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509 Pecan Street 

Fort Worthjexas 76102-4060,1 

riieret, Associate E 

Felix Llamas, Contributing Spanish Editor 

Dpto, cle Botanica, Facultad de Biologia 

©sida, contribuiions to botany 

Volume 20, Numbfr 1 , pagfs 1 -460. 

19 July 2002 

Copyright 2002 

Botanical Research Institute of Texas 

Printed in the United States of America 

ISSN 0036-1488 

Table of Contents 

Key to the species of Ericaceae of Be 

W.Takeuchi and M.Golman — 55 



Richard Spellenberg— 151 
BiLLiE L.Turner— 157 


Documented chromosome numbers 2002: 1 . Chromosome number of Stenanthium (L 
Wendy B. Zomlefer and Gerald L. Smith— 221 

J.K. Morton— 227 

Alina Freire Fierro, Diana FernAndez, and Catalina Quintana— 233 


Vascular plants of Konza Prairie Biological Station: an annotated checklist of speci 

\ Sierra "El Viejo," Sonora, reg oi 
S— 349 

Eric L Keith and N. Ross Carrie— 387 

Robert F.C. Naczi, Ronald L. Jones, F, Joseph Metzmeier, Mark A. Gorton, and Timothy J. Weckman— 397 

Eric Sundell, R. Dale Thomas, Carl Amason, and Chris Doffl 

F Calcasieu Parish, L 

Rupert Barnfby Award— 446 

Book reviews and notices 30, 46, 1 04, 1 1 4, 1 48, 1 56, 1 60, 1 78, 1 88, 232, 268, 312, 348, 372, 396, ^ 





STAT NOV — 170 




NOV — 170 




Lelong ( 

-ombnov— 170 


"sTATNOV — 171 


3, COMB.. 

& STAT NOV.— 171 


NG, comb. NOV.— 171 

Eriogonum riogran 

disNesom,sp.nov,— 32 

Festuca cuzcoensis 

5tanc!ik& P.M. Peterson, SRN. 

ov.— 22 


5rAN(riK & PM. Peterson, sr nov.— 24 


Ipomoea costellata 



Medinilla anamalai. 


5R NOV.— 110 


Panicum sect. Hemitc 

)maH(tchc.& Chase EX Freo 

;MANN &LELONG, sect NOV.- 


Panicum AMARUM SUf 


AT NOV — 171 

Panicum anceps subs 




EAT NOV — 171 

Panicum capillars si 

jbsrhillmanii (Chase) FRECKf 

. NOV.—' 


Panicum dichotomifi 




Panicum dichotomifi 





?w York Botanical Gard 
lew York 10458-51 26, U 

Luteyn y Themistocksia unduavensn Luteyn. Se presenta una clave en ingles y en espanol para los 19 
generos y las 53 especies nativas. Los nombres de todos los taxones de Ericaceae atribuidos a Bolivia 

The Ericaceae are a large, cosmopolitan family of over 125 genera and 4500 spe- 
cies, which inhabit the temperate regions of the world and montane areas in 
tropical latitudes (Luteyn 1998, m press). In the Neotropics, the Ericaceae are 
composed of 46 genera (70% endemic) and about 800 species (ca. 94% endemic), 
although generic limits m the Andean Vaccinieae are still poorly understood 
(Kron et al. 2002; Luteyn 1997, 2001). Neotropical Ericaceae are an Andean-cen- 
tered family, adapted to moist, open, cool montane environments. Overall spe- 
cies richness in the Neotropics increases nearer the Equator, with the highest 
species numbers concentrated in Colombia and Ecuador between 1000 m and 
3000 m. Nineteen genera and 53 species are native to Bolivia, Two of those gen- 
era and 18 of the species are endemic, while 18 additional species have ranges 
that overlap between Bolivia and adjacent Peru and one species range overlaps 
between BoLvia and extreme northwestern Argentina; one species is occasion- 
ally cultivated. In Bolivia, the Ericaceae occur almost exclusivelv in the wet 

