), 1900 m, 23 Sep 2000, M. Nino et al. 1351 (PORT, US). 


~~ 


ACKNOWLEDGMENTS 
We are grateful t to the Ministerio del Ambiente and INPARQUES (Venezuela) 
for issuing g permits for our Flora de Guaramacal project. Gerrit Davidse 


(MO), Michael Nee (NY), and Basil Stergios (PORT) located and verified some of 
the material cited. 


REFERENCES 


Apams, C.D. 1972. Flowering plants of Jamaica. University of the West Indies, Mona, 
Jamaica. 

BeRENDSOHN, W.G. and A.E. Araniva DE GONZALEZ. 1991. Listado bdsico de la flora Salvadorensis. 
Monocotyledonae: Iridaceae, Commelinaceae, Gramineae, Cyperaceae. Cuscatlania 
1(6):1-29. 

Brown, L.E. and J. ScHuutz 1991. Arthraxon hispidus (Poaceae), new to Texas. Phytologia 
71:379-381. 

CARVAJAL HERNANDEZ, S. 1980. Notas sobre la flora fanerogamica de Nuevo Galicia, |.Phytologia 
46:145-153. 

Cusick, A.W. 1986. Distributional and taxonomic notes on the vascular flora of West 
Virginia. Castanea 51:56-65 

Fournet, J. 1978. Flore illustrée des phanérogames de Guadeloupe et de Martinique. Institut 
National de la Recherche Agronomique, Paris. 

GouLp, F.W. 1979. Poaceae. In: R.A. Howard. Flora of the Lesser Antilles, Vol. 3. 
Monocotyledonae. Arnold Arboretum of Harvard University, Jamaica Plain. Pp.25-220. 

Hatcn, S.L. 1982. Arthraxon castratus (Poaceae: Andropogoneae) new to the New World. 
Sida 9:266-267. 

HitcHcock, A.S. 1936. Manual of the grasses of the West Indies. U.S.D.A. Misc. Publ. 213: 

9 


Kicer, RW. 1971. Arthraxon hispidus (Gramineae) in the United States: taxonomic and 
floristic status. Rhodora 73:39-46. 

Nasu, G.V. 1912. Poaceae (pars). North American Flora 17(2):99-196. 

Newsotp, A. 1979. Additions to the check-list of the flora of Montgomery County, II. Bartonia 
46:49-50. 

USDA, NRCS. 2001. The PLANTS Database, Version 3.1 (http://plants.usda.gov). National 
Plant Data Center, Baton Rouge, Louisiana. [Accessed 28 September 2001]. 

Wetzen, PC. van. 1981. A taxonomic revision of the genus Arthraxon Beauv. (Gramineae). 
Blumea 27:255—300. 


1194 BRIT.ORG/SIDA 19(4) 


Book REVIEW 


] ,AN listi Approach 


Coral G. Guest. 2001. Painting Flowers in \ 
(ISBN 0-88192-509-8, pbk.), Timber Press, 133 SW Second Ave, oule4 0, 
Portland, OR 97204-3527, U.S.A. Published in association with The Royal 
Botanic Gardens, Kew. (Orders: www.timberpress.com, 800-327-5680, 503- 
227-2878, 503-227-3070 fax). $19.95, pbk., 128 pp., 78 color illustrations, 
43 color figures, 6 b/w figures, 1] line drawings. 


t ] like 


You've heard the expression, “Gee, I wish I could paint like that!” or “Where did you | 
that?” To many, art is often considered a gift, but it is a gift one must work for. This is why many 
latent artists never make that first step. They may feel the task too pone For those ae have 
wanted to try painting and in particular, botanical painting, t al nspira- 
tional ook i is the place to start. In Painting Flowers in Watercolour, a Guest has ce and 
organized her techniques so that anyone, whatever their level of expertise, can benefit 
Coral G. Guest is a full-time botanical artist who conducts courses 0 samncnisue botanical 
painting for the Royal pseu ee Kew, and at a number of other venues. Her work has been 
} : ; 1] jo} } oy J ] ; aay 1A Vf } R ] 
8 y 


Horticultural Society. She frequently lectures and has written for various magaz 
Coral’s techniques in watercolor are impeccably classical and she has ae laid out 
instructions in a step- eas step Ne with co ee full color visuals and illustrations. Her naturalistic 
helps th creative way, helping one develop not only tech- 


ach h paint 


approa 
nique, but obser eae skills as well 
renee lowers in Watercolour is quite oe from the get-go with an 
by Dr. Shirley Sherwood, noted connoisseur and collector of some of the finest contemporary ovate 
cal art in private hands. _ Sherwood has collected oe commissioned quite a few of Coral’s works, 
in particular, Lilium longiflorum which is included as part of the instructional lesson of the book. 
yok is divided into two parts. The chapters of Part | are: Equipment, Materials, Plant Mate- 
rial, Work in Progress, The Workspace, The Properties of Watercolour, Watercolour Wash chniques, 
Dry Brush Techniques, The Colour Palette, Mixing Greens, The Neutral Tint Mixture, Colour Assessment, 
Drawing, Composition, The Thumbnail Sketch, Light and the Tonal Sketch, The Colour ai Method. 
Part cigs V) oo ee ieee to ae au the se Color and eur of plant 
f work- 


ble Fo d 


F 
ing with Saneparent watercolor, which stilises che whiteness of he paper to achieve eas 
pale tints, rather than using white watercolor or opaque white watercolor (gouache). Part II is de- 
voted to: 1) observation of cime flowers in a systematic procedure, ee ane the eee and 
ee described in Part I. 2) specific issues, such as highlight on leaves and various surface tex- 
tures, are analyend as 5 they occur. Plants featured are: Lilium longiflorum, Anemone coronaria, Del- 
phinium, F iviaimperialis, Canna, Camellia japonica, Tulipa, Strelitzia reginae. The last part of 

the book is ee to: ‘Bohanitedt Terms, ns Bibliography Suppliers directory and Index 
om ents humble artist’s Spo of view, I find it one of the best instructional ee on flower 
e afew criticisms, however. There are 


cs 


printed and | well designed. Idoha 

the odd‘ typos’ here and there—those kind of pesky on ae errors that the ieee check 

doesn’t catch. That is minor stuff of course. More problematic is in the inconsistency of including 

e numbers vaca the text an meletencine a visual. eae it is a bit tedious trying to 
ced i 


find referen 1 ng a page number as reference 
to ALL the Scones visuals in the text would have made a quicker reference for the reader, espe- 


cially in some of the more — instructions.—Linny Heagy, Linny/Designer, Illustrator, Arling- 


VNETOIWED 


ton, TX, a00 @airmail.ne 


SIDA 19(4): 1194. 2001 


ANAGALLIS ARVENSIS SUBSP. FOEMINA (PRIMULACEAE) 


NEW TO LOUISIANA 
Charles M. Allen, Sara Thames, Selena Dawn Newman 
hillip Paul 
Department of Biology 3737 Government Street 
University of Louisiana at Monroe Alexandria, LA 71302, U.S.A: 


Monroe, LA 71209, U.S.A. 


Ona recent outing with the Louisiana Native Plant Society, a blue flowered “scar- 
let pimpernel” (Anagallis arvensis L.) was observed on Fort Polk in Vernon Par- 
ish. A search of the literature revealed that scarlet pimpernel can be blue flow- 
ered (Correll & Johnston 1970; Diggs et al. 1999; Fernald 1950; Radford et al. 
1968). Some of the blue flowered plants are only a color variant while others are 
separated as subsp. foemina (P. Mill.) Schinz & Thellung (Blamey & Grey-Wil- 
son 1989). The leaves of our plants were lanceolate and narrower than the com- 
mon and scarlet flowered var. arvensis. The blue petals were narrower than those 
of var. arvensis and were toothed but glabrous along the margins. Thus, our col- 
lection corresponds to subsp. foemina, as identified in Blamey and Grey-Wil- 
son (1989). The common name is usually poorman’s weathervane or weather- 
glass. The name is suggested by the closing of the flowers with cloudy conditions 
associated with inclement weather (Diggs et al. 1999; Fernald 1950). Synonymy 
for this subspecies includes A. foemina P. Mill (Blamey & Grey-Wilson 1989) 
and A. arvensis var. caerulea (Schreb.) Gren. & Godr. (Fernald 1950). The plants 
are annual/biennial and introduced from Europe. Prior to this report, the United 
States distribution included California, Colorado, lowa, Kentucky, Maine, Ohio, 
Pennsylvania, and Vermont (USDA, NRCS 2001). 

Voucher specimen: LOUISIANA. Vernon Parish: roadside shoulder of LA 467, western border of 
Fort Polk,ca 4 mi ESE of Leesville Section 34 T2N R8W, 31 Mar 2001,Allen, Thames, Paul,and Newman 
18437 (NLU). 


REFERENCES 


Biamey, M. and C. Grey-Witson. 1989. The illustrated flora of Britain and Northern Europe. 
Hodder & Stoughton, London 

Corrett, D.S. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner. 

Dices, G.M., B.L. Lirscome, and R.J. O’Kennon. 1999. Shinners & Mahler's illustrated flora of 
north central Texas. BRIT and Austin College, Fort Worth, Texas. 

Fernato, M.L. 1950. Gray's manual of botany, 8th ed. Reprinted 1987. Dioscorides Press, 
Portland, OR. 


SIDA 19(4): 1195 - 1196. 2001 


1196 BRIT.ORG/SIDA 19(4) 


Raprorb, A.E., H.E. AHLes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
Univ., North Carolina Press, Chapel Hill. 

USDA, NRCS. 2001. The PLANTS Database, Version 3.1 (http://plants.usda.gov). Accessed 
September 2001. National Plant Data Center, Baton Rouge, LA 70874-4490 USA. 


ALTERNANTHERA SESSILIS (AMARANTHACEAE) 
NEW TO ARKANSAS 


Christopher S. Reid and David X Williams 


University of Arkansas Herbarium 


Biomass Research Center, Rm. 141 
Fayetteville, AR 72701, U.S.A. 


Fieldwork for the Arkansas Vascular Flora Project has resulted in the discovery 
of Alternanthera sessilis (L.) R. Br ex DC. (chaff-weed, sessile joyweed) in Ar- 
kansas. This species was collected in Arkansas and Prairie counties, which lie 
in the Mississippi Alluvial Plain Natural Division of the state. Both occurrences 
were located in the White River watershed. Alternanthera sessilis is not reported 
for the state by Smith (1988, 1994). It is recorded in five parishes in northeast 
Louisiana, including East Carroll Parish, which borders Chicot County, Arkan- 
sas (Thomas & Allen 1996). 

Alternanthera sessilis has a pantropical distribution (Buckingham 1996) 
and appears on the United States Federal Noxious Weed List (Coile 1996). Fur- 
ther fieldwork is necessary to document the range of this species in Arkansas 
and to determine its status in the state. 

Voucher Specimens: ARKANSAS. Arkansas Co.: Abundant on either side of a small backwater 
inlet Johnson Bay) on the White River, NE1/4 $13 12S R3W, growing on mud, associated with 
7 ee aa nnaaee ae psa: oe Stance Sh 


hbristylic¢ 

J a f r r F : ao r. beau Enda Ca 
vahlii, 8 Aug 2000, Reid & Williams 1358 (UARK). Prairie Co.: Rare on mud around boat ramp at NE 
access to Peckerwood Lake, NE1/4 S27 TIN R5W,ca 2 mi SSE of Tollville,8 Oct 2000, Reid 1965 (UARK). 


ACKNOWLEDGMENTS 


We wish to thank Kenneth R. Robertson (ILLS) for verifying the identification 
of duplicate specimens. We would also like to thank the curators of the follow- 
ing herbaria for checking their collections for Arkansas specimens of A. sessilis: 
BRIT, MO, NLU, STAR, and UAM. 


REFERENCES 


BUCKINGHAM, G.R. 1996. Biological control of alligatorweed, Alternanthera philoxeroides, the 
world's first aquatic weed success story. Castanea 61:232—243. 

Come, N.R. 1996. Invasion of the South: The ecological impact and control of exotic weeds 
in the southeastern United States (Symposium Preface). Castanea 61:209—213. 

Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, second 
edition. Published by the author. 

Smith, E.B. 1994. Keys to the flora of Arkansas. The University of Arkansas Press, Fayetteville. 

THomas, R.D. and C.M. ALLEN. 1996. Atlas of the vascular flora of Louisiana. Vol. I]. Louisiana 
Department of Wildlife & Fisheries Natural Heritage Program, Baton Rouge. 


SIDA 19(4): 1197. 2001 


1198 BRIT.ORG/SIDA 19(4) 


Book NOTICES 


FREDERICK J. SIMOONS. 1998. Plants of Life, Plants of Death. (ISBN 0-299-15904-3, 
pbk.). University of Wisconsin Press, 2537 Daniels Street, Madison, W1 53718, 
US.A. $34.95, pbk., 568 pp., | map, 43 b/w photos, 6" x 9" 


Synthesizing evidence from the fields of geography, botany, religion, medicine, folklore, and anthro- 
pology, author Frederick J. Simoons has produced a groundbreaking volume that covers plant use 
shaped by two primary human concerns: life and death. Whether he is debunking the theories of 
Pythagoras’s ban of fava beans or bringing to light the Hindu use of the holy basil plant tulsi, the 
hor has compiled an erudite volume that is both exceptionally written and well researched. 
Geographically this tome covers the se from Europe through the Near East to India and China. 
The author covers subjects such as ritual purity, fertility, good health, ae sickness, ill fate, and 
death in his quest to decipher the historical and etl raphic dat lants of life and death. This 
edition is profusely illustrated and comes with a biblioataphy and general index. | would recom- 
mend this book to those of us interested in Old World ethnobotany and the perception and use of 
plants in religious and medical contexts—Kevin D. Janni, Botanical Research Institute of Texas, Fort 
Worth, TX 76102-4060, U.S.A, kjanni@britorg. 


a 


N.M. Nayar and T.A. More. 1998. Cucurbits. (ISBN 1-57808-003-7, hbk.). Science 
Publishers, Inc., RO. Box 699, May Street, Enfield, NH 03748, 603-632-7377, 
603-632-5611 fax). $75.00, 340 pp. 


The Cucurbitaceae or gourd family is what this book is all about. Cucurbits are “of considerable eco- 
nomic importance as a source of vegetables, fruits, edible seeds rich in oil and proteins, edible and 
industrial seed oils, domestic utensils and drugs.” This is a comprehensive book with 17 chapters 
ne cover every nane from Eta ton morpneeey as yology, physiology and biochemistry, and 
plant protection, and to seed production 
of oe ouch, Botanical Research Institute of e xds, Fort hia TX 76102-4060, 
v@) it. g. 


