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-VoLuMe 19° Numer 1 ‘SEPTEMBER, 2000 oo
Barney L. Lipscomb, Editor
Botanical Research Institute of Texas
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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).
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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
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
Ac-
cessed 1999 September 30) and in the Texas A&M University Grass Images (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 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) .$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
FACULTAD DE CIENCIAS FORESTALES
UNIVERSIDAD AUTONOMA DE NUEVO LEON 0%
x 9 g
IV CONGRESO LATINOAMERICANO DE 3 og
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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:
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renee os mallee de Ciencias Forestales aed que se llevara a cabo en la Facultad
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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
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Presidente del Comité Organizador Comision de Contenido
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VOLUME 19, NUMBER 2, PAGES 235-443.
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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 ;
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)
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te
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‘ ‘ \
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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\ 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
sete 2000. Trees: Their Natural History. (ISBN 0-521-45963-X, pbk.) Cambridge
versity Press, 110 Midland Avenue, Port Chester, NY 10573-4930, U.S.A. $24.95
bk, 286 pp. Line drawings.
From the first sentence in Chapter One,“Everyone knows what a tree is: a large woody thing that
provides shade,” you know that the book you are about to read is written by someone who has
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 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....
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SHINNERS & MAHLER’S
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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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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
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
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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
; » - 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
» 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
BARBARA L. Bow inc. 2000. The Berry Grower’s Companion. (ISBN 0-88192-489-
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2. Berries in Mie Landscape
5. Blueberries
6. Grapes
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
reach type of berry plant, information includes aneete Sue notes on cultivation history, recom-
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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 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 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.
SIDA 19(3): 766. 2001
ms Mav en's
TRATED
Fon op
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 SHINNERS & MAHLER'S
projects! Use this one book instead of five, it has it all.” ILLUSTRATED
—Sharon Foster, 7th grade teacher,
Fort Worth Country Day School ELORA OF
“The best available, user-friendly source of information about NORTH CENTRAL
our native and naturalized plants. This book should be in the TEXAS
library of every gardener, landscape architect, horticulturist,
urban planner, farmer, rancher, and wildflower enthusiast.” Now IN ITS SECOND PRINTING !
—Howard Garrett, The Dirt Doctor DETAILED BROCHURE WITH ORDER FORM
AVAILABLE BY REQUEST ON THIS GROUND
BREAKING NEW FLORA.
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ALL
To be more specific: COMPREHENSIVE, ENCYCLOPEDIC, 817 332-4441 or
INNOVATIVE. With every species illustrated - it’s a new EMAIL: SIDA@BRIT.ORG
. FOR HOW TO ORDER.
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 RL
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“Si
ee
ISSN 0036-1488
At
Pt é
Z t ei; .
' ARL
Lift HOY AA, ¥ ARG! INIA
SOURCE OF
SIDA CONTRIBUTIONS TO
Botany was founded
by Lloyd H.
Shinners, above left
in 1962. Inherited
by William F.
Mahler, Director
Emeritus of BRIT,
above right, in
1971, and since
1993 it has been
published by BRIT.
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.orc
Home page at the URL: http://www.brit.org/sida/
John W. Thieret, Associate Editor
Dept. of Biological Sciences
Northern Kentucky University
Highland Heights, KY 41099 USA
Félix Llamas, Contributing Spanish Editor
Dpto. de Botanica, Facultad de Biologia
Universidad de Leon
E-2471 Leon, SPAIN
Guidelines for contributors are available upon request
and on our SIDA home page as well as the last page
of each issue.
Subscriptions for year 2001:
$27. Individual
$50. USA Institutions
$60. Outside USA
numbers issued twice a year
OSIDA, CONTRIBUTIONS TO BOTANY
VOLUME 19, NUMBER 4, PAGES 767-1233.
28 DECEMBER 2001
COPYRIGHT 2001
BOTANICAL RESEARCH INSTITUTE OF TEXAS
PRINTED IN THE UNITED STATES OF AMERICA
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.—
¥
oO
oD
Ww
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
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AGRA
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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 = —
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= ‘.
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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
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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)
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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
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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
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6 O
we O O 7 timida
ZO w+ 0 o “ s . d
se
= o- 5 2 1, a
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Co WW! 20 $0 a oa
= oO & O | iW
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= 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)
Book NOTICES
STUART GENTLING and Scott GENTLING. 2001. Of Birds and Texas. (ISBN 0-292-72834-
4, hbk.) University of Texas Press, PO. Box 7819, Austin, TX 78713-7819, US.A.