cool, montane cloud forests, mostly in the " Yungas" regions of the eastern slopes 
of the Cordillera Real, ranging between 2000 m and 4000 m elevation. One of 
these species (Satyria negkcta) ranges primarily below 1000 m elevation, 12 
species range primarily between 1000 m and 2000 m, 18 species range prima- 
rily between 2000 m and 3000 m, and 20 species range primarily between 3000 
m and 4000 m. Agarista holiviensis and Gaylussacia cardenasii, in contrast, 
occur in relatively dry montane regions in south-central Bolivia at about 1200- 
2500 m elevation. Although Bolivia is a very large country within the Neotropics, 
it lies at the southern end of the distributional range of the family and many of 
Its mountainous regions are dry Therefore, relatively few Ericaceae occur in 
Bolivia compared to countries in the Northern Andes and adjacent southern 
Mesoamerica. There are no economic uses known for the Ericaceae of Bohvia. 

There are very few general publications about the flora of Bolivia and only 
the out-dated checklist of Foster (1958) gives an idea of the numbers of species 
of Ericaceae that occur there. Killeen et al. (1993) is misleading for Ericaceae, 
since virtually all the taxa in Bolivia are shrubs not trees. There is, however, a 
project currently being organized by the Missouri Botanical Garden in collabo- 
ration with all the major herbaria in Bohvia and the New York Botanical Gar- 
den that has begun a modern checkhst of the plants of Bolivia. With regards to 
the Ericaceae of Bolivia, only the larger overall treatment of the tribe 
Thibaudieae (=Vaccinieae) by A.C. Smith (1932) gives details of some of the 
Bolivian blueberries. The most up-to-date list of Ericaceae of Bolivia, along with 
some descriptions and photos, may be found on the website "Neotropical Blue- 
berries" (Luteyn 1998). A list of all species of Ericaceae attributed to Bolivia, 
including the current status of names given by Foster (1958), is herein presented 
in an Appendix. The largest herbarium collection of Ericaceae in Bolivia is found 
in La Paz, at the National Herbarium (LPB); there is also a moderate sampling 
in Cochabamba (BOLV). The geographical regions of Bolivia in greatest need of 
more collecting and with greatest possibilities for more Ericaceae records would 
be the areas northeast of La Paz towards the frontier with Peru. 

Neotropical Ericaceae, in general, have undergone dynamic speciation and 
extensive adaptive radiation due to their ecological and life-form plasticity colo- 
nization abilities, adaptation to epiphytic habits, and co-evolution with hum- 
mingbirds (Luteyn 2002). Given the high diversity and endemicity within 
neotropical Ericaceae, along with high levels of habitat alteration, protection of 
Andean montane ecosystems should become a priority for their c 


Terrestrial or epiphytic shrubs, suhshruhs, sometimes lianoid, rarely trees, often 
rhizomatous; indumentum when present of simple unicellular hairs, that are 
usually translucent or whitish, these sometimes glandular tipped, or often (es- 