U.S.A 


Book ANNOUNCEMENT 


Ropert D. Dorn. 2001. Vascular Plants of Wyoming. Third Edition. Illustrations 
by Jane L. Dorn. Mountain West Publishing, Cheyenne, Wyoming. Rocky 
Mountain Herbarium, Department of Botany, University of Wyoming, 
Laramie, WY 82071-3165, US.A., http://wwwrmbh.uwyo edu. (Orders: Copies 
sell for $20.00, shipping included in USA; Wyoming orders add sales tax 
for ahs cack [4% - $.80, 5% - $1.00, 6% - $1.20). 412 pp. 


Pl } } dD ] 


erbarium. Questions, contact Ronald L. Hartman, 
Curator, Rocky Mouncin Herbarium: ere ae edu; 307-766-2236; fax 307-766-2851. 


SIDA 19(4): 1198. 2001 


VASCULAR PLANTS NEW TO KENTUCKY 
J,Richard Abbott Ralph L. Thompson & Rudy A. Gelis 


Marie Selby Botanical Gardens Herbarium, Biology Department 
811 South Palm Ave. Berea College 

Sarasota, FL, 34236-7726 U.S.A. College PO.2121 
galearis@yahoo.com Berea, KY, 40404 U.S.A. 


ralph_thompson@berea.edu 


ABSTRACT 
Fourteen species of vascular 1 new to the state of Kentucky, U.S.A. Five of these are 
European weeds: Anthiosanthain pes Aphanes microcarpa, Erysimum hieraciifolium, Lathyrus 
tuberosus, and wierd tetrasperma. dns ome nine are native epee snows from acacent states ome 
iC C 


Elymus wiegandii, an x pence spent in aie Lupinus perennis, Polygonum 


cilinode, and Silene nivea 


RESUMEN 


Se presentan 14 especies de plantas l f d biert ara el estado 

de Kentucky, EE.UU. Cinco de éstas son malezas de Europa: Anthoxanthum aristatum, Aphane 

us s wunerOsus, y Vicia tetrasperma. Las otras nueve son 
y, por eso, no fueron totalmente inesperadas: 


microcarpa, Erysimum Meractotnitte L t} 


q 


= 


Acdiena deamii, Carex austrocaroliniana, Centunculus minimus, Elymus wiegandii, Equisetum x 
ferrissii, Leucothoe fontanesiana, Lupinus perennis, Polygonum cilinode, y Silene nivea. 


DISCUSSION 


A recent dissertation (Medley 1993) and a recent book (Browne & Athey 1992) 
provided lists of the vascular flora of Kentucky and are the comparative basis 
for our records. We report the following species to make knowledge of their pres- 
ence available not only to other in-state workers but also to the botanical com- 
munity at large. Gleason and Cronquist (1991) was used for distributional in- 
formation outside Kentucky and is the main source of the nomenclature used 
here. Taxa not present in Gleason and Cronquist were found in Radford et al. 
(1968). All specimens are deposited at the Berea College Herbarium [BEREA]. 
Acalypha deamii (Weath.) Ahles (Euphorbiaceae)—Reported in mesic sites from 
southern Indiana and Ohio to Tennessee and Arkansas. Levin (1999) did not 
report this species from Kentucky. 


on Co.: Fort Boonesborough State Park, SW of junction of KY 627 and the Kentucky River 
upper ae rare; associates: sats ace americana, Commelina virginica, Lobelia Siphilitice: 
Stachys spp., Te um cand ; 26 Sep 1992, Abbott 4464. Verified by John Thieret [KNKI. 


Anthoxanthum aristatum Boiss. (Poaceae]|—Native to Europe; introduced 
throughout the northeastern United States. 


SIDA 19(4): 1199 — 1202. 2001 


1200 BRIT.ORG/SIDA 19(4) 


Trigg Co.: Land Between The Lakes, Elk-Bison Prairie, entranceway and the Trace Road, locally fre- 
quent, associates: a poatonaeon BLS Bromus japonicus, Carex hirsutella, Chamaecrista 
H i ta, L. striata, Panicum lanuginosum, Smilax glauca, 
Solidago eanatenky Iripsacum dactyloides Vuleracetoit ord, 27 Jun 1998, Thompson 98-282. Verified 
by John Thieret [KNK]. 


fasciculata, 


Aphanes microcarpa (Boiss. @ Reuter) Rothm. (Rosaceae)—Native to southern 
Europe. In United States from Long Island to Alabama. 
Madison Co.: Fort Boonesbor ae State Park, sk of junction of uo al ane as Hanna River, 


mowed yard in campground, rare, only one | pulina, Plantago 


spp., Poa spp., Trifolium spp., are spp. 21 Mey 1994, Abbott 6898. 

Carex austrocaroliniana Bailey (Cyperaceae)—A southern Appalachian endemic 
(Radford et al. 1968). First found in Kentucky in Laurel County by Thompson 
(Campbell et al. 1994), and now known to be found in other southeastern counties. 


Laurel Co.: Rock Creek Research ss Area, ca 30 km S of London off KY 1193 and 2 km W on FS. 
131, mixed mesophytic forest ravine at Rock Creek and Rockcastle River, occasional; associate: Carex 
laxiflora; 21 Apr 1985, Thompson 85- ee Verified by Rob Naczi [KNK]. 


Centunculus minimus L. [= Anagallis minima (L.) Krause] (Primulaceae)—An 
irregularly cosmopolitan species, especially in warmer areas. Reported from 
Kentucky, without explanation, by Radford et al. (1968). Medley (1993) cited an 
1840, Short s.n., Muhlenberg Co. collection at Philadelphia [PH], and he also 
stated that the specimen should be verified. David Hewitt [PH] was kind enough 
to send usa digital scan of the specimen, which is indeed this species and was 
reportedly common around Greenville in Muhlenberg Co. in western Kentucky. 
Our collection represents the first known population in Kentucky in over 150 
years and its first documentation in east-central Kentucky. 


Madison Co.: A wet ditch on as E side a northbound I-75 at KY 21 (Exit 77); associates: Daucus 
carota, Rhexid virginica, S floril s, 17 Jun 1995, Thompson 95-374. Verified by John Thieret 
IKNK]. 


Elymus wiegandii Fern. (Poaceae)—In northeastern North America and scat- 
tered to the west. 


Lewis Co.: Manchester Island #1 (27 acres), in the Ohio I ie at mile a8, l. ‘“ mi FE of Tr tae ee 
forested wet es saccharinum- Sloe Platanus|,d 


field, near middle of island, rare; associates: Boehmeria cylindrica, Laportea canadensis, Leersia 
virginica, Ai carees frondosa, eae virginianum: 29 Jul 1995, Gelis MI-753. 


Equisetum x ferrissii Clute (Equisetaceae)—According to the distribution maps 
in Hauke (1993), this sterile hybrid is more widespread in North America than 
E. laevigatum A. Braun, the parent it most closely resembles. Medley (1993) cited 
literature reports for E. laevigatum in Kentucky, but he said no voucher speci- 
mens are known. 

Laurel Co.: Along Laurel River downstream of Laurel River Dam cee oh hitley C euaty line), 


f 
L ie) oO 


sandy banks scattered among rocks and boulders, i LUVeX 


ABBOTT ET AL., VASCULAR PLANTS NEW TO KENTUCKY 1201 


eae Phlox maculata, Physostegia virginiana, ee arborescens, Rudbeckia laciniata, 

elaginella apoda, Spiraea angen, Ee CHE ee qutvetteria caroliniensis, 1 Aug 1993, 
Abbott 5970. Whitley Co.: The above popu ation | I vera few hundred 
feet downstream from the powerplant es off the end of County Road 1193; handheld GPS coordi- 
nates 36'57' 41"N 84° 16' 31" W [NAD = projection System vel ce les tt eae This time strobili 
were present, which yielded sp g the identity. 


Erysimum hieraciifolium L. ie to Europe; naturalized in 
Canada, Michigan, and Wisconsin. 


urel Co.: NNW of Hazelpatch, scattered along railroad tracks N of junction with C.R. 760 and S of 


Shetland Trace, E of the Little Rockcastle River, in gravel along railroad adjacent to mesic woods, 23 
Jun 1993, Abbott saline Penden, sist E of - S.250n R. 106 (toward Levi Jackson Wilderness Road 
State Park), ti Camelina microcarpa, 


Plantago aristata, Poa een Triodanis nerfoliata: Tragopogon sp. 19 Jun 1994, Abbott 6979. Veri- 
fied by John Thieret [KNK]. 


Lathyrus tuberosus L. (Fabaceae)—Native to Europe and western Asia. Known 
from a few areas in the United States, mostly in the north. Isely (1990) reported 
this species from Tennessee. 


Madison Co.: Berea College Campus; S of the music building, in unmowed field adjacent to the NW 
end of the athletic fields, locally abundant sprawling vine, no record of it having been planted; 2 Jun 
1992, Abbott 2374. 


Leucothoe fontanesiana (Steud.) Sleumer (Ericaceae)—Native to the southeast- 
ern United States. 


Madison Co.: Lower Dogfoot Springs, Berea College Forest, 1.6 mi S of Bighill (at junction of KY 21 
and U.S. 421), and 0.2 mi S of Grant House entrance, in a NNW mesic mixed hardwood-hemlock 
ravine  transyersed vy ean aan Creek, infrequent, several shrubs; associates: Acer saccharum, 
is, Cornus florida, Lindera benzoin, Lirioc ape alas Tilia 

americana, sar tins radicans, Tsuga canadensis, 12 Jul 1995, Soe 95-407. This popula- 
tion was first found in 1982 by David D. Taylor [#1943, BEREA]. Later in the 1990s, - cee 
was buried and extirpated during road construction on U.S. 421, but some shrubs were transplanted 
to the nearby Upper Dogfoot Springs. Some skeptics had claimed that this population may have just 
persisting from cultivation by some 19th century settler, despite the lack of homesite remnants. 
ntly, however, a second po Sao was brought to our attention by Ben Begley, ames the 


Hee that this species is naturally in entee and has just been ear rlan Co.: Pine 
eee Settlement School, Pine Mountain, 3.1 mi SSE from Big Laurel on KY 221 at junction w 

Y 510, 2.2 mi Eof Letcher Co.,and 9. die ESE from a 42lina Fagueprandiolioa and ee ee ee 
a na yee alba, H tyli lia, 1 Rhodo- 


dendron maximum, Thelypteris niovebonacensis and Tiarella sea 5 May 2001, ee 01-51. 
Lupinus perennis L. (Fabaceae )—Widely native in the eastern United States. Isely 
(1990) reported this species to the north, east, and south of Kentucky. 

Rockcastle Co.: Horselick Surface-mined Area, 1.8 mi S of KY 1955 off Daugherty Road, open bench, 
rare; associates: Cardamine hirsuta, Trifolium pratense, T. repens, Veronica arvensis, 14 Jun 1987, 
Thompson 87-735. 

Polygonum cilinode Michx. (Polygonaceae)—Native to northeastern North 
America and south, in the mountains, to North Carolina. 


1202 BRIT.ORG/SIDA 19(4) 


Lewis Co.: Manchester Island #2 (115 acres), in the Ohio River at mile 396, 1.3 mi E of Trinity Station, 
old field [Rubus-Sambucus- Vitis], E rim of island head, trailing down from upper terrace, rare; associ- 


ates: Erigeron philadelphicus, Ipomoea pandurata, Rudbeckia ae Urtica ee Verbesina 
alternifolia, Vernonia gigantea; 21 Jul 1995, Gelis M2-522. 

Silene nivea (Nutt.) Otth. (Caryophyllaceae)—Widespread in the northeastern 
United States. 


Lewis Co.: Manchester Island #1 (27 acres), in the Ohio River at soe = 13 mi E of baeuacse - ane, 
late old pepsi’ ies us-Acer age Rosa- wet hb Idatisland | 


st bottomlan Phalari li 1, Stellaria aquatica, 
te dioica, eee alternifolia, 54 tun 1995, Gag M1-389. Verified by John Thieret [KNK]. 
Vicia tetrasperma (L.) Moench. (Fabaceae)—Native of Eurasia and introduced 
at scattered stations in the eastern United States to Texas. Isely (1990) reported 
this species as occurring primarily on the coastal plain, but he also said that it 
was obey present in every southeastern state. 


adison Co.: Berea College Forest, Red Lick Reservoir, exposed south side of dam, lower slope, ne 
cattail marsh, rare, one small population of 20-25 individuals; associates: Melilotus alba, ras 
repens; 19 Jun 1996, Gelis 393. 


REFERENCES 


Browne, E.T., Jk. and R. AtHey. 1992. Vascular plants of Kentucky: an annotated checklist. 
University Press of Kentucky, Lexington. 

Campseit, J.J.N., J.R. Aggort, R.R. Ciceretto, J.D. Kiser, JH. MacGrecor, and J.G. Pats. 1994. Coop- 
erative inventory of endangered, threatened, sensitive and rare species, Daniel Boone 
National Forest: London Ranger District. Kentucky State Nature Preserves Commis- 
sion, Frankfort. 

Gteason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada. New York Botanical Garden, Bronx. 

Hauke, R.L. 1993. Equisetaceae. In: Flora of North America Editorial Committee, eds. Flora 
North America 2:/6-84. Oxford University Press, New York. 

Isety, D. 1990. Leguminosae (Fabaceae). Vascular flora of the southeastern United States 
3(2):1-258. University of North Carolina Press, Chapel Hill. 

Levin, G.A. 1999. Notes on Acalypha (Euphorbiaceae) in North America. Rhodora 101:217- 


Meotey, M.E. 1993. An annotated catalog of the known or reported vascular flora of Ken- 
tucky. Ph.D. dissertation, University of Louisville, Louisville, Kentucky 

Raprorb, A.E., H.E. Ades, and C.R. Beit. 1968. Manual of the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill. 


BOOK REVIEWS 1203 


Book REVIEWS 


Dan Graurand Wen-HsiunG Li. 2000. Fundamentals of Molecular Evolution, Second 
edition. (ISBN 0-87893-266-6, pbk.). Sinauer Associates, Inc., PO Box 407, 
23 Plumtree Road, Sunderland, MA 01375 U-S.A. (Orders: Internet: 
publish@sinauer.com, wwwsinauercom, 413-549-1118 fax). $48.95, pbk., 
482 pp., 7" x 9" 

Having been trained in biosystematics and comparative secondary chemistry, | feel a bit like the 

driver of an old 70s model car who has just been left behind in the dust of a new sporty model, the 

molecular systematics model, that is. Hence, | approached reviewing this book from the standpoint 

of whether it could bring me up to date in my understanding of molecular biology and how the data 

are applied to systematics. 


After reading this book, I do feel more comfortable discussing introns, reverse transcription, 
Sete and soon. In hat sense, the book succeeded in passing the test of teaching me. However, 


1 


readings. This was particularly true for concepts that, 
meeauced early in the book, were the basis for models or for more complex concepts later in the 
book (e.g, replication of leading vs. lagging strands; Holliday structures and heteroduplexes, calcu- 
lation of mutation ange: supSrtunion rates a 


nly tl I ducti st that the book assumes the reader 
hasa | 1 ling of molecular ap DIE ys and Bene Presumably. cheaa main Ere of this book 
7 f ‘} -| + 
primary li | Pp ide t] ith tk ls f learning h techni However, it is 
not a “cookbook” of techniques i lecul 1 populati ti Unfortunately, the 


1 


_ 


book also tries to be a primer many basic topics, the primer 
part becomes shallow and inaccurate. eo: example, the first ee ee with an explanation of 

.In the second enon the four bases are equated with nucleotides: 

chain is a linear polynucleotide consisting of four kinds of nucleotides. There are two 

purines, oe )and guanine (G), and two pyrimidines, thymine (T) and cytosine (C). 