(Orders: http://www.utexas.edu/utpress, 512-471-4032). $75.00 hbk., 228
pp., 50 color plates, 28 color remarques, 8 figures, 9 7/8" x 12 7/8".
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essays detailing the history o I John James Audubon
to whom the folio edition was dedioated: In addition each plate oi oneal by a brief ee
of their own relationship to the subject of the painting. The volume was meant to read and so in-
cludes an essay, Self Portrait with Birds, by the nature writer, John Graves. But the paintings are Se
focus after all. Naturally there are color differences between the folio eee and this one, but the
accessibility of the book more than compensates for any loss. The emotional identification of the
artists with the landscapes and birds and the careful attention to detail make each painting memo-
rable.
The book is an exceptional tribute to Audubon. Pl Ginsburg, Botanical Research Institute of
Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, |
Editor’s not If Birds and Texas.—Of the 50 ee lates 10 are of landscapes of Texas and the
remaining 40 are of birds. ager the Gentlings have painted more than just birds, there are at
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flower, us: and tl their i tae ular foliage include, Water Hyacinth, Yellow Lotus, White Water-L ily,
Sweetgum, é ; ir Elm, Common Trumpet-Cree per, Texas Bluebonnet, Sycamore. Don't
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“God hath made three beautiful things, birds, and women, and flowers,’—John Sturat Blackie,
[he Botanist’s song, 1869. This is almost the perfect book.
CHRISOPHER KeLty (Editor). 2000. Jean-Jacques Rousseau The Reveries of the Solitary
Walker, Botanical Writings, and Letter to Franquieres. Translated and
annotated by Charles E. Butterworth, Alexandra Cook,and Terence E. Marshall.
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In the middle of the 20" century, the taxonomist, Lloyd H. Shinners, wrote that he was testing the
validity and hope of the most romantic of sciences in an age hostile and alien to it. The writings of
ean-Jacques Rousseau, the ‘Father of the Romantic Movement offer an interesting perspective on
that earlier time when the science of botany was emerging. Jean-Jacques Rousseau offers an admi-
rable translation of the nature essays and letters about botany, and thus makes available to the En-
glish reader a lesser known side of the great philosopher. The copious and well-written notes pro-
vide a valuable backgrounc
to the circumstances of the walks as well as a short history of botany.
Rousseau'’s deep appreciation of the beauties of nature and the lyricism of his writing are well served
by this excellent book.—Ruth Gins
Worth, TX 76102-4060, U.S.A.
ourg, Botanical Research Institute of Texas, 509 Pecan Street, Fort
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
4)
(
BRIT.ORG/SIDA 19
946
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J ens x
oe
MOIBIOQE |
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University of California Riverside
Herbarium
Amaranthus cruentes L
Det. by Mihai Costes
CULTIVATED PLANTS
-OF-THE
Ue SHNAHONAL ARBORETUM, WASHINGTON,
Dc
from Lehmann, AMES 5602 (NA)
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
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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)
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= J SJ Ww i CASTER |OTOE
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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!
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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
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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
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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 . 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)
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
ESPECIES 0 R | G E
NT NA CA CB SA AT EU EA AS AF OC
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
Ss)
2 =
S
OE ses
Ba*s
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
CA CB SA AT EU EA AS AF OC
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
NT NA CA CB SA AT EU EA AS AF OC
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
ESPECIES 0 R I
NT NA CA CB SA AT EU EA AS AF OC
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
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relevance of shamanism to West mallangles
and ee ines. From the back cover: “The shaman occupies a key role as a healer, mediating be-
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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,
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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,
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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
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Lewis (1774-18 Portulacaceae.“A genus of succulent asta
numbering appr vosimae Ean speties Lewisia has been adopted asa favorite by rock gardener
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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 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
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erie
DE AGRICULTURA
Lime - rexu
RBARIO NACIONAL
te Breast,
OME sncreemcee ov +
TLE,
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Localidad LA yl {}
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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
Subscriptions and distribution.—Swa is distributed in over 90 countries. There
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We thank all authors, reviewers, subscribers, and readers for your contin-
ued interest and support.—Barney Lipscomb (BRIT), Editor; John W. Thieret
(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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