pecially on leaves beneath) of multicellular, multiseriate, swollen-headed, usu- 
ally glandular, hair-like structures called fimbriae which are brownish to red- 
dish-brown. Leaves alternate, simple, usually petiolate, exstipulate but rarely 
bud scales appearing pseudostipular; lamina coriaceous to membranous, ever- 
green, the margin usually entire but sometimes serrulate-crenate, the venation 
pinnate or plinerved; leaf scars usually with a single vascular bundle scar, nodes 
usually with one trace and one gap. Inflorescences axillary racemose, panicu- 
late, fasciculate, or flowers solitary; individual flowers pedicellate or rarely 
sessile in axils of small or large, deciduous or persistent floral bracts; pedicel 
bibracteolate; bracteoles persistent, small or large. Flowers bisexual, actinomor- 
phic or slightly zygomorphic, 5(-7)-merous, obdiplostemonous, hypogynous 
or epigynous and with a biseriate perianth, typically without floral odors, rarely 
with extraf loral nectaries, the superior-ovaried genera pollinated by bees and 
the inferior-ovaried genera by hummingbirds; aestivation valvate, imbricate, 
or reduplicate; calyx continuous or articulate with the pedicel, synsepalous, the 
sepals rarely fleshy and accrescent to the fruit, the hypanthium when present 
terete, angled, or winged; corolla membranous to thick-carnose, polypetalous 
or more commonly sympetalous, cylindric, campanulate or urceolate, terete or 
angled to winged opposite the lobes; stamens 8-10(-14), in 2 whorls, usually 
twice as many as the petals or rarely just as many, equaling the corolla in over- 
all length or 1/2-1/3 the corolla length, equal with each other or alternately 
unequal, borne on the edge of an obscure to prominent nectariferous disc; fila- 
ments equal or unequal, usually straight or rarely S-shaped (geniculate), ligu- 
late but sometimes basally dilated, sometimes also basally papillose, distinct 
or connate, with or without spurs, shorter or longer than the anther; anthers 
inverting during development, 2-celled, equal or unequal, often distally with 2 
distinct or connate tubules or terminal awns, sometimes provided with abaxial 
spurs; disintegration tissue present or lacking; thecae smooth to coarsely granu- 
lar, the base rounded to apendiculate; tubules when present conical and rigid 
or cylindric and flexible, of equal or ca. 1/2 the diameter of the thecae, longer to 
shorter than the thecae; dehiscence introrse by longitudinal or more typically 
by apical to subapical clefts or pores, rarely perfectly terminal; pollen grains in 
tetrahedral tetrads; carpels 4-5(-10), fused; ovary superior or inferior, usually 
with as many locules as carpels or with twice as many locules as carpels or 
rarely loculate m lower portion and Mocular above; placentation axile, rarely 
intruded parietal; ovules numerous per locule or rarely solitary, anatropous to 
campylotropous with a single integumentary layer; style single, fluted, hollow; 
stigma simple but occasionally weakly lobed. Fruit a loculicidal or septicidal 
capsule, berry, or drupe, with a usually persistent, rarely accrescent and fleshy 
calyx; seeds small, ca. 1-1.5 mm long, usually numerous (1 per locule in 
Gaylussacia), winged or tailed only in Bejaria, sometimes enclosed in a muci- 
laginous sheath, the testa thm with elongated or isodiametric cells, the en- 

dosperm fleshy, the embryo straight, usually white or sometimes green. Chro- 
mosome numbers: x=6, 8, 11, 12(?), 13, 19, 23. 

Ericaceae comprise ca. 125 genera and 4500 species worldwide, and are 
cosmopolitan with the exception of Antarctica. Nineteen genera and 53 spe- 
cies are native to Bolivia. Rhododendron simsh, native to China, is sometimes 
cultivated in the montane areas throughout the Neotropics (including Bolivia), 
but is not treated here. 

Notes on using the keys 

The keys below are based primarily on herbarium specimens, although living 
{in situ and greenhouse-grown plants), alcohol-preserved material, and photo- 
graphs taken from field or greenhouse-grown material have been used. Floral 
measurements are taken from herbarium material at anthesis unless otherwise 
stated; colors are from fresh material observed by the author unless otherwise 
stated; if a range of measurements is not available, the known measurement is 
preceded by the abbreviation "ca." (about); calyx limb length includes measure- 
ment of the lobes, and anther length includes thecae and tubules. 


. Fruit a capsule, surrounded by the fleshy, accrescent calyx 
4. Flowers solitary in axils of normal (or only slightly reduce 

_Gaultheria buxifolia 

Gaultheria vacinioides 

< glabrous; corolla glabrou: 

subsetose, rarely glandular hairs;ovary densely short-white pilose 
or cinerous; inflorescence not congested at anthesis, the flowers 
widely spaced Gaultheria bracteata 

. Young twigs and inflorescences glabrous or variously spreading pu- 

obovate with the apex rounded _ 
. Erect, subshrubs to shrubs, 0.1-8 

veined on both surfaces, the base usually acute to rounded; 
inflorescences clustered at branch tips and conspicuously ex- 
ceeding the leaves in length;calyx and corolla eglandular;co- 
rolla white, pilose within Gaultheria n 


2. Filaments equal and connate over entire length;anthers with tubules widening 

3. Leaves elliptic-lanceolate, 6-10 cm long, 1.5-3 cm broad, basally obtuse to 
rounded, 3-plinerved; inflorescence densely pilose (except corolla); pedicels 
16-21 mm long; corolla ca. 1 1 mm long Satyi 

:-oblong, 3-6 cm long, 1.2-1.8 cm broad, basally c 

lants essentially glabrous. 