Before the term “base” is explained, the concept of canonical base pairs is presented. 

This brings up another weakness. There is no glossary. Certain words are boldface in both the 
text and index. These d idered to be “defined” where first presented in the text. Often, 
this definition must be gleaned from context a 


Writers of conceptually oriented bool d to | ful to p pti hat underlie 
the various concepts; for the most part, th hors | led. However, they were not always 


aware of their assumptions, especially of their bias of the supremacy of molecular methods over 
more “Haginonal ones, as is evident on peer a 


1 } }] ] ‘ } Te 


“the introduction of molecular 
footing, and has turned iti ich rel d, counted, or com- 


puted from empirical data, and theories are tested against objective reality. 
Such seca reveal that the authors really do not understand that molecular data faces 
— oA ee same limitations Sion more severe limitations in the case of homoplasious base dif- 


feren rganismal data n both data types are subjected to algorithms of population 
genetics and cladistics. Lies: authors seem - me to As nae the assumption that the 
genome contains all necessary i | pathways differentiat- 


ing taxa, whereas recent research is also pointing to the: pioteinone and maternal inheritance as 
major factors, as well. 


s explained in the i lucti he book f 


g in two major areas: 1) molecu- 


SIDA 19(4): 1203. 2001 


1204 BRIT.ORG/SIDA 19(4) 


c 


lar evolution as oe a a ea and character ) molecular evolution 
“ hv No 


pothese of differences among organisms. The topics related to the two ma- 
areas are eae in ee nee long chapters 
hapter One, where basic terms and concepts are introduced, suffers many of the problems 
already mentioned. 
ntroduces concepts of population genetics espenally: rates ? eee ae 


a 
under differing assumptions and conditions. The connection t 


otide substitutional differences among species and high her oop Ds is poorly ¢ developed. The ongoing 


controversy between the hypotheses of the neo-Dar ts and neutral mutation advo- 
cates is well covered. 


| aS jl ] : # ] 


otide sequences among individual organisms and taxa. Methods for aligning eanences from ee 
inspection to computer algorithms, are covered. 


Based on these assumptions, Chapter Four shows ho i dto draw 
conclusions about the mechanisms and rates of Maen subst itution. Becanise some sites and re- 
gions show greater differences than do other regions, an important part of the chapter examines 
mechanisms that might explain these nonrandom biases 

For the systematist, Chapter Five contains the most meat—the use of nucleotide as data to 


construct hypothetical phylogenies. Not only does it introduce the vocabulary of phylogenetics, but 
also covers ee problems, as well. aie! of f distance measures and the aaa for 
UPGMA group method f ae measures, transformed 
distance methods (true parsimony), neighbors-relati d neigh! and minimum 


Sparta ing Siecle eis et in si hecho I a the ‘book could’: rave been improved if the 


I p work thro pA by nang ai these Bynetent 
] . ] eae 


methods. Because the math of 


and tree d maximum likeliHieod is ey complicate, only the seschipion oF phese 
methods is sven: Alcoinelaaed are rooting methods, est 


BeScssmnemisieis, bootstrap). ae empire On Hope eric were re provided (human-ape; ceta- 
igins; tree-of -lif gths of the various analytical methods. 


I ealiia beyond point mutations, Chapter Six examines duplication and modification of gene 


eee especially pseudogenes, alae pathways for cnn ee new funertOns ane une lack 
li Th 


of independent evolution of d 


concept of evolution by ‘ amine eeries* “Le. insertion or elon of transposable elements, includ- 
ing both transposition and retroposition, is covered in Chapter Seven. Here the authors give a rather 
complete account o | retroviruses, retroposons, and re eal a I ieee the de- 
tailed explanations of LINE 1 SINE be help 


ful. Lateral gene transfer among unrelated species isan micgeiaiee ecncent of special interest: can it 
account for discontinuity in the taxonomic distribution of isolated genetic sequence 

Chapter Eight relates attempts to account for veaehone current! differences at the mo- 
lecular level. That is, what h to th what is heterochromatin, 


and why are there vast difieeaces in sizes Sor genomes of die organism? Also two appendices 
provide acne! bi pare oun iO Sie unfamiliar with 1) the geologic column, geologic history, 


and current sp thematical probabilities. 


r the goals the auithots have set ae themselves, | would give the book a grade of “B.” If other 
books that do a better job are not available in this niche, then, for the taxonomist seeding a better 
understanding of molecular systematics, it is worth the investment.—Roger W. Sanders, Associate 
Collections plas aes scl iaacik Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102- 
4060, U.S.A, rsanders@br 


‘: 


SIDA 19(4): 1204. 2001 


BOOK REVIEWS 1205 


Jou J. Wiens (Editor). 2000. Phylogenetic Analysis of Morphological Data. (ISBN 
1-56098-841-X, hbk. 1-56098-816-9, pbk.). Smithsonian Institution Press, 
PO Box 960, Herndon, VA 20172-0960 U.S.A. (Orders: MLitts@sipress.si.edu, 
1-800-782-4612; 202-287-3738 x343). $49.95 hbk,, 26.95, pbk., 220 pp., tables 
and numerous figures, 6" x 9", 
What ever happened to morphology-based systematics? A perusal of titles of such traditional jour- 
nals as Systematic Botany reveals that studies employing morphological data have been replaced 
largely by those focusing on molecular. If that has you feeling depressed, a quick read-through of this 
book will have you pumped up about the future of morphological studies 
symposium, “Morphological Data in Phylogenetic Analysis: Recent Progress and Unresolved 
Problems,” at the 1996 annual meeting of the Society of Systematic Biologists gave birth to this col- 
lection of papers. The symposium’s title clearly re elects the theme of the ule —no nape WHE 


] 


Insteac 


over the demise of morpholo 
bution (Chapter 1, “Molecules V Versus Meee) in Systematics”) as a means of oe morpho- 
logical analyses in the perspective of the current state of systematics and providing a context for the 
rest of the papers. The senior author, David M. Hillis with John Wiens as junior author), is a promi- 
nent advocate of using both morphological and molecular data in systematics. Hence, a reasoned 
argument is made that the best studies take advantage of the strengths and avoid the weaknesses of 
both morphological and molecular data. Moreover, ae training in systematics should generate 
researchers who have a command of both m 

In Chapter 2, “Character Selection and a Mees of Morphological Sosa Steven 
Poe and John Wiens begin the study of morphological data sets in earnest. In t tryin g to answer the 
question of how systematists decide on the set of characters they actually includ ladistic stud- 
ies, the authors subveyed 23 journals between 1987 and 1997 for research papers that explicitly state 
criteria for character inclusion. Only about 20 Sales of ae papers gave any cured which can be 


categorized as: 1) excessive variation in terminal ta 
titative, 3) missing data, 4) too much homoplasy in oe and 5) polarity unknown. Theyre point 
out that rejection of characters generally was based on assumptions taihier than on L emmipitical evi- 
dence. After analyzing each of the criteria, they g after one has 
objective evidence to support rejection 
Chapter 5 (‘Coding eee Variation within Species and Higher Taxa for Phylogenetic 
Analysis”) actually follows in logical sequence after Chapter 2. However, because its sole author is 
John Wiens, its placement probably reflects modesty. The first part of the paper focuses on variation 
within species. Using simulations of an artificial (ie. “known”) phylogeny and nee analyses 
of a real data set, he compared the effect of eight different fem methods under parsimony algo- 
rithms to that of continuous data under maximum likelihood, as well as ee all combinations of 
three diffe cl g methods (UPGMA, neighbor-joining, Fitch-Margoliash) with two distance 
measures or s, Cavalli- Sforza & Edwards's). The results were surprising. The most highly resolved, 
and potentially most accurate, cladograms resulted from parsimony of continuous characters coded 
as be oeees ae from Hg non-parsimony methods. Lowest resolution came from excluding all 
, the standard practice of phylogenetics) and from coding the character 


as as missing or equal soe two or more states. wena coding met hods to include data pe oy 


1 


seco ine to modal eudinen ee aie onc scored as a state falermmediate or alternative 
to fixed apomorphy) performed at ee levels. eas — concludes that analytical 
information is lost when polymorphic cl 1 or not allowed to contribute to tree 


construction. Wiens also Somme the various methods of coding eee states in higher taxa 


SIDA 19(4): 1205. 2001 


1206 BRIT.ORG/SIDA 19(4) 


with iatl ies. In those cases, | | Se Parmar ] 
= dee 3 
ener as han aoaing higher taxa ore a characters. 


orphological research in which advances 
are eccutring- morphometric, ontogeny, hybridization, seller) of fossils, and adaptation. 

nd the study of morphometrics (both intraspecific and interspecific variation) to be fasci- 
aie a was pleased by Zelditch, Swiderski, and Fink’s article (Chapter 3, “Discovering Phyloge- 
netic Characters in Morphometric Data). They doa superb job of explaining the field and summarizing 
its state as practiced today. If one is not familiar with morphometrics, this article would make a good 
introduction for entering the literature. 


aula Mabee (Chapter 4, “The pees of Satbpeny in a tee Mier phological Charac- 
ters) summariz s the state of affairs She 


reviews the recent history of the role of éatieenyy in cladistics, eriticeine agjeetilied uses (especially 


in pattern Aadisites) with recent empirical studies. She al commends ways of coding ontogeny 
as characters. However, ontogeny has not proved useful for a ree or ordering characters or es- 
tablishing homology. There must first be a better understanding of morphogenesis. 

Chapter 6 (Hybridization and Phylogenetics: Special Insights from Morphology), Lucinda 
McDade reviews the state of dealing with hybrids in this context. In particular she discusses the 
meaning of hybrid intermediacy, how it can be recognized, and the expected placement of hybrids 
in cladistic analyses. She makes a very good case that new computer algorithms are needed to per- 
form hybrid reticulations in addition to species’ cladogenesis. 


John Huelsenbeck and Bruce Rannala’s treatment of fossil evidence (Chapter 7, “Using Strati- 
graphic Information in Phylogenetics”) is enlightening but rather tedious, except for someone inter- 
ested in maximum-likelihood mathematics. They provide the historical setting for the new interest 
(since about 1985) of using fossil data for phylogenetics. Rather than focusing on using morphologi- 
cal characters of fossils, they are concerned most with using the stratigraphic occurrences to evalu- 
ate or choose among alternative cladograms. Apparently, maximum likelihood statistics have be- 
come an important factor in such studies, and they devote a major portion of their article to the topic. 

In Chapter 8 (Logical Problems Associated with Including and Excluding Characters During 
Tree Reconstruction and Their Implications for the Study of Morphological Character Evolution), 
Kevin de Queiroz is concerned about circular reasoning. That is, can interpretations about adapta- 
tions be related to characters that were included in the tree-bui Iding analysis? Despite the impor- 
tance of avoiding circular reasoning, | found de eee beatin saunas tedious. He spends con- 
siderable time building a philosophical t 
circularity of including the data versus excluding the daa, He seems to build a straw man of the 
practice of excluding data to replace it with the superior method of including data. In frustration 
before reaching his conclusion, I thought to ae —— not just ee eo and compare the effects 
on the tree?”—to later discover de Queiroz arriving a 
me down that path or did he get there himself by caer 


. Was he actually leading 


Allin all, the papers are = consistently well written and edited am ppears to be an important 
contnipouenst the systematic literature and : d f and challenges 
for logical systematics. | recommend it to anyone wanting to catch upon tl ing edge of 


sonpholesine or peas encouragement to continue using “out-dated” characters. ee ee it should 
be required reading of molecular phylogenetics who have not been trained to recognize the apex of a 
leaf or the skull of a mammal.—Roger W. Sanders, Associate Collections setae ae Research 
Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060. U.S.A iders@ 


SIDA 19(4): 1206. 2001 


BOOKS RECEIVED 1207 


Books RECEIVED 


c¢ # IRA 1 EIT [fowid 


JouN M. Gittett, Norman L. Tayitor, and MicHaeL CoL.ins (edited by Michael Col- 
lins). 2001. The World of bia (ISBN 0-8138-2986-0, hbk.). lowa State 
University Press, 2121 SouthS t, Ames, [A 50014-8300, wwwisupress.com, 
800-862-6657, 515-292-0155, 515-292-3348 fax). $144.95 hbk. (includes CD- 
ROM), $99.95 (CD-ROM only), 488 pp, illustrated, 6" x 9" 


IRWIN M. Bropo, SytviA DURAN SHARNOFF, and STEPHEN SHARNOFF. 2001. Lichens of 
North America. (ISBN 0-300-08249-5, hbk.). Yale University Press, 302 Temple 
Street, PO. Box 209040, New Haven CT 06520-9040, U.S.A. (Orders: 
www.yale.edu/yup, 203-432-0960, 203-432-0948 fax). $69.95, hbk., 795 pp., 
939 color photos, maps, 91/2" x 11". 

Jim Cootes. 2001. Orchids of the Philippines. (ISBN 0-88192-516-0, hbk.). Timber 
Press, 133 SW Second Ave., Suite 450, Portland, OR 97204-3527, US.A. (Orders: 
www.timberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). 
$49.95, hbk., 232 pp., 339 color ee 1 diagram, 1 map, 7 1/2" x 10". 


c5 VE (Medicinal/Useful Plants/Toxic 


Cuar_es R. Hart, TAM GARLAND, A. CATHERINE BARR, BRUCE CARPENTER, and JOHN C. 
ReEAGOR. 2000. Toxic Plants of Texas. Integrated Management Strategies to 
Prevent Livestock Losses. (no ISBN listed, TAES B-6105 12/00). Texas Ag- 
ricultural Extension Service, PO. Box 1209, Bryan, TX 77806, U.S.A. (Or- 
ders: http://texaserc.tamu.edu, 888-900-2577, 979-862-1566 fax). $20.00, 
spiral bound, 247 pp., color photos, 51/2" x 81/2". 


Grorce E. Burrows and RonaLp J. Tyre. 2001. Toxic Plants of North America. (ISBN 
0-8138-2266-1, hbk.). lowa State University Press, 2121 South State Street, 
Ames, [A 50014-8300, U.S.A. (Orders: www.isupress.com, 800-862-6657, 515- 
292-0155, 515-292-3348 fax). $174.95 hbk., 1350 pp, illustrated, 81/2" x 11". 


Hans ZocBELEIN (editor). 2001. Dictionary of Renewable Resources. Second, Revised 
and Enlarged Edition. (ISBN 3-527-30114-3, hbk.). Wiley-VCH Verlag GmbH, 
D-69469 Weinheim, Germany (Orders: John Wiley & Sons, Inc., One Wiley 
Drive, Somerset, NJ 08875, 732-302-2300 fax). Price not listed, 408 pp., 6 
3/4" x 91/2" 

JAMES LANG. 2001. Notes of a Potato Watcher. (ISBN: 1-58544-154-6, pbk.). Texas 
A&M University, 4354 TAMU, College Station, TX 77843-4354, 
www.tamuedu/upress, 979-458-3982, 979-847-8752 fax). $24.95, 388 pp, 
8 color, 9 b/w photos, 5 line drawings, 8 tables, bibliography, index, 6" x 9". 


SIDA 19(4): 1207. 2001 


1208 BRIT.ORG/SIDA 19(4) 


Botany/Molecular 


MOHAMMAD PessaRAkLI (editor). 2002. Handbook of Plant and Crop Physiology. 
Second Edition, Revised, and Expanded. (ISBN 0-8247-0546-7, hbk.). Marcel 
Dekker, Inc. Cimarron Road, PO. Box 5005, Monticello, NY 12701-5185, U.S.A. 
(Orders: www.dekker.com, 212-696-9000, 212-685-4540 fax). $225.00, 973 
pp., figures, 7" x 10". 

Ecology/Conservation/Landscape Ecology 


CLARK C. Gipson, MARGARET A. McKean, and ELinor Ostrom (editors). 2000. People 
and Forests. Communities, Institutions, and Governance. (ISBN 0-262-57137- 
4, pbk.). The MIT Press, 5 Cambridge Center, Cambridge, MA 02142-1493, 
US.A. (Orders: http://mitpress.mitedu, mitpress-orders@mit.edu, 800-356- 
0343). $21.00, pbk., 274 pp., 22 illustrations, 6" x 9" 


Mito L Harpsteap, THOMAS J. SAURER, and WILLIAM F. Bennett. 2001. Soil Science 
Simplified. Fourth Edition. (ISBN 0-8138-2942-9, hbk.). lowa State University 
Press, A Blackwell Science Company, 2121 South State Street, Ames, [A 50014- 
8300, U.S.A. (Orders: www.isupress.com, 800-862-6657, 515-292-0155, 515- 
292-3348 fax). $42.95 hbk., 268 pp, illustrated, 6" x 9" 


Horticulture/Gardening 


DIANA GRENFELL. 2001. The Gardener’s Guide to Growing Hostas. (ISBN 0-88192- 
518-7, pbk.). Timber Press, 133 SW Second Ave., Suite 450, Portland, OR 97204- 
3527, US.A. (Orders: www.timberpress.com, 800-327-5680, 503-227-2878, 
503-227-3070 fax). $19.95, pbk., 160 pp., 74 color photos, 5 line drawings, 
7x 91/2" 

GEOFF STEBBINGS. 2001. The Gardener’s Guide to Growing Irises. (ISBN 0-88192- 
388-5, pbk.). Timber Press, 133 SW Second Ave., Suite 450, Portland, OR 97204- 
3527, U.S.A. (Orders: www.timberpress.com, 800-327-5680, 503-227-2878, 
503-227-3070 fax). $19.95, pbk., 160 pp., 74 color photos, 3 line drawings, 
Peo. 

GRAHAM Rice and ELizABETH STRANGMAN. 2001. The Gardener’s Guide to Growing 
Hellebores. (ISBN 0-88192-517-9, pbk.). Timber Press, 133 SW Second Ave., 
Suite 450, Portland, OR 97204-3527, U.S.A.(Orders: www.timberpress.com, 
800-327-5680, 503-227-2878, 503-227-3070 fax). $19.95, pbk., 160 pp., 74 
color photos, 20 line drawings, 7" x 91/2" 

JANE McGary (editor). 2001. Bulbs of North America. (ISBN 0-88192-517-9, pbk.). 
Timber Press and North America Rock Garden Society (Orders: Tim ber Press, 
133 SW Second Ave., Suite 450, Portland, OR 97204-3527, U.S.A., 


SIDA 19(4): 1208. 2001 


BOOKS RECEIVED 1209 


www.timberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). 
$34.95, hbk., 308 pp., 101 color photos, 4 line drawings, 2 maps, 6" x 9". 


ALLAN M. ArmitaGe. 2001. Armitage’s Manual of Annuals, Biennals, and Half-Hardy 
Perennials. (ISBN 0-88192-505-5, hbk.). Timber Press, 133 SW Second Ave., 
Suite 450, Portland, OR 97204-3527, US.A. (Orders: www.timberpress.com, 
800-327-5680, 503-227-2878, 503-227-3070 fax). $39.95, hbk., 604 pp., 125 
color photos, 107 line drawings, | map, 75 tables, 6" x 9" 


Pat Hatuipay. 2001. The IHlustrated Rhododendron. Their Classification Portrayed 
Through the Artwork of Curtis’s Botanical Magazine. (ISBN 0-88192-510- 
1, hbk.). Timber Press, 133 SW Second Ave., Suite 450, Portland, OR 97204- 
3527, U.S.A. (Orders: www.timberpress.com, 800-327-5680, 503-227-2878, 
503-227-3070 fax). $69.95, hbk., 274 pp., 120 color illustrations, 4 line drawings, 
81/4" x 113/4" 

Emy S. SIEGERIST. 2001. Bulbophyllums and Their Allies. A Grower’s Guide. (ISBN 
0-88192-506-3, hbk.). Timber Press, 133 SW Second Ave., Suite 450, Portland, 
OR 97204-3527, U.S.A. (Orders: wwwtimberpress.com, 800-327-5680, 503- 
227-2878, 503-227-3070 fax). $34.95, hbk., 284 pp., 77 color photos, 6" x 9". 

Mary Toomey and Everett Leeps (photograhy editor Charles Chesshire). 2001 
An Illustrated Encyclopedia of Clematis. (ISBN 0-88192-508-X, hbk.). Timber 
Press, 133 SW Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: 
wwwtimberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). 
$59.95, hbk., 428 pp., 652 color photos, 5 b/w illustrations, 19 line draw- 
ings, 2 maps, 81/2" x 11" 


BarBARA JOE Hosuizakiand Rosin C. Moran. 200L. Fern Grower’s Manual. Revised 
and Expanded Edition. (ISBN 0-88192-495-4, hbk.). Timber Press, 133 SW 
Second Ave., Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: 
www.timberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). 
$59.95, hbk., 624 pp., 50 color photos, 165 b/w photos, 826 line drawings, 
S.y2 i. 

J.D. Vertrees. 2001. Japanese Maples. Momiji and Kaede. Third Edition Revised 
and Expanded by Peter Gregory. (ISBN 0-88192-501-2, hbk.). Timber Press, 
133 SW Second Ave., Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: 
wwwtimberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). 
$49.95, hbk., 332 pp., 312 color photos, 81/2" x 11". 

STEPHEN ANDERTON. 2001. Urban Sanctuaries. Peaceful Havens for the City Gar- 
dener. (ISBN 0-88192-502-0, hbk.). Timber Press, 133 SW Second Ave., Suite 
450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, 800- 


SIDA 19(4): 1209. 2001 


1210 BRIT.ORG/SIDA 19(4) 


327-5680, 503-227-2878, 503-227-3070 fax). $29.95, hbk., 144 pp, 148 color 
photos, 4 color plans, 4+ b/w plans, 81/2" x 11" 
Biographical 
MARGARET LYNN Brown. 2001. The Wild East. A Biography of the Great Smoky 
Mountains. (ISBN 0-8130-2093-X, hbk.). University Press of Florida, 15 NW 
15'" Street, Gainesville, FL 32611-2079, US.A. (Orders: www.upfl.com). $55.00, 
hbk, $24.95, pbk., +79 pp, 68 b&w photos, notes, bibliography, index, 6" x 9". 


SIDA 19(4): 1210. 2001 


SIDA, CONTRIBUTIONS TO BOTANY UPDATE AND 
REVIEWERS FOR VOLUME 19, 2000-2001 


All manuscript submissions are peer-reviewed. Below are names of 153 indi- 
viduals who kindly supported Sia through their time and expertise in review- 
ing manuscripts for volume 19. Your support is deeply appreciated. If by chance 
you reviewed a manuscript and your name was left out, the error rests solely on 
the shoulders of the editor and I truly apologize. Please bring any omissions to 
my attention. Volume 19 contains 1233 pages, 109 contributions, and 97 new 
names and new combinations. 


Dates of publication 
No. 1, pp. 1-234: 23 Sep 2000 
No. 2, pp. 235-444: 20 Dec 2000 
No. 3, pp. 445-766: 23 Aug 2001 
No. 4, pp. 767-1233: 28 Dec 2001 
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(NKU), Associate Editor; Félix Llamas (LEB), Contributing Spanish Editor. 


SIDA 19(4): 1211. 2001 


1212 


Ackerman, James D. 
Allen, Charles M. 
Allred, Kelly W. 

Amos, Bonnie 
Anderson, Gregory J. 
Anderson, Loran C 
Arias, Salvador 

Arnold, Anne Elizabeth 
Atwood, John T. 

Austin, Daniel F 


Baranski, Michael 


Simin Sarah 
Bossart aes nic 
Boyd, S 
a. oor S. 
Slice Dawid. 
Brown, Larry E 
ie ee T. 
Burks, Kathleen 


Calie, Patrick 
Chamberlain, David 
Chase, M 

eee ane Ib 
Clark, Lynn G. 
ne Theodore S. 
Coile, Nancy C 
Collins, Turner 

oe Edward A 

fill, Baynend 
Croat, Thomas B. 
Cusick, Allison W. 


SIDA 19(4): 1212. 2001 


REVIEWERS FOR VOLUME 19 


Davidse, Ger 
DeLaubentels, St 
Delprete, Piero G. 


Denham, Miriam L. 
Denton, Amy L. 
DeVore, Melanie 


Eckenwalder, James E. 
Endress, Mary E. 


Fishbein, Mark 

Ford, Bruce 
Freckmann, Robert W. 
Fryxell, Paul A 

Gabel, Mark L. 


Gandhi, K.N. 
Goetghebeur, Paul WS. 


Gomez-Sanchez, M. Sc. 
Maricel 


ricela 
Greuter, Werner 


Hatch, Stephan L. 
Hayden, W. John 
Henrickson, James 
Hess, William J. 
Hill, L. Michael 
Hils, Matthew H. 
Holmes, Walter C. 
Hong, Suk-Pyo 
Horner, John 
Janni, Kevin 
ee Richard J. 
ns, Robert 

ae Ronald L. 
Jones, Stanley 
Judziewicz, Emmet J. 
Keating, Richard 
Keil, David J. 
Kennedy, Robin C. 

Kiger, Robert W. 
oe H. 


Knapp, Sandra 
Knox, Eric 
seat 
Kral, Rober 

Krings, Alexander 
Kroh, Glenn C 


Lammers, Thomas G. 
Lamont, Eric 
Lasseigne, Alex A. 

7 ; 


Lelong, Michel G. 
Lemke, David E. 

Li, Jianhua 

Llamas, Félix 
Lopez-Ferrari, Ana R. 
Lott, Emily Jane 
Lowrey, Timothy Kk 


Macklin, James A. 


Mosyakin, Sergei L 
Naczi, Rob FC. 
Nelson, John B. 
Nesom, Guy 
Nixon, Elray 
O’Kennon, Robert J. 
Olsen, John S. 
Peterson, Paul M. 
Pinkava, Donald J. 
Pipoly, Il, John J. 
Pittman, ae Bertis 
Price, Michael G. 
Pruski, John F. 


BRIT.ORG/SIDA 19(4) 


Rabeler, Richard K. 


Reznicek, Anton (Tony) A. 
tson, Jon 
Robinson, Harold 
Sanders, Roger 
Schmid, Rudolf 
Smith, Alan R 
Solomon, James C. 
Sorrie, Bruce 
Spellenberg, Ric hard 
Stern, Pe 
Stevens, Peter E 
Stoynoff, Nick A. 
Strother, John L. 
Sundell, Eric 
Sutton, David L. 
Taylor, David D. 
Thieret, John W. 
Thomas, R. Dale 
Timbrook, Steven L. 
Trock, Debra 
Tucker, Gordon C. 
Turner, B.L. 


yan Welzen, PC. 
Villasenor, Jose Luis 
Vincent, Michael A. 


Watson, Frank D. 
Webster, Robert 
Wetter, Mark A 
Wiersema, John H. 
Wilbur, Robert L. 


Ae , Joseph K. 
worhingo, Richard D. 
Wujek, Daniel E. 


Wu eee Richard P. 


Yatskievych, George A. 


INDEX 


1213 


INDEX TO VOLUME 19, 2000-2001 
TITLES OF ARTICLES WITH AUTHORS 


A floristic and ethnobotanical account of 
the Josephstaal Forest Management 
Agreement Area, Papua New Guinea by 
W. TAKEUCHI—1:1 

A leaf blade anatomical survey of Muhlen- 
bergia (Poaceae: Muhlenbergiinae) by 
Paut M. Peterson and Yolanda Herrera- 
Arrieta—3:469 

A new combination for Sarcostemma 
cynanchoides var. hartwegii (Asclepia- 
daceae) by ALEXANDER KRINGS—1:13/ 

A new combination in Archibaccharis 
(Asteraceae: Astereae) by Guy L.Nesom— 


A new species of Saccoglossum (Orchi- 
daceae) from the Hans Meyer Range, 
New Ireland, Papua New Guinea by N.H.S. 
Howcroft—3:519 

A new species of Trepadonia (Astera- 
ceae: Vernonieae) from Peru by Haro.o 
RoBiNson and Hamitton BeLTRAN——1:111 

A new synonym for Eragrostis pilgeri 
(Poaceae: Eragrostideae) by J. Jost ALeGRIA 
OtlverA and Arturo GRANDA PAucAR— 
4:1157 

A new variety and four new combinations 
in Pittocaulon and Telanthophora 
(Asteraceae: Senecioneae) from Mexico 
by Bonnie L. CLARK—2:235 

A revision of Salvia section Heterosphace 
(Lamiaceae) in western North America 
by Jay B. Wacker and Wayne J. ELiseNs— 
3:571 

A Scutellaria (Lamiaceae) new to North 
Carolina and a key to the small-flowered 
Carolina congeners by ALEXANDER KRINGS 
and JosepH C. NeaL—3:735 


SIDA 19(4): 1213. 2001 


A synopsis of the ferns and fern allies of 
Nebraska, with maps of their distribution 


by Steven B. Rotesmeier, Rogert B. Kaul, and 
Davio M. SUTHERLAND —4: 1015 

A vascular flora survey of Calcasieu Parish, 
Louisiana by Ray NeyLANo, BiLue J. HOFFMAN, 
Mark MayrieLo, and Lowett E. UrsatscH— 
2:361 

Ada Nesta Ewan (19??—2000). A memoir 
by Anne S. BRaDbBURN—2:42 1 

Additions to the flora of Crater Mt., Papua 
New Guinea by W. TakEUCHI—2:237 

Alopecurus myosuroides and Sclero- 
chloa dura (Poaceae) new to Louisiana 
by JoHNn K. SaicHuk, CHARLES M. ALLEN, and 
WiLuiAM D. Reese —2:41 1 