. Corolla 15-33 mm long. 
8. Staminal filaments distinct. 

Cavendishia pubescens 

ted; leaves glabrous to glabrate Cavendishia b 

. Stamens equal with filaments and anthers of equal lengths (rarely anthers incon- 
spicuously alternately unequal). 
1 3. Bracteoles located at apex of pedicel and surrounding calyx (and sometimes 

Disterigma pernettyoide; 

. Corolla subcylindric to somewhat urceolate, 6-9 mm long, red Disterigma 

aves more than 5 mm broad. 
. Corolla 10-12 mm long, narrowly cylindric, red Disterigma pallidum 

Tubules elongate,thin, very graceful. 

prominent,3-5 in number,often ir 

corolla 1 8-35 mm long; stamens 9 

22. Leaves chartaceous to soft coriae 

24. Corolla elongate-tLibular,(7-)l 

ous with regard to anthers. 
25. Corolla 20-27 mm long;s 

a macrocalyx 

'; calyx 5.5-6 mm long, the h 

apiculate, less than 1 mm long Thibaudia crenulata 

24. Corolla urceolate to campanulate, 7-12 mm long, normally of 

ments proportionally long with regards to anther length. 
29. Ovary with a single ovule in each of the 10 locules;fruit a 

drupe with 10 pyrenes Gaylussacia cardenasii 

30. Leaves glabrous or lacking subfasciculate hairs be- 

_ Themistoclesia 

)-angled,8-11 mm long, 

1 or more long, if less than 1 cm then staminal 
ice as long as thecae;staminal tubules 2-5 time^ 

. Corolla 0.6-0.8 cm long, white to pinkish 

37. Corolla 2-3 cm long and 3-8 mm diam., slightly 

zygomorphic Demosthenesia mandonii 

nomorphic Demosthenesia spectabilis 

irollas small, thin-membranaceous, up to 10 mm long, 
t if longer then filaments proportionally much longer 

. Flowers usually in few- to many-flowered fascicles or 

39. Inflorescence a 

sessile); filaments usually longer than anthers. 

40. Flowers sessile Sphyrospermum sessiliflorum 

40. Flowers conspicuously pedicellate. 

0.9-1 .5(-l .8) cm long, the apex rounded or ob- 

incipalmente en ejemplaresde herbario, aunque 
le invernadero), material 
campo y en invernaderos. 
Las medidas f lorales se tomaron del material de herbario en la antesis a menos 
que se diga lo contrario; los colores se tomaron del material vivo observados 
por el autor a menos que se diga lo contrario; si no se tiene un interval© de 

medidas, entonces las medidas conocidas se presentan precedidas por la 
abreviacion "ca." (cerca); la longitud del limbo del caliz incluye la medida de los 
lobulos y la longitud de la antera incluye las tecas y los tubulos. 


L Corola con petalos separados;fruto una capsula septicida Bejaria aestuans 

L Corola con petalos unidos;fruto una capsula loculicida o una baya. 

2. Estambres con filamentos geniculados.anteras sin aristas terminales y sin tejido 

bianco desintegrado en el lado abaxial Agarista boliviensis 

2. Estambres con filamentos rectos; anteras con aristas terminales y con tejido 

3. Fruto una baya, raras veces el caliz se vuelve carnoso en la base pero nunca 

. Flores solitanas en las axilas de las hojas normales (o ligeramente reducidas), 

ancha en el apice,verde-amarillenta Gaultheria buxifolia var.secunda 

5. Subarbusto de tallo delgado hasta 0,4 (raras veces 1-2) m de altura; 
corola urceolada a estrecfiamente subglobosa,infladaen la basey muy 

contraida en la garganta, rosada a rosado-roja Gaultheria vaccinioides 

. Flores en racimosaxilares. 

aguda a redondeada; i 


-lanceoladas, 6-1 cm de largo, 1 ,5-3 cm de ancho, obtusas a 
basalmente, 3-plinervias; inflorescencia densamente pilosa 

Satyria boliviana 

las basalmente, pinnatinervias; inflorescencia glabra; pedicelos 6- 
3rgo;corola ca.6 mm de largo Satyria negle< 

. Raquis 3-5 cm de largo; pedicelos 8-13 mm de largo Orthaea 

0. Dehiscencia estaminal por poros terminales o subterminales Orthaea 

0. Dehiscencia estaminal por suturas laterales Orthaea ferreyrae 

la parte baja de la corola; 

. Corola campanulado-c 

15. Corola subcili'ndrica a a 

estaminales cohesionados en la base Disterigma ovatum 

)s alargados,delgados,casi tan ancho como la mitad (o menos) de 

20. Corola ca. 25 mm de largo _ 

20. Corola 43-48 mm de largo 

Tubulos casi tan anchos como la 

proporcionalmente mucho mas cort 

s que la teca;dehiscencia por suturas 


21 . Teca conspicuamente papilosa;tu 

bulos rigidos.alargados a cortamente 

c6nicos;estambres a menudo 1/3-1/2 tan largos como la corola 

22. Hojas rigidamente coriaceas 

pedicelos 20-40(-55) mm de largo; 


,3-5,a menudo algo fusionados, 1-3 

Psammisia paucifloi 

5;estambres generalmente tan largos como la corola. 

:uos con respecto a las anteras. 

en la base Thibaudia axillaris 

superficies de la planta glabras o esencialmente glabras, 

los lobulos 1 mm de largo Thibaudia 

Thibaudia crenulata 

:ampanulada,verde Vaccinium dependens 

H pedicelo, rio articulado. 

1 Demosthenesia 

?zigomorfa Demosthenesia mandonii 

Demosthenesia spectabilis 

Tiembranacea, h 

39. Inflorescenciaenfascicu 

39. Inflorescencia en racim 

raquis 2,5-6 cm de large 

. Floresl-2 

1 ,5(-l ,8) cm de largo,el apice redondeado 
tuso;flores generalmente extendiendose 

rgo;estambres4 Sphyrospermum buxifolium 


Siphonandra boliviana Luteyn, sp. nov. (Fig. l). TYPE: BOLIVIA. Depto, LA Paz. Prov. 
Bautista Saavedra: Charazani, W of Chullina, 3400 m, 1 Aug 1994 (fl), B. Herzog 

Shrub (size unknown); mature branches terete, glabrous, the bark exfohating 
m thin strips; twigs subterete, sometimes shallowly angled or ribbed, short- 
pilose with white hairs; buds axillary, the scales 2, valvate, ca. 2 mm long, short- 
pilose. Leaves alternate, coriaceous, elliptic to oblanceolate, 2.5-5.5 cm long, 1-1.8 
cm broad, basally cuneate, apically broadly acute to nearly obtuse, marginally 
entire, essentially glabrous above or sparsely short-pilose proximally along 
midrib, sparsely pilose beneath especially along midrib, also provided with 
reddish-brown, basally swollen, glandular fimbriae beneath; pmnately nerved 
with 4-6 secondary veins anastomosing near margin, the midrib and second- 
ary veins impressed above and raised beneath, the reticulate veins plane to 
slightly impressed above but inconspicuously raised beneath; petiole rugose, 
subterete, broadly flattened above, 4-5 mm long. Inflorescence axillary, race- 
mose, ca. 20-flowered, apparently nodding; rachis subterete, striate to angled, 
densely short-pilose with white hairs, at least 5 cm long (still m bud, apparantly 
still elongating); floral bracts ovate, acuminate, 3-4 mm long, densely short- 
pilose; pedicels subterete, striate to angled, densely short-pilose as rachis, 11-13 
mm long; bracteoles located m proximal 1/3 pedicel, similar to floral bracts, 2- 
3 mm long. Flowers: calyx articulate with pedicel, 7-8 mm long, densely short- 
pilose as rachis; hypanthium cylindric, ribbed, 4-5 mm long, rounded at base; 
limb spreadmg-campanulate, 3.8-4.5 mm long; lobes 5, deltate, acute, 1.5-2 mm 
long; sinuses obtuse; corolla of 5 fused petals, long-cylindnc, 43-48 mm long, 
6-7 mm diam., short-pilose throughout, with white hairs, the lobes 5, deltate, 
acute, ca. 2 mm long; stamens 10, equal, ca. 32 mm long; filaments connate, gla- 
brous, ca. 7 mm long; anthers ca. 28 mm long; thecae granular, ca. 6 mm long, 
incurved at base; tubules thin, delicate, about half as wide as thecae, ca. 22 mm 
long, dehiscing by perfectly terminal, flaring pores; ovary 5-locular; style about 
equaling corolla. Berry not seen. 