Alternanthera sessilis (Amaranthaceae) 
new to Arkansas by CHristorHer S.Reio and 
Davio X WiLuiAMS—4:1197 

An anomalous population of Aster 
(Asteraceae: Astereae) sensu lato in 
Michigan by Guy L.Nesom—3:625 

Anagallis arvensis subsp. foemina 
(Primulaceae) new to Louisiana by 
CHarces M. ALLEN, SARA THAMES, PHILLIP PAUL, 
and Sevena DAWN NewmMan—4:1195 

Another new species of Tripogon 
(Poaceae) from India by C.N. Sunit and 
A.K. PrapeeP—4:803 

Arthraxon (Poaceae: Andropogoneae) 
new to South America by L.J.Dorr and S. 
Micue Nito—4:1191 

Batopilasia (Asteraceae: Astereae), a new 
genus from Chihuahua, Mexico by Guy 
L. Nesom and RicHarD D. Noves—1:79 

Bibliographical notes on the publication of 
Woolward’s monograph on the genus 


1214 


Masdevallia (Orchidaceae) by Guivo J. 
BRAEM—3:633 

Biological status of Argythamnia laevis 
(Euphorbiaceae) by B.L. Turner —3:621 

Blooming “behavior” in five species of 
Boerhavia (Nyctaginaceae) by RicHarp 
SPELLENBERG—2:3 11] 

Bolbitis thommankuthiana 
(Lomariopsidaceae), a new species of 
from India by SANTHOsH NampY—1:1 33 

Caesalpinia bonduc (Fabaceae) new to 
Louisiana by WILLIAM G. VermMILLION— 


Carex abscondita  (Cyperaceae: 
Careyanae) in Louisiana by Davin J. 
RoseN—3:727 

Carex cumberlandensis, a new species of 
section Careyanae (Cyperaceae) from 
the eastern United States of America by 
Rosert F.C. Naczi, and Rosert Kral, and 
Cuartes T. Bryson—4:993 

Cinna and Limnodea (Poaceae): not con- 
generic by Davio M. BRANDENBURG and JOHN 
W. THIERET—1:195 

Consideraciones sobre el origen de la flora 
arvense y ruderal del estado de Querétaro, 
Mexico by José Auretio COLMENERO RoBLes, 
CONCEPCION RODRIGUEZ JIMENEZ, aNd FERNANDEZ 
Nava RarAtL—4:1123 

Cryptocoryne beckettii (Araceae), a new 
aquatic plant in Texas by Davin J. Rosen— 
2:399 

Cupressus arizonica (Cupressaceae) new 
to the Davis Mountains of west Texas by 
JOHN P. Karces and James C. ZecH—3:719 

Cyperus (subg. Queenslandiella) 
hyalinus (Cyperaceae) new to the 
United States and the Western Hemi- 
sphere by RicHarD Carter and Ranpy L. 
Mears—2:345 


SIDA 19(4): 1214. 2001 


BRIT.ORG/SIDA 19(4) 


Cyperus sanguinolentus (Cyperaceae) 
new to the southeastern United States, 
and its relation to the supposed en- 
demic Cyperus louisianensis by Rich- 
ARD Carter and CHARLES T. Bryson—2:325 

Dipsacus fullonum (Dipsacaceae) and 
Verbesina walteri (Asteraceae), new to 
Texas by Jason R. SiIncHURST and Watter C. 
Ho_mes—3:723 

Discovery of Ardisia subgenus Acrardisia 
(Myrsinaceae) in Mesoamerica: Another 
boreotropical element? by JouNn J. Pipoty 
Ill and Jon M. Ricketson—2:275 

Distinction between Vitis blancoi and V. 
cinerea var. tomentosa (Vitaceae) by 
Barry L. Comeaux and Jianc Lu—1:123 

Documented chromosome numbers 
2001:1. Chromosome number of 
Lupinus havardii (Fabaceae) by BL. 
Turner and A.M. Poweit—3:639 

Eragrostis ancashensis (Poaceae: 
Chloridoideae), a new species from 
Ancash, Peru by Paut M. Peterson, NANCY 


ReFuLio Ropricuez, and Oscar TOVAR—1:65 

Establishing ethnobotanical conservation 
priorities: A case study of the Kallawaya 
pharmacopoeia by Kevin D. JANNi and Jo- 
SePH W. BasTIEN—2:387 

Expansion of the exotic aquatic plant 
Cryptocoryne beckettii (Araceae) in 
the San Marcos River, Texas by Rosert D. 
Doyie—4:1027 

Floristic documentation imperatives:some 
conclusions from contemporary surveys 
in Papua New Guinea by W. TakeucHi and 
M. GolmMan—3:445 

Gamochaeta simplicicaulis (Asteraceae: 
Gnaphalieae) in Georgia by Guy L. 
Nesom—2:413 

Genetic diversity in the Carex jamesii 


INDEX 


complex (Cyperaceae: sect. 
Phyllostachyae) with insights into the 
evolution and origin of the newly de- 
scribed species Carex timida by Bruct 
A.Forb and Rosert F.C. Naczi—4:885 

Geographic distribution patterns of the 
genus Ariocarpus (Cactaceae) in 
Tamaulipas, Mexico by José. G. Martinez- 
Avaos and Humperto SuzAn-Azpiri—1:1 75 

Geographic variation and taxonomy of 
North American species of Mirabilis, 
section Oxybaphoides (Nyctaginaceae) 
by RicHARD SPELLENBERG and Sercio R. 
RODRIGUEZ TUERINA—32539 

George Jones Goodman (1904-1999). A 
memoir by CHeryt A. LawSON—2:415 

Glochidion puberum (Euphorbiaceae) 
naturalized in southern Alabama by 
Miriam L. FeaRN and Lowett E. UrsatscH— 
3:71] 

Gnaphalium exilifolium (Asteraceae: 
Gnaphalieae) in Colorado and South 
Dakota by Guy L.Nesom—3:641 

Houstonia longifolia (Rubiaceae): newly 
documented for the flora of Texas by 
JASON R. SinGHURST and W.C. Holmes— 
4:1183 

Ipomoea sororia (Convolvulaceae),a new 
species from Yucatan, Mexico by Daniel 
F. Austin and Jose Luis Taria Munoz —4:807 

Joseph Andorfer Ewan, October 24, 
1909-December 5, 1999. A memoir by 
Anne S. BRaDBURN—1:221 

Laennecia turnerorum (Asteraceae: 
Astereae), a new species from Trans- 
Pecos Texas by G.L. Nesom—4:/89 

Legumes from the central part of the state 
of Chihuahua, Mexico by A. Eouarvo Es- 
TRADA C.and AtFonso Martinez M.—2:351 

Lespedeza cuneata (Fabaceae), a first 


SIDA 19(4): 1215. 2001 


1215 


record of its occurrence in Mexico by 
Epuarpo Estrapa C. and Carmen Yen M.— 
3:74] 

Neotypification of Enslenia albida and a 
new combination in Ampelamus for 
Cynanchum laeve (Apocynaceae: 
Asclepiadoideae) by ALEXANDER KriNGs— 
4:925 


New combinations in Chionolaena 
(Asteraceae: Gnaphalieae) by GL. 
NesomM—4:849 

New records in Pseudognaphalium 
(Asteraceae: Gnaphalieae) for the United 
States by G.L. Nesom—4:1185 

New subtribes for North American 
Astereae (Asteraceae) by Guy L.Nesom— 
2:263 

Nodulating legumes from the Tahoe Basin, 
California by MoHammad ATHAR and JAMES 
HARDING —1:205 

Nomenclatural change in the Digitaria 
cognata complex (Poaceae: Paniceae) 
by JoserH K. Wiprr—4:923 

Nomenclatural changes in Pennisetum 
(Poaceae: Paniceae) by JoserH K. Wiprr— 
3525 

Notes on some little known Amaranthus 
taxa ( thaceae) in the United 
States by Mixai CosTea, ANDREW SANDERS, and 
Gites WaINeES——4:9 7/5 

Notes on the increasing proportion of non- 
native angiosperms in the Missouri flora, 
with reports of three new genera for the 
state by Georce YarsKievycH and Jay A. 
RAVEILL—3:701 

Notes on variation in Pseudognaphalium 
obtusifolium (Asteraceae: Gnaphalieae) 
by Guy L.Nesom—3:615 

Noteworthy plants from north Florida. Vil 
by Loran C. ANDERSON—1:213 


1216 


Occurrence, distribution, and ecology of 
Alocasia, Caladium, Colocasia, and 
Xanthosoma (Araceae) in the south- 
eastern United States by Brett E. Serviss, 
Sipney T. McDaniet, and CuHartes T. BRYson— 
1:149 

Phyllostachys bambusoides (Poaceae: 
Bambuseae) previously unreported 
from Louisiana by Davio J. Rosen, STANLEY 
D. Jones, and JoserH K.WirrF—3:731 

Plant species-area relationships in ten 
north central Texas protected natural 
areas by Monica Granavos, Rosert J. 
O’KeNNOoN, and Bruce F. BEn7—4:1061 

Plateilema (Asteraceae: Helenieae) a new 
generic report for the United States by 
Bicuie L. TuRNeR—1:185 

Preliminary results toward a revision of the 
Amaranthus hybridus species com- 
plex (Amaranthaceae) by Minai Costea, 
ANDREW SANDERS, aNd Gites WAINES—4:93 1 

Pseudognaphalium austrotexanum 
(Asteraceae: Gnaphalieae),a new species 
from southeastern Texas and adjacent 
Mexico by Guy L. Nesom—3:507 

Radiate and eradiate individuals in Grind- 
elia nuda (Asteraceae) by R.T. Harms— 
B:/15 

Rediscovery of Ribes niveum (Grossu- 
lariaceae) in Colorado by Timotuy W. 
CHumtey and Ronatp L. HartMAN—2:407 

Reevaluation of Aylacophora and 
Paleaepappus (Asteraceae: Astereae) 
by José M. Boniracino and GiseLa SANCHO— 
B50 | 

Revision of Lobelia sect. Tupa (Campanu- 
laceae: Lobelioideae) by THomas G. 
LAMMERS— 1:87 

Rhynchospora leptocarpa (Cyperaceae), 
an overlooked species of the southeast- 


SIDA 19(4): 1216. 2001 


BRIT.ORG/SIDA 19(4) 


ern United States by Bruce A. Sorric— 
1:139 

Rhynchospora zacualtipanensis and 
Eleocharis moorei, two new Cypera- 
ceae from Mexico by Mark T. StronG and 
M.S. GONZALEZ 1:115 

Rupert C. Barneby (October 6, 1911- 
December 5, 2000).A memoir by RicHaRD 
SPELLENBERG—3:745 

Scleria lacustris (Cyperaceae), an aquatic 
and wetland sedge introduced to 
Florida by Cotette C. Jacono—4:1 163 

Senecio quaylei (Asteraceae: Sene- 
cioneae),a new species from north cen- 
tral Texas, U.S.A by THeopore M. Barkley — 
2:285 

Solidago villosicarpa (Asteraceae: 
Astereae), a rare new southeastern 
coastal plain endemic by RJ. LEBLono— 
2:29) 


Fiizanir 
CLIZONDC 


— 


South American skullcap (Scutellaria 
racemosa: Lamiaceae) in the southeast- 
ern United States by ALEXANDER KRINGs and 
Josep C. Neat—4:1171 

Status of Quercus _ leana and Quercus _ 
runcinata (Fagaceae) in Illinois by Gor- 
DON C. Tucker and JOHN E. EsinceR—4:1073 

Stenotis (Rubiaceae), a new segregate 
genus from Baja California, Mexico by 
Epwarb E, TerrELL—4:899 

Structure of populations of otate (Otatea 
acuminata subsp.aztecorum: Poaceae) 
in harvested stands by Jose Martin 
Vazquez Lopez, Bruce F. Benz, Micuel OLvERA 


9° 20)) 
Z2.5U 1 


VARGAS, aNd SERGIO GraF MONTERC 
Stylogyne aguarunana (Myrsinaceae) a 
new species from Amazonas, Peru by 


JOHN J. Pipoty Hl and Jon M. RickeTtson— 


Systematics of the Carex jamesii complex 


INDEX 


(Cyperaceae: sect. Phyllostachyae) by 
Rosert F.C. Naczi and Bruce A. FoRD—4:853 

Taxonomic notes on Keysseria and 
Pytinicarpa (Asteraceae: Astereae, 
Lageniferinae) by Guy L. Nesom—3:5 13 

Taxonomic review of Chrysogonum 
(Asteraceae: Heliantheae) by GL. 
Nesom—4:8 1 1 

Taxonomic review of Houstonia acerosa 
and H. palmeri, with notes on Hedyotis 
and Oldenlandia (Rubiaceae) by Eowarbd 
E. TERRELL —4:9 13 

Taxonomy of Stenaria (Rubiaceae: Hedyo- 
tideae),a new genus including Hedyotis 
nigricans by Eowaro E. TerretL—3:591 

Taxonomy of the Dichotoma group of 
Dichanthelium (Poaceae) by RicHaro J. 
LeBLOND—4:821 

The genus Hesperoyucca (Agavaceae) in 
the western United States and Mexico: 
new nomenclatural combinations by 
Karen H. CLARY —4:839 

The intriguing case of Cypripedium 
crossii, its priority over Cypripedium 
callosum and its transfer to the genus 
Paphiopedilum by Guivo J. Bracm and 
KARLHEINZ SENGHAS—2:249 

The vascular flora of Amite County, Missis- 
sippi by Mac H. ALroro—3:645 

The vascular flora of Madison County, Texas 
by Amanpa K. Nett and HucH D. Witson— 
4:1083 

Themeda quadrivalvis (Poaceae: 
Andropogoneae) in Kansas: an exotic 
plant introduced from birdseed by E. 
Gene TOWNE and IRALEE BARNARD—1:201 


SIDA 19(4): 1217. 2001 


1217 


Thymelaea passerina (Thymelaeaceae) 
new to Texas by Watter C. HOLMes, JOHN F. 
Pruski, and JASON R. SINGHURST—2:403 

Two new species of Impatiens (Balsami- 
naceae) from India by MukresH Kumar and 
STEPHEN SEQUIERA—4:795 

Two new species of Liatris series 
Punctatae (Asteraceae: Eupatorieae) 
centered in north central Texas by G.L. 
Nesom and R.J. O'KENNON—4:/67 

Use of variety and subspecies and new 
varietal combinations for Styrax 
platanifolius (Styracaceae) by B.L. Turner 
and Guy L. Nesom—2:257 

Validation of the name Orobanche 
ludoviciana subsp. multiflora 
(Orobanchaceae) by HeatHer L.Wuite and 
Watter C. HOLMES—3:623 

Vascular plant types in the Arizona State 
University Herbarium by STEFANIE M. ICKERT- 
Bono and Donato J. PINKAvA—4:1039 

Vascular plants new to Kentucky by J. RicH- 


ARD Apgott, RALPH L. THOMPSON, and Ruby A. 
GELIS—4:1199 

Vines of a temperate state: still 
undercollected? by ALEXANDER KRINGS— 
4:1147 

Which non-native plants are included in 
floristic accounts? by Guy L. Nesom— 
1:189 

Who named Euthamia (Compositae: 
Astereae) and when? by Joun L. 
STROTHER—1:219 

Wood and bark anatomy of 
Achatocarpaceae by SHERWIN CARLQUIST— 
1:7] 


1218 


BRIT.ORG/SIDA 19(4) 


AUTHOR INDEX 


Abbott, J. Richard—4:1199 
Alegria Olivera, J. Jos¢—4:1 157 
Alford, Mac H. Alford —3:645 
Allen, Charles M.—2:411 

Allen, Charles M.—4:1195 
Anderson, Loran C.—1:213 
Athar, Mohammad—1:205 
Austin, Daniel F—4:807 

Barkley, Theodore M.—2:285 
Barnard, lralee—1:201 

Bastien, Joseph W.—2:387 
Beltran, Hamilton—1:111 

Benz, Bruce F—2:301; 4:1061 
Bonifacino, José M.—3:53] 
Bradburn, Anne $.—1:221; 2:421 
Braem, Guido J.—2:249; 3:633 
Brandenberg, David M.—1:195 
Bryson, Charles 1—1:149; 2:325; 4:993 
Carlquist, Sherwin—1:71 

Carter, Richard—2:325, 345 
Chumley, Timothy W.—2:407 
Clark, Bonnie L.—2:235 

Clary, Karen H—4:839 
Colmenero Robles, José Aurelio—4:1123 
Comeaux, Barry L—1:123 
Costea, Mihai—4:931,975 

Dorr, L.J—4:119] 

Doyle, Robert D.—4:1027 
Ebinger, John E.—4:1073 
Elisens, Wayne J—3:571 

Estrada C., A. Eduardo—2:351;3:741 
Fearn, Miriam L.—3:/11 

Ford, Bruce A.—4:853, 885 

Gelis, Rudy A.—4:1199 

Golman, M.—3:445 
Gonzdlez-Elizondo, M.S—1:115 
Graf Montero, Sergio—2:301 
Granados, Monica—4:1061 
Granda Paucar, Arturo—4:1157 


SIDA 19(4): 1218. 2001 


Harding, James—1:205 

Harms, R.1.—3:/15 

Hartman, Ronald L.—2:407 

Herrera-Arrieta, Yolanda —3:469 

Hoffman, Billie J —2:361 

Holmes, Walter C.—2:403; 3:623, 723: 
4:1183 

Howcroft, N.H.$.—3:519 

Ickert-Bond, Stefanie M—4:1039 

Jacono. Colette C—4:1163 

Janni, Kevin D.—2:387 

Jones, Stanley D.—3:731 

Karges. John P-—3:719 

Kaul, Robert B.—4:1015 

Kral, Robert—4:993 

Krings, Alexander—1:137; 3:735; 4:925, 
1147,1171 

Kumar, Muktesh—4:795 

Lammers, Thomas G.—1:87 

Lawson, Cheryl A.—2:415 

LeBlond, R.J.—2:291; 4:821 

Lu, Jiang—1:123 

Martinez M., Alfonso—2:35 1 

Martinez-Avalos, José. G.—1:175 

Mayfield, Mark—2:361 

McDaniel, Sidney T.—1:149 

Mears, Randy L.—2:345 

Naczi, Robert F.C.—4:853, 885, 993 

Nampy, Santhosh—1:133 

Nava Rafael, Fernandez—4:1123 

Neal, Joseph C.—3:735; 4:1171 

Neill, Amanda K.—4:1083 

Nesom, G.L.—1:79, 85, 189; 2:257, 263,413; 
3:507,513,615, 625,641; 4:789, 767, 811, 
849,1185 

Newman, Selena Dawn—4:1 195 

Neyland, Ray—2:361 

Nino, S. Miguel—4:1191 

Noyes, Richard D.—1:79 


AUTHOR INDEX 


O’Kennon, R.J.—4:/67, 1061 

Olvera Vargas, Miguel—2:301 

Paul, Phillip —4:1195 

Peterson, Paul M.—1:65; 3:469 

Pinkava, Donald J.—4:1039 

Pipoly, John J, II]l—2:269, 275 

Powell, A.M.—3:639 

Pradeep, A.K.—4:803 

Pruski, John F—2:403 

Raveill, Jay A.—3:701 

Reese, William D.—2:411 

Refulio Rodriguez, Nancy—1:65 

Reid, Christopher S.—4:1197 
Ricketson, Jon M—2:269, 275 
Robinson, Harold—1:111 

Rodriguez Jiménez, Concepcion—4:1 123 
Rodriguez Tijerina, Sergio R.—3:539 
Rolfsmeier, Steven B.—4:1015 

Rosen, David J.—2:399; 3:727, 731 
Saichuk, Jonn K—2:411 

Sancho, Gisela—3:531 

Sanders, Andrew—4:931,975 

Senghas, Karlheinz—2:249 

Sequiera, Stephen—4:795 

Serviss, Brett E—1:149 

Singhurst, Jason R.—2:403; 3:723; 4:1183 
Sorrie, Bruce A.—1:139 

Spellenberg, Richard—2:31 1; 3:539, 745 


SIDA 19(4): 1219. 2001 


Strong, Mark T.—1:115 

Strother, John L.—1:219 

Sunil, C.N.—4:803 

Sutherland, David M—4:1015 
Suzan-Azpiri, Humberto—1:175 
Takeuchi, W.—1:1; 2:237; 3:445 
Tapia Munoz, Jose Luis—4:807 
Terrell, Edward E.—3:591;4:889, 913 
Thames, Sara—4:1195 

Thompson, Ralph L—4:1199 
Thieret, John W.—1:195 

Tovar, Oscar—1:65 

Towne, E.Gene—1:201 

Tucker, Gordon C.—4:1073 

Turner, B.L.—1:185; 2:257; 3:621, 639 
Urbatsch, Lowell E.—2:361;3:711 
Vazquez Lopez, Jose Martin 
Vermillion, William G.—4:118] 
Waines, Giles —4:931,975 

Walker, Jay B—3:571 

White, Heather L.—3:623 

Williams, David X—4:1197 

Wilson, Hugh D.—4:1083 

Wipff, Joseph K—3:523, 731; 4:923 
Yatskievych, George—3:/01 

Yen M., Carmen—3:741 

Zeck, James C.—3:/19 


1219 


1220 


BRIT.ORG/SIDA 19(4) 


BOTANICAL NAMES 
New names in bold face 


Abutilon theophrasti 1:213 
Acalypha deamii 4:1199 
Achatocarpaceae 1:71 
Achatocarpus 
nigricans 1:71 
praecox 1:71 
Aglaia saxonii 1:11, 12 (illus) 
Ajuga reptans 1:211 
Alocasia 1:149 
lancifolia 1:19 
macrorrhizos var. macrorrhizos 
1:153, 154 (illus) 
odora 1:156 
plumbea 1:156 
Alopecurus myosuroides 2:41 1 
Alstroemeria psittacina 3:645 
Alternanthera sessilis 4:1197 
Amaranthaceae 4:931, 1197 
Amaranthus 
blitum subsp. blitum 4:975, 980 
(photo), 987 (SEM) 
blitum subsp. emarginatus var. 
emarginatus 4:979, 982 (photo), 
987 (SEM) 
blitum subsp. emarginatus var. 
pseudogracilis 4:981, 983 (photo) 
blitum subsp. oleraceus 4:984, 985 
(photo), 987 (SEM 
caudatus 4:940 (SEM), 943 
cruentus 4:945, 946 (photo) 
graecizans 4:977,987 (SEM), 988, 989 
graecizans subsp. sylvestris 4:987 
(SEM), 989, 990 (photo) 


= 


7 


hybridus 4:931,938 (SEM), 940 (SEM), 


hybridus subsp. hybridus 4:950, 951 
(photo) 


SIDA 19(4): 1220. 2001 


hybridus subsp. quitensis 4:940 (SEM), 
955 


hypochondriacus 4:957, 958 (photo) 
powellii 4:960 
powellii subsp. bouchonii 4:938 
(SEM), 940 (SEM), 964, 965 (photo) 
powellii subsp. powellii 4:938 (SEM), 
940 (SEM), 961, 962 (photo) 
retroflexus 4:938 (SEM), 940 (SEM), 966 
viridis 4:986 
Ampelamus 4:925 
laevis 4:927 
Anagallis arvensis subsp. foemina 4:1195 
Andropogoneae 1:201 
Anthoxanthum aristatum 4:1199 
Antidesma katikii 1:23 
Aphanes microcarpa 4:1200 
Apocynaceae: Asclepiadoideae 4:925 
Araceae 1:149; 23399; 4:1027 
Archibaccharis trichotoma 1:85 
Archidendron hispidum 2:239, 240 (photo) 
Ardisia 2:275 
rarescens 2:278, 279 (illus) 
Argythamnia laevis 3:621 
Ariocarpus 1:175 
agavoides 1:175,179 
kotschoubeyanus 1:175, 179 
retusus subsp. retusus 1:175, 180 
retusus subsp. trigonus 1:175, 180 
Arthraxon 4:1191 
hispidus 4:1191 
Asclepiadaceae 1:137 
Aster laevis 1:211 
Asteraceae 1:79, 85, 111, 185, 217; 2:235, 
200):20 1,413; 31507, 513,531.615,025, 
639, 723; 4:767, 789, 811,849 
Asteraceae subtribe Astranthiinae 2:265 


BOTANICAL NAMES 


Asteraceae subtribe Boltoniinae 2:266 

Asteraceae subtribe Chaetopappinae 
2:264 

Asteraceae subtribe Pentachaetinae 


Asteraceae: Gnaphalieae 4:1185 
Astereae 1:79,85,217;2:263,291;3:513,531, 


Aureolaria virginica 1:213 
Aylacophora 3:53] 
deserticola 3:533 (illus) 
Balsaminaceae 4:795 
Bambuseae 3:731 
Barringtonia josephstaalensis 1:9, 19 
(illus) 
Batopilasia 1:81 
byei 1:81, 82 (illus) 
Boerhavia 2:31] 
Bolbitis 
semincordata 1:133 
thommankuthiana 1:1 33, 134 (illus), 
135 (photos) 
Boltonia 1:80 
asteroides 1:213 
Cactaceae 1:175 
Caesalpinia bonduc 4:1181, 1182 (map) 
Callisia repens 1:213 
Calycosia mamosei 1:14, 15 (illus) 
Campanulaceae 1:87 
Canarium acutifolium var. pioriverensis 


Carex 4:853 

abscondita 3:727, 728 (map); 4:1004 
(ma 

austrocaroliniana 4:1 200 

crebriflora 1:213 

cumberlandensis 4:993,994, 996, 
(illus), 997 (illus), 1002 (map) 

jamesii 4:853, 862 (photo), 863 (illus), 


we 


SIDA 19(4): 1221. 2001 


1221 


864 (photo), 865 (map), 873, 885 
juniperorum 4:862, 864 (photos), 866 
(map), 877, 885 
styloflexa 1:213 
timida 4:862 (photo), 863 (illus 
(photo), 866 (map), 879, 885 
Casearia erythrocarpa 1:20 
Centunculus minimus 4:1 200 
Chasmanthium latifolium 1:213 
Chenopodium murale 1:212 
Chinolaena 4:849 
costaricensis 4:850 
cryptocephala 4:850 
durangensis 4:850 
macdonaldii 4:850 
salicifolia 4:850 
Chloracantha 1:80 
Chloridoideae 1:65 
Chrysogonum 4:81 1 
virginianum 4:814 (illus), 815, 816 
(map) 
virginianum var. brevistolon 4:8 17 
Cinna 1:195 
Cladium 1:149, 1:157 
bicolor 1:157, 1:158 (illus) 
Cleistanthus 1:20 
Clinopodium gracile 3:645 
Colocasia 1:149, 1:159 
esculenta 1:160 
var.antiquorum 1:164 
var. aquatilis 1:161, 1:162 (illus) 
var.euchlora 1:166 
var.esculenta 1:165 
var. fontanesii 1:166 
var. globulifera 1:166 
var. illustris 1:167 
var.nymphaeifolia 1:167, 168 
(illus) 
gigantean 1:167 


ear 


, 864 


1222 


Coniogramme macrophylla 2:239 

Conobea multifida 1:212 

Convolvulaceae 4:807 

Croton willdenowii 1:213 

Cryptocoryne beckettii 2:399, 400 (illus); 
4:1027 

Cupressaceae 3:/19 

Cupressus arizonica 3:719 


Cyperaceae 1:115, 139; 2:325, 345; 3:727; 
4:1163 

Cyperaceae sect. Cayeyanae 4:993 
Cyperaceae sect. Phyllostachyae 4:853, 885 
Cyperus 

hyalinus 2:345, 348 (color photos, 

map) 
louisianensis 2:325, 333 (color photos) 


pseudovegetus 1:213 
retrofractus 1:212 
sanguinolentus 2:325, 333 (color pho- 
tos), 337 (illus), 338 (map) 
Cypripedium 
callosum 2:249 
crossil 2:249 
Desmodium rotundifolium 1:213 
Dichanthelium 4:82 1 
annulum 4:826 
caerulescens 4:828 
dichotomum var.dichotomum 4:828 
var. nitidum 4:829 
var. ramulosum 4:830 


— 


var. roanokense 4:83 1 
lucidum 4:83 1 
mattamuskeetense 4:832 
nudicaule 4:833 
sphagnicola 4:834 
yadkinense 4:835 

Dichotoma group (Poaceae) 4:821 


SIDA 19(4): 1222. 2001 


BRIT.ORG/SIDA 19(4) 


Digitaria 

cognata 4:923 

pubiflora 4:923 
Dipsacaceae 3:723 
Dipsacus fullonum 3:723 
Ditaxis humilis 3:622 (map) 

forma laevis 3:622 
Doellingeria umbellate 3:625 
Dryopteris ludoviciana 3:645 
Eleocharis moorei 1:118, 119 (illus) 
Elymus wiegandii 4:1 200 
Enslenia albida 4:925 
Equisetum x ferrissii 4:1200 
Eragrostis 

ancashensis 1:66, 67 (illus) 

ancashensis 4:1157 

macrothyrsa 1:70 

magna 1:66 

pilgeri 4:1157,1158 
Erigeron byei 1:81 
Erysimum hieraciifolium 4:1201 
Etlingera 1:24 
Euphorbia texana 2:361, 364 
Euphorbiaceae 3:621, 711 
Euthamia 1:217 

graminifolia 1:218 

tenuifolia 1:218 
Fabaceae 1:205; 2:351; 3:643, 741; 