Distribution.-Endemic to Bolivia and known only from the type collection, 
which was made in a Weinmannia forest. 

Etyn^ology-The species is named for the country Bolivia to which it is endemic. 

Siphonandra is a small, distinctive, high-elevation genus of three species: 
5. elliptica which is common and ranges from south-central Peru to northern 
Bolivia, S. magnijica which is endemic to Bolivia and is herewith maintained 
despite the fact that the type and only specimen was destroyed during World 
War II, and the new species herein described 5. holiviana, for which only the 
type collection is known. With only one extant collection of the latter two species, 
it is difficult to assess the relationships between any of the three species in this 
genus. Nevertheless, S. holiviana is easily distinguished morphologically from 
the other two species by the characters mentioned in the key and diagnosis. 


Themistoclesia unduavensis Luteyn, sp. nov. (Fig. 2). Type: BOLIVIA. Depto. La Paz. 
Prov. Nor Yungas: trail to Rio Coscapa, ca. 2.5 km E of Unduavi, along new rd to 
Coroico, and ca. 6 km W of Cotapata, ca. 16' 17'S, 67" 53'W, 3200-3350 m 19 Mar 
2000 (fDj.L. Luteyn, E. Ann Powell & S. Beck 15m (holotype: NY; isotypfs AAU 
CAS, F, K, LPB, MO, TEX, plus 6 others to be distributed by LPB). 

;, terrestrial or epiphytic subshruhs, to 30 cm tall; 
, glabrous, the bark cracking longitudinally but m 

s G. Stamens showing back, f 

t, and side views (kUom Solomon U 

twigs subterete, flattened to broadly and obtusely ribbed, weakly striate, gla- 
brous; buds axillary, the scales 2, valvate, ovate, acumuiate to acicular, 4-5 mm 
long. Leaves alternate, coriaceous, slightly bullate, clustered near apex, ovate to 
elliptic-ovate, 4.4-9 cm long, 1.4-4.5 cm broad, basally rounded to obtuse, 
apically long-acuminate, marginally entire, glabrous on both surfaces but pro- 
vided with brownish, glandular fimbriae beneath; 3-5-plmerved with inner 
nerves arising 7-11 mm above base, midrib, lateral nerves, and reticulate vem- 
lets impressed above and raised beneath; petioles rugose, subterete, broadly flat- 
tened above, 3.5-5 mm long, glabrate. Inflorescences axillary but usually from 
older, leafless nodes, flowers l-2(-3) per node, when 2-3 then with very short 

rachis; rachis (when present) subterete, 1-4 mm long, glabrous, obscure; floral 
bract ovate, acute, somewhat cucullate, ca. 2 mm long, marginally glandular- 
fimbriate, glabrous; pedicels slender, subterete, striate, sharply angled distally 
14-18 mm long, glabrous; bracteoles nearly basal, ovate, acuminate ca. 3 mm 
long, glabrous but marginally glandular-fimbriate. Flowers: calyx contmuous 
with pedicel, 6-8.5 mm long, essentially glabrous; hypanthium obpyramidal, 
strongly 5-winged, 3.5-5 mm long, sparsely short-pilose along proximal por- 
tions of wings; limb spreading, ca. 3-3.5 mm long; lobes 5, broadly ovate, sharply 
and shortly acuminate, ca. 2-2.8 mm long, sparsely short-pilose along margins; 
sinuses acute; corolla of 5 fused petals, carnose, urceolate-turbinate, broadly 
swollen basally broadly and bluntly 5-angled, 8-11 mm long, 7-12 mm diam. 
at base, glabrous, reddish-orange, the lobes 5, ref lexed, deltate, acute, ca. 1.5 mm 
long, green in bud but becoming light green to white at anthesis; stamens 10, 
equal, nearly as long as corolla, ca. 8 m m long; filaments distinct, sparsely short- 
pilose, ca. 2.5-3 mm long; anthers ca. 7~7.5 mm long; thecae smooth, incurved 
at base, ca. 3.5 mm long; tubules distinct to base, ca. 3.5-4 mm long; ovary 5- 
locular; style about equaling corolla. Berry not seen. 

Distribution.-Endemic to Bolivia and known only from the province of 
Nor Yungas, between Unduavi and Cotapata, at ca. 3000-3500 m. The plants 
are found in the wet, moss-covered, cloud forest associated with Clusia, 
Weinmannia, Hedyosmum, Brunellia, Miconia, and Desfontainea. 

Etymology -The species is named for the small village of Unduavi, close to 
the locality from which all collections have been made. 

Themistodesia is a neotropical genus of ca. 25 species, ranging from Costa 
Rica and Panama through the Andes of South America from Venezuela into 
northern Bolivia. Themistodesia unduavensis may be distinguished by its rhi- 
zomatous, subshrub habit, somewhat buUate leaves, 1-3-f lowered, axillary and 
often ramif lorous inflorescences, carnose corollas that are bluntly 5-angled and 
basally swollen, and reddish-orange corollas with green to white lobes. Indi- 
vidually each of these characters is either absent or uncommon amongst the 
other species in the genus, and the com bmation of characters effectively isolate 
this species from all others. 

appearing in it 

synonyms; names appearing in Roman are currently accepted; ! = new specie: 
BO = species endemic to Bolivia (or nearly so); * = species endemic to norther 
Bolivia and adjacent central to southern Peru; ** = species not found m Bolivi: 
CULT = introduced ornamental species.] 
Agarista boliviensis (Sleumer) Judd BO (but see (type and only specimen destroyed durin 

Befaria boliviensisB.f edtscl 

n.&Basilevsk.= Bejana 

uemostnenesia graeonena 

Befaria denticulata Remy = Bejaria aestuans 
Befariaglauca Bonpl.= Bejaria aestuans 
Befaria glauca var. coarctata (Bonpl.) Mansf & 

Befaria glauca var. glandulosa Mansf. & Sleumer 

Diogenesia racemosa (He 

Befaria glauca var. setoss 
Bejaria aestuans 

Befaria glauca var. tomeril 
= Bejaria aestuans 

Befaria hispida Poeppi 


Disterigma alaternoides ((■ 
Distengma alaternoides va 
A.C.Sm.= Distengmac 
Disterigma empetrifolium 
Disterigma ovatum(Rusb: 
Disterigma pallidum A.C.S 

3, only in Colombia 
loides Rusby = G 

Cavendishia strobilifera (Kunth) Hoer. - 
Ceratosfema hoo/cen'Britton = Siphonandra elliptica 

Gaultheriaodorata Bredem 
Gaultheria pichinchensis 

Gaylussacia pseudogaulthena Cham 

Orthaea boliviensis B.Fedtsch.& Basilevsk.B( 


Orthaea ferreyrae A.C.Sm.* 

Orthaea ignea Sleumer * 

Orthaea pinnatinervia Mansf.* 

Orthaea rusbyi Luteyn BO 

Orthaea weberbaueri Hoerold * 

Pernertya densa Rusby = Pernettya prostrata 

Pernettya phyllyraefolia (Pers.) DC. =not fou 

in Bolivia, only Argentina and Chile 
Pernettya prostrata (Cav.) DC. 
Pernettya prostrata var.pentlandii (DC.) Sleun 

G.Don) Sleumer = Pernettya prostrata 

sufficient for determii 
obusta Rusby = Sphyrc 

ccir^ium floribundurv 

var. ramosissimi 

(Dunal) Sleumer = Vac 


ccinium floribundum var. 

fore/ (Rusby) Sleun- 



Dunal =VacciniL 


Stephan Beck encouraged me to write this paper I thank the Herbario LPB (La 
Paz, BoHviaXjim Solomon, the Missouri Botanical Garden, and the late David N. 
Smith for facilities and logistical support. Thanks and acknowledgments are 
also given to thejessie Smith Noyes Foundation and the National Science Foun- 
dation under whose grants field work was undertaken and this paper was writ- 
ten. 1 thank Stella Sylva for translating the key into Spanish and computer help, 
Bobbi Angell for the beautiful illustrations, and Carmen Ulloa U. and Walter 

Foster, R.C.I 958. A cal 




plants of Bolivia.C 


184:1 -223. [Ericao 




RMANN. 1990.( 



]ansta boliviensis ( 

Ericaceae). Sida 


KiLLEEN.TJ., E. Garcia E.„ 

and S.G. Beck ( 



Doles de Bolivia. He 


de Bolivia and Mis 

souri Botanic 

:al Garden, La Pa. 

St Louis. [Ericacea 


Kron,K.A., E.A.Powell, 





n the blueberry 

tribe (Vaccinieae,i 

Ericaceae) ba 

sed on sequenc 


:a from mafK and 

nuclear riboso- 

mal ITS regions, wi 

th comment 




LuTEYN, J.L. 1995. Gaultheria. Pp. 

384-488. In: J.L 

.. Lut 

eyn, ed. Ericacea 

e-Part II. The 


genera (Monotropoideae 

', Pyroloideae, Rhododendroideae, 

Vaccinioideae p.p. 

). PL Neotrop. Monogr, 67:1 -560. 

LuTEYN, J.L. 1997. Are 


axonomic realignments within the nee 

^tropical genus 

Madeania (Ericac^ 




Esp. No. 6:455-46! 


tropical btuel 

3erries:The plant 


ily Ericaceae, www 



LuTEYN, J.L. 2001. Two r 

lew species a 

ind two new con 



rican Ericaceae. 

Brittonia 53:437-4 


LuTEYN, J.L. 2002. Dive 


tion,and endem 


in neotropical Ericaceae: Biogeo- 

graphical patterns 

■ intheVaccir 





eds. Plant evolutic 


nism in Andean 


h America. BoL Re 

V. 68:55-87. 

LuTEYN, J.L (In press). 1 

Ericaceae (He 

^ath Family), In: ^ 



Henderson, S.A. 

Mori and D.W.Ste\ 




n Tropics. Princ- 

eton University Pr 

ess/New Yorl 

(Botanical Card, 


ess, Princeton, NJ and Bronx, NY. 

Smith, A.C.1932.The/ 





rb. 28:3 11 -547. 


Daniel Stancik 

Paul M.Peterson 



tment of Systematic Biology - fit 
tional Museum of Natural Histo 

Smittisonian Institution 
ashington. DC 20560-0166, USj 




M.Peterson a 

nd E tovflrensis Stancik & P.MT 

While making determinations of South American material in the United States 
National Herbarium the senior author recognized several specimens with 
unique features. The two new species discussed here appear to be members of 
Festuca subg. Suhulatae (Tzvelev) E.B. Alexeev sect. Suhulatae. The position of 
sect. Suhulatae in subg. Suhulatae is clear, however, the relationships with other 
sections in this subgenus are unresolved. Alexeev (1980, 1982, 1986) recognized 
three sections in subg. Suhulatae: sect. Suhulatae, sect. Elmera E.B. Alexeev, and 
sect. Glahricarpae E.B. Alexeev Alexeev (1986) and Clayton and Renvoize (1986) 
also recognized two other subgenera in these flat-bladed South American spe- 
cies of Festuca: subg. Suhuliflorae E.B. Alexeev and subg. Ohtusae E.B. Alexeev 
Aiken's et al. (1997) treatment included species of subg. Suhuliflorae and subg. 
Ohtusae, sensu Alexeev, in subg. Suhulatae sect. Ohtusae, and moved a single 
species from sect. Elmera, sensu Alexeev, to sect. Suhulatae. In addition, Lu (1992) 
described sect. Longiglumes S.L. Lu to include some Chinese species within subg. 
Suhulatae. Clearly, there are considerable differences of opinion among