4:1181,1073 
Fatoua villosa 3:701, 702 
Fimbristylis decipiens 1:213 
Funastrum cynanchoides var. hartwegii 
2 


— 


Gamochaeta simplicicaulis 2:41 3 
Ghaphalieae 2:413 
Glochidion 

chondrocarpum 1:20 

puberum 3:711, 713 (color photo) 
Gnaphalieae 3:507, 615, 639; 4:849 
Gnaphalium exilifolium 3:639 


BOTANICAL NAMES 


Grindelia nuda 3:715, 716,717 (photos) 
Grossulariaceae 2:407 
Hedyotis 4:913 
Helenieae 1:185 
Heliantheae 4:811 
Helicia affinis 1:21 
Hesperoyucca 4:839, 842 
newberryi 4:845 
peninsularis 4:845 
whipplei 4:842, 843 (illus) 
Homalomena magna 1:19 
Homochrominae 2:267 
Houstonia 
acerosa 4:91 3 
var. acerosa 4:916 
var. polypremoides 4:916 
var. potosina 4:918 
var. tamaulipana 4:916 
longifolia 4:1183 
palmeri 4:913,919 
var. muzquizana 4:919 
var. palmeri 4:919 
Hypericum gymnanthum 1:214 
Impatiens 4:795 
sholayarensis 4:795 
violacea 4:798 
Ipomoea 4:807 
indica 3:645 
sororia 4:807 
Keysseria 3:514 
sect. Sandwicactis 3:514 
Kyllinga squamulata 1:214 
Laennecia 4:789 
turnerorum 4:789 
Lamiaceae 3:571 
Lathyrus tuberosus 4:1201 
Leersia lenticularis 1:214 
Lespedeza cuneata 3:741; 3:742 (map) 
Leucothoe fontanesiana 4:1201 


SIDA 19(4): 1223. 2001 


1223 


Liatris 4:767 
aestivalis 4:/768 
aestivalis 4:775 
glandulosa 4:778 
glandulosa 4:785 
mucronata 4:775, 785 
series punctatae 4:767 
Limnodea 1:195 
Linum macrocarpum 1:212 
Lipocarpha aristulata 1:212 
Lithospermum incisum 1:214 
Lobelia 1:87 
bridgesii 1:105 
excelsa x L. polyphylla 1:105 
excelsa 1:98 
polyphylla 1:94 
tupa 1:100 
Lomariopsidaceae 1:133 
Lupinus 
harvardii 3:643 
perennis 4:1201 
Masdevallia 3:633 
Mecardonia procumbens 1:212 
Microcos 1:23 
Mirabilis 3:539 
laevis 3:541 (photo of fruits); 3:545, 
546, 547 (maps) 
var. crassifolia 3:549 
var. retrorsa 3:553 
var. Villosa3:551 
oligantha 3:556, 557 (map) 
oxybaphoides 3:541 (photo of fruits), 
543,557 (map) 
tenuiloba 3:541 (photo of fruits), 555, 
(map) 
Muhlenbergia 3:469 
asperifolia 3:478 (photo) 
brevivaginata 3:476 (photo) 
ciliata 3:476 (photo) 
curvula 3:480 (photo) 


1224 


dubia 3:476 (photo) 
expansa 3:480 (photo) 
gigantea 3:478 (photo) 
japonica 3:478 (photo) 
lehmanniana 3:478 (photo) 
lindheimeri 3:480 (photo) 
microsperma 3:480 (photo) 
pauciflora 3:476 (photo) 
pubigluma 3:476 (photo) 
rigida 3:478 (photo) 
schreberi 3:478 (photo) 
Muhlenbergiinae 3:469 
Murdannia keisak 1:214 
Myrsinaceae 2:269; 2:275 
Nardophyllum 
bracteolatum 3:533 (illus) 
bryoides 3:533 (illus) 
deserticola 3:531 
patagonicum 3:531 
Nyctaginaceae 2:311;3:539 
Oclemena 
xblakei 3:629 
nemoralis 3:625 
Oenanthe javanica 3:701, 704 
Oldenlandia 4:913 
Orchidaceae 2:249; 3:519, 633 
Orobanchaceae 3:623 
Orobanche ludovici b 
33623 
Otatea acuminata subsp. aztecorum 2:301 
Ottelia alismoides 3:701, 706 
Oxycaryum cubense 1:214 
Paederia foetida 1:214 
Paleaepappus patagonicus 3:533 (illus) 
Paniceae 3:523 
Paphiopedilum 2:249 
crossii 2:253;2:252 (color photo) 
var. potentianum 2:254 
var. sublaeve 2:254 
Paspalum conjugatum 1:214 


p.multiflora 


SIDA 19(4): 1224. 2001 


BRIT.ORG/SIDA 19(4) 


Pennisetum 
pennisetiforme 3:527, 528, 529 
(photos) 
setigerum 3:5 26 
somalensis 3:52/7 
Phaulothamnus spinescens 1:71 
Photinia serratifolia 3:645 
Phyllostachys bambusoides 3:731 
Physalis carpenteri 3:645 
Piper arfakianum 2:241 
Pittocaulon 
hintonii var. cerrograndensis 2:235 
velatum var. tzimolensis 2:235 
Plantago rugelii 1:214 
Plateilema 1:185 
palmeri 1:185, 1:186 (map) 
Platycladus orientalis 1:214 
Poaceae 1:65, 195, 201; 2:301, 411; 3:469, 
523,737 803 621 
Poaceae: Eragrostideae 4:1 
Polygonum 
cilinode 4:1201 
lapathifolium 1:214 
Primulaceae 4:1195 
Pseudognaphalium 
arizonicum 4:1185 
attenuatum 4:1186 
austrotexanum 3:507, 508 (illus), 510 


—_— 


af 


(map) 
helleri 4:1189 
jaliscense 4:1187 
luteoalbum 4:1188 
micradenium 3:618 
obtusifolium 3:615 
var.micradenium 3:618 
Stramineum 4:1189 
viscosum 3:509, 510 (map) 
Psychotria 1:22 
dipteropoda 1:21 
mayana 1:1 7, 18 (illus) 
Pycnanthemum flexuosum 1:214 


BOTANICAL NAMES 


Pytinicarpa 

neocaledonica 3:517 

pickeringii 3:516 

sarasinil 3:517 
Quercus 

imbricaria 4:1079 (illus) 

x leana 4:1073, 1079 (illus) 

x runcinata 4:1073, 1079 (illus) 

velutina 4:1079 (illus) 
Rauvolfia moluccana 1:19 
Rhynchospora 

capitellata 1:139, 155 (map) 

leptocarpa 1:139, 144 (map) 

macrostachya 1:215 

thornei 1:212 

zacualtipanensis 1:115, 116 (illus) 
Rhyticaryum novoguineense 1:21 
Ribes niveum 2:407 
Rubiaceae 3:591;4:1183, 899,913 
Saccoglossum takeuchii 3:519, 520 (illus) 
Salvia 3:571 

henryi 3:576 (illus), 579 (map), 580 


— 


— 


roemeriana 3:576 (illus), 579 (map), 


582, 583 (illus) 
sect. heterosphace 3:57] 
summa 3:576 (illus), 579 (map), 586 
Sarcostemma cynanchoides var. hartwegii 
1:137 
Schisandra glabra 1:215 
Scirpus koilolepis 1:215 
Scleria lacustris 4:1163, 1164 (map), 1168 
(illus) 
Sclerochloa dura 2:411 
Scutellaria 
laterifolia 3:738 (illus) 
nervosa 3:738 (illus) 
parvula 3:738 (illus) 
racemosa 3:735, 737, (illus), 738 (illus) 
racemosa 4:1171,1173 (map) 


SIDA 19(4): 1225, 2001 


1225 


Senecio quaylei 2:286, 287 (illus), 288 
(color photo) 
Senecioneae 2:235, 285 
Silene nivea. 4:1202 
Solidago 
auriculata 3:645 
villosicarpa 2:292, 293 (illus), 294,295 
(color photos) 
Spermacoce 
glabra 1:215 
verticillata 1:215 
Stenaria 3:591,592 
butterwickiae 3:595, 596 (illus) 
mullerae 3:593, (photo of seeds), 597 
var. mullerae 3:598, 599 (photo) 
var. pooleana 3:598 
nigricans 3: 593 (photo of seeds), 600 
var. breviflora 3:605 
var. floridana 3:605 
var. gypsophila 3:606 
var. nigricans 3:601, 602 (photo) 
rupicola 3:593 (photo of seeds), 608, 
609 (photo) 
Stenaria umbratilis 3:593 (ohoto of seeds), 
610,611,612 (photo) 
var. brevipedicellata 3:611, 613 
(photo) 
Stenotis 4:899, 901 
Stenotis arenaria 4:900 (seed), 902 
Stenotis asperuloides 4:900 (seed), 903, 
905 (illus) 
var. asperuloides 4:904 
var. brandegeana 4:904 
australis 4:900 (seed), 906 
brevipes 4:907 
gracilenta 4:908 
mucronata 4:900 (seed), 909 
peninsularis 4:910 


Stylogyne aguarunana 2:269, 271 (illus), 


272 (map) 


1226 


Styracaceae 2:257 
Styrax 
platanifolius var. mollis 2:261 
var. texanus 2:26] 
var. youngiae 2:261 
Syzygium hylochare 2:241, 242 (photo) 
Tapeinochilos 1:20 
recurvatum 1:20 
Taraxacum laevigatum 1:212 
Telanthophora 
cobanensis var.molinae 2:236 
grandifolia var. serraquitchensis 
22250 
sublaciniatus 2:236 
Themeda quadrivalvis 1:201 
Thunbergia alata 1:212 
Thymelaea passerina 2:403, 404 (illus) 
Thymelaeaceae 2:403 


BRIT.ORG/SIDA 19(4) 


Thymophylla tenuiloba 1:212 
Trepadonia oppositifolia 1:1 11,112 (photo) 
Tripogon 4:803 
ravianus 4:803 
Vaccinium tenellum 1:215 
Verbesina walteri 3:723 
Vernonieae 1:111 
Versteegia grandifolia 1:22 
Vicia tetrasperma 4:1202 
Vitaceae 1:123 
Vitis 
blancoi 1:123 
cinerea var. tomentosa 1:123, 129 
Wenzelia dolichophylla 1:22 
Xanthosoma 1:149, 169 
sagittifolium 1:169, 170 (illus), 213 
violaceum 1:172 
Zanthoxylum conspersipunctatum 1:23 


SUBJECT 


Alabama 2:325;3:711 

Amazonas, Peru 2:269 

Amite County, Mississippi 3:645 

Anatomy 1:71;3:469 

Ancash, Peru 1:65 

Aquatic 2:399; 4:1027, 1163 

Arizona 4:1039 

Arizona State University Herbarium 4:1039 

Arkansas 4:1197 

Baja California, Mexico 4:899 

Barneby, Rupert C. (1911-2000) 3:745, 751 
(color photo) 

Birdseed 1:201 

Blooming behavior 2:311 

Boreotropical 2:275 

Calcasieu Parish, Louisiana 2:361 

California 1:205 

Central America 4:849 

Chihuahua, Mexico 1:79; 2:351 


SIDA 19(4): 1226. 2001 


Chromosome number 3:643 

Colorado 2:407; 3:639 

Crater Mountain Wildlife Management 
Area 2:237 

Davis Mountains 3:719 

Eastern United States 4:993 

Endemic 2:291, 325 

Ethnobotany 1:1, 89; 2:301, 387 

Ewan, Joseph Andorfer 1:219, 221 (photo) 

Ewan, Nesta Dunn 2:421, 422, 423 (color/ 
bw photos) 

Exotic 1:201;2:399: 4:1027 

Fern allies 1:133;4:1015 

Ferns 4:1015 

Florida 1:211;2:345; 4:1163 

Floristics 1:1, 189; 2:361; 3:445, 645 

Georgia 2:325, 413 

Goodman, George Jones 2:415, 418, 419 
(color/bw photos) 


SUBJECT 


Hans Meyer Range, New Ireland, Papua 
New Guinea 3:519 

Hawail 4:1186 

Illinois 4:1073 

India 1:133; 4:795, 803 

Indiana 4:1189 

Introduced 4:1163 

Josephstaal Forest Management Agree- 
ment Area 1:1 

Kallawaya pharmacopoeia 2:387 

Kansas 1:201 

Kentucky 4:879, 1199 

Kerala, India 4:795, 803 

Louisiana 2:325, 361,411;3:727,731;4:1181, 
1195 

Madison County, Texas 4:1083 

Mesoamerica 2:275 

Mexico 1:79,115,175;2:235;3:351,507, 741; 
4:1123, 839,849 

Michigan 3:625 

Mississippi 2:325; 3:645 

Missourl 3:701 

Nebraska 4:1015 

New Mexico 4:1188 

New York 4:1189 

Nodulating legumes 1:205 

Non-native angiosperms 3:701 

North America 2:263 

North Carolina 2:291;3:735;4:1147 

North Central Texas 2:285; 4:767, 1061 

North Florida 1:211 

Oklahoma 4:767 

Otate 2:301 


SIDA 19(4): 1227. 2001 


1227 


Papua New Guinea 1:1; 2:237;3:445,519 

Parataxonomist 3:449 

Peru 1:65, 111;2:269; 4:1157 

Phenology 2:31] 

Protected natural areas 4:1061 

Queretaro, Mexico 4:1123 

Rare species 2:291 

San Marcos, Texas 4:1027 

Sea bean 4:1181 

South America 2:387; 4:1191 

South American skullcap 4:1171 

South Dakota 3:639 

Southeastern Coastal Plain 2:291 

Southeastern Texas 3:507 

Southeastern United States 1:139, 149; 
2:291, 325;4:1147,1171 

Tahoe Basin, California 1:205 

Tamaulipas, Mexico 1:175 

Texas 1:185:2:285, 399, 403:3:507;621,623, 
719, 723; 4:767, 789, 1027, 1061, 1083, 
1183,1188 

Trans-Pecos Texas 4:789 

United States 1:185; 2:345;4:1185,975 

Vascular plant types 4:1039 

Vines 4:1147 

Weeds 4:1123 

West Texas 3:719 

Western Hemisphere 2:345 

Western North America 3:571 

Western United States 4:839 

Woolward, Florence Helen 3:633 

Woolward’s Monograph 3:633 

Yucatan, Mexico 4:807 


1228 


BRIT.ORG/SIDA 19(4) 


New NAMES AND NEw ComsiNaTIONS 
Volume 19, 2000-2001 


Aglaia saxonii Takeuchi, sp. nov.—1:1 1 

Amaranthus bli bsp. emarginatus var. 
pseudogracilis (Thell.) Costea,comb.et 
stat. nov.— 4:98 1 

Amaranthus blitum subsp. oleraceus (L.) 
Costea, comb. nov.—4:984 

Amaranthus hybridus subsp. quitensis 
(Kunth) Costea & Carretero, comb.nov.— 
4:955 

Amaranthus powellii subsp. bouchonii 
(Thell.) Costea & Carretero, comb.nov.— 
4:964 

Ampelamus laevis (Michx.) Krings, comb. 
nov.—4:927 

Archibaccharis trichotoma (Klatt) Nesom, 
comb. nov.—1:85 

Asteraceae subtribe Ast 


gte ss KI 


Carex timida Naczi & B.A. Ford, sp. nov.— 


Chionolaena costaricensis (Nesom) 
Nesom, comb. nov.—4:850 
Chionolaena cryptocephala 
Nesom, comb. nov.—4:850 
Chionolaena durangensis (Nesom) 
Nesom, comb. nov.—4:850 
Chionolaena macdonaldii 
Nesom, comb. nov.—4:850 
Chionolaena salicifolia (Bertol.) Nesom, 

comb. nov.—4:850 
Chrysogonum virginianum var. 
brevistolon Nesom, var. nov.—4:817 
Dichanthelium annulum (Ashe) R.J. 
LeBlond, comb. nov.—4:826 
Dichanthelium dichotomum (L.) Gould var. 


— 


Nesom) 


(Nesom) 


subtr. nov.— 265 

Asteraceae subtribe Boltoniinae Nesom, 
subtr. nov.—266 

Asteraceae subtribe Chaetopappinae 
Nesom, subtr. nov.—264 

Asteraceae subtribe Pentachaetinae 
Nesom, subtr. nov.— 264 

Barringtonia josephstaalensis Takeuchi, 
sp. nov.— 1:9 

Batopilasia byei Nesom & Noyes, comb. 
nov.—1:8] 

Batopilasia Nesom & Noyes, gen. nov.— 
1:8) 

Bolbitis thommankuthiana Nampy, sp. 
nov.—1:133 

Calycosia mamosei Takeuchi, sp. nov.— 


Canarium acutifolium (DC.) Merr. var. 
pioriverensis Takeuchi, var. nov.— 237 

Carex cumberlandensis Naczi, Kral, & 
Bryson, sp. NoV.—4:994 


SIDA 19(4): 1228. 2001 


nitidum (Lam.) R.J. LeBlond, comb. 
NOV.—4:829 

Dichanthelium dichotomum (L.) Gould var. 
ramulosum (Torr.) RJ. LeBlond, comb. 
nov.—4:830 

Dichanthelium dichotomum (L.) Gould var. 
roanokense (Ashe) RJ. LeBlond, comb. 
nov.— 4:83 ] 

Dichanthelium lucidum (Ashe) RJ. 
LeBlond, comb. nov.—4:831 

Dichanthelium sphagnicola (Nash) RJ. 
LeBlond, comb. nov.—4:834 

Digitaria pubiflora (Vasey) Wipff, comb. et 
Stat. Nov.—4:923 

Ditaxis humilis forma laevis (A. Gray ex 
Torrey) B.L. Turner, forma nova.—622 

Eleocharis moorei M. Strong, sp. nov.— 


Eragrostis ancashensis P.M. Peterson, 
Refulio & Tovar, sp. nov.—1:66 


NEW NAMES AND NEW COMBINATIONS 


Funastrum cynanchoides var. hartwegii 
(Vail) Krings, comb. nov.—1:137 

Hesperoyucca newberryi (McKelvey) Clary, 
comb. nov.— 4:845 

Hesperoyucca peninsularis (McKelvey) 
Clary, comb. nov.—4:845 

Houstonia acerosa (A. Gray) Bentham & 
Hooker f. var. polypremoides (A. Gray 
Terrell, comb. nov.—4:916 

Houstonia acerosa (A. Gray) Bentham & 
Hooker f. var. tamaulipana (B.L. Turner 
Terrell, comb. nov.—4:916 

Houstonia palmeri var. muzquizana (B.L. 
Turner) Terrell, comb. nov.—4:919 

Impatiens sholayarensis M. Kumar & 


Sequiera, sp. Nov.—4:795 


— 


~S 


Impatiens violacea M. Kumar & Sequiera, 
sp. NOV.—4:798 

lpomoea sororia D.F. Austin & J.L. Tapia, sp. 
nov.— 

Keysseria Lauterbach sect. Sandwicactis 
Nesom, sect. nov.—514 

Laennecia turnerorum Nesom, sp. nov.— 
4:/89 

Liatris aestivalis Nesom & O’Kennon, sp. 
nov.—4:768 

Liatris glandulosa Nesom & O’Kennon, sp. 
nov.—4:778 

Mirabilis laevis var. crassifolia (Choisy) 
Spellenb., comb. nov.—549 

Mirabilis laevis var. villosa (Kellogg) 
Spellenb., comb.nov.—551 

Orobanche ludoviciana Nutt. subsp. mul- 
tiflora (Nutt.) Collins ex H.L.White & W.C. 
Holmes, stat. nov.—623 

Paphiopedilum crossii (Morren) Braem & 
Senghas, comb. et stat. nov.—253 

Paphiopedilum crossii var. potentianum 
(Gru & R6th) Braem & Senghas, comb. 
et stat. nov.—254 


SIDA 19(4): 1229. 2001 


1229 


Paphiopedilum crossii var. sublaeve 
(Rchb.f) Braem & Senghas, comb. et stat. 
nov.—254 

Pennisetum pennisetiforme (Hochs. & 
Steud. ex Steud.) Wipff, comb. nov.—527 

Pennisetum setigerum (Vahl) Wipff, comb. 
nov.— 

Pennisetum somalensis (Clayton) Wipff, 
comb. nov.— 527 

Pittocaulon hintonii H. Rob. & Brettell var. 
cerrograndensis B.L. Clark, var. nov.— 
235 

Pittocaulon velatum (Greenm.) H. Rob. & 
Brettell var, lensis (T.M.Barkley) BL. 
Clark, comb. nov.—235 

Pseudognaphalium austrotexanum 
Nesom, sp. nov.—507 

Pseudognaphalium micradenium 
(Weatherby) Nesom, comb. et stat. 
nov.— 618 

Psychotria mayana Takeuchi, sp. nov.— 
1:17 

Pytinicarpa pickeringli (A. Gray) Nesom, 
comb. nov.—5 16 

Rhynchospora zacualtipanensis M. 
Strong, sp. nov.—1:115 

Saccoglossum takeuchii Howcroft, sp. 
nov.—519 

Senecio quaylei T.M.Barkley, sp. nov.— 286 


Solidago villosicarpa LeBlond, sp. nov.— 
292 

Stenaria (Raf.) Terrell, stat. nov.— 592 

Stenaria butterwickiae (Terrell) Terrell, 
comb. nov.—595 

Stenaria mullerae (Fosb.) Terrell, comb. 
nov.—597 

Stenaria mullerae var. pooleana (B.L. 
Turner) Terrell, comb. et stat. nov.— 598 

Stenaria nigricans (Lam.) Terrell, comb. 
nov.— 


1230 


Stenaria nigricans var. breviflora Terrell, var. 
nov.—605 

Stenaria nigricans var. floridana (Standl.) 
Terrell, comb. nov.—605 

Stenaria nigricans var. gypsophila (B.L. 
Turner) Terrell, comb. nov.—606 

Stenaria rupicola (Greenman) Terrell 
comb. nov.—608 

Stenaria umbratilis (B.L. Robinson) Terrell 
comb. nov.—610 

Stenaria umbratilis var. brevipedicellata 
Terrell, var. nov.—611 

Stenotis Terrell, gen. nov.—4:901 

Stenotis arenaria (Rose) 
comb.nov.—4:902 

Stenotis asperuloides (Benth.) Terrell, 
comb. nov.—4:903 

Stenotis asperuloides var. brandegeana 
(Rose) Terrell, Comb. nov.—4:904 

Stenotis australis (1M. Johnst.) Terrell, 
comb. nov.—4:906 

Stenotis brevipes (Rose) Terrell, comb. 
nov.—4:907 

Stenotis gracilenta (I.M. Johnst.) Terrell, 
comb. nov.—4:908 

Stenotis mucronata (Benth,) Terrell, comb. 
nov.—4:909 


Terrell, 


SIDA 19(4): 1230. 2001 


BRIT.ORG/SIDA 19(4) 


Stenotis peninsularis (Brandegee) Terrell, 
comb. nov.—4:910 

Stylogyne aguarunana Pipoly & Ricketson, 
Sp. NOV.—2:269 

Styrax platanifolius var. mollis (PW. Fritsch) 
B.L. Turner, comb. et stat. nov.—2:261 

Styrax platanifolius var. texanus (Cory) B.L. 
Turner, comb. et stat. nov.—2:261 

Styrax platanifolius var.youngiae (Cory) BL. 
Turner, comb. et stat. nov.—2:261 

Telanthophora cobanensis (J.M.Coult.) H. 
Rob. & Brettell var. molinae (H. Rob. & 
Brettell) B.L. Clark, comb. et stat. nov.— 


Telanthophora grandifolia (Less.) H. Rob. & 
Brettell var. serraquitchensis (Greenm.) 
B.L. Clark, comb. et stat. nov.—2:236 

Telanthophora sublaciniatus (Greenm.,) 
B.L. Clark, comb. et stat. nov.—2:236 

Trepadonia oppositifolia H. Rob. & H. 
Beltran, sp.nov.—1:111 

Tripogon ravianus Sunil & Pradeep, sp. 
nov.—4:803 

Vitis cinerea var. tomentosa (Planchon) 
Comeaux, comb. nov.—1:129 


GUIDELINES FOR CONTRIBUTORS 1231 


SIDA GUIDELINES FOR CONTRIBUTORS 


Sida, Contributions to Botany is an international journal of systematic botany 
containing primary research papers sensu lato, including anatomy, biogeogra- 
phy, chemotaxonomy, cladistics, ecology, floristics, genetics & evolution, nu- 
merical taxonomy, paleobotany, and palynology. Sida is open to all authors any- 
where; coverage is not restricted to any geographical area. 

All manuscripts submitted to Sida are considered by at least two review- 
ers. Manuscripts may be submitted in English or Spanish. Page costs are $35 per 
page but may be waived or reduced under certain circumstances. Please con- 
tact the editor and make arrangements before publication. Manuscripts are not 
rejected due to lack of financial support. Submit manuscripts and editorial ques- 
tions to Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan 
Street, Fort Worth, TEXAS 76102-4060, U.S.A. Phones: 1-817-332-7432 voice; 
1-817-332-4112 fax. Our electronic mail address is sida@brit.org. Access our Sida 
home page at the URL http://www. brit.org/sida/ 

Manuscripts must be double spaced throughout. Three hard copies and an 
electronic file of the manuscript are sent to the editor. Upon acceptance for pub- 
lication, please return two copies along with a copy in your word-processing 
program format (with format commands) on a 3'/2" data disk. You may also 
send your electronic copy attached to an email. Consult the latest issue of Sida 
for format of articles and notes regarding title, author, and address. Ranges of 
numbers are separated by double hyphens (6--8). Foot notes, figure legends, ap- 
pendices, and tables should be on separate pages and inserted at end of manu- 
script following references. 


ABSTRACT 
Every paper no matter the length should include both an English and Spanish, 
or another major language abstract. 
SPECIMEN CITATIONS 


Use the collector’s last name and number or (s.n.) and underline. Abbreviate 


the months of the year by the first three letters of the month without a period. 
Example: Texas. Jeff Davis Co. 1 mi W of Fort Davis, 3 Jan 1972, Smith 118 (BRIT). 


AUTHOR CITATIONS 


Use the author abbreviations as given in the Author Abbreviations compiled at 
the Herbarium, Royal Botanic Gardens, Kew (1992). Use et or the ampersand 
(&) between two authors; for more than two authors, restrict to the first one 


SIDA 19(4): 1231. 2001 


1232 BRIT.ORG/SIDA 19(4) 


followed by et al. Do not italicize or underline in the manuscript the terms et, et 
alex Or in: 

LATIN DESCRIPTIONS OR DIAGNOSES; ENGLISH DESCRIPTIONS 
Please provide a Latin diagnosis for each new taxon. After the Latin diagnosis, 
an English or Spanish version is Eecommenden highlighting the diagnostic char- 
acters. Write plant descripti with phrases separated 
by semicolons. 


I 


FIGURES 


Please present ALL figures asa unified, single sequence of figures. A scale should 
be in or on the illustration or photograph; reduction will automatically reduce 
the scale and subject proportionately. In halftones, sharp glossy photographs 
with good contrast are necessary for good reproduction. Please mount illustra- 
tive material (halftones, line drawings, etc.) on flexible paper so that may be 
mounted and on a high-speed drum scanner. You may also submit graphic 
materials in electronic format. Please consult the editor for proper graphics for- 
mat (tif, jpeg, gif, bmp, etc.). Color figures can be used and an additional cost 
maybe charged. Color figures reproduced best from a color transparency. Please 
submit color prints for reviewing purposes. 


REFERENCES 


This includes all of the literature cited in the text and may include other article 
citations the author may deem desirable. Normal text references should be cited 
as follows: ‘Ricketson and Pipoly (1997) stated... or ‘the latest revision (Ricketson 
S& Pipoly 1997) when reference is used as authority for a statement. When there 
are three or more authors use only the name of the first author followed by et 
al.: ‘Barrie et al. 1992a) stated’. References at the end of the article are arranged 
alphabetically and chronologically making use of a,b, etc. if an author had more 
than one publication in a given year. Author's names are typed in lower case 
except for initial capitals. Only the first letter of the initial word, proper nouns, 
and proper adjectives of titles should be capitalized. For journal abbreviations, 
use Botanico-Periodicum-Huntianum. Reference examples: 
Barrie, FR., C.E. Jarvis, and J.L. Reveal. 1992a.The need to change Article 8.3 of the Code. 
Taxon 41:508-512. 
Ricketson, J.and J. Pipoly. 1997. A synopsis of the genus Gentlea (Myrsinaceae) and a key 
to the genera of Myrsinaceae in Mesoamerica. Sida 17:697—-707 
Crawford, D.J. 1983. Phylogenetic and systematic inferences from electrophoretic stud- 
ies.In:S.D.Tanksley and T.J.Orton,eds.|sozymes in plant genetics and breeding, Part A. 
Elsevier, Amsterdam. Pp. 257-287. 


SIDA 19(4): 1232. 2001 


GUIDELINES FOR CONTRIBUTORS 1233 


Sivinski,R.C.and K. Lightfoot (eds.). 1995. Inventory of rare and endangered plants of New 
Mexico. New Mexico Forestry and Resources Conservation Division. Energy, Minerals 
and Natural Resources Department. Misc. Pub. No. 4. 


ABBREVIATIONS 
When the following abbreviations are used the period is omitted. Distance: mm, 
dm,cm,m, km, ft, mi; directions (in caps): S, N, E, W, months: first 3 letters only, 
Jan, Feb, etc. Example: Potter Co.: 2 km W of Dot, 5 Jun 1971, Smith 118 (SMU). 


DOCUMENTED PLANT CHROMOSOME NUMBERS 
Refer to Sida 18(3):909-925. 1999; 19(3):639. 2001. 


SIDA 19(4): 1233. 2001 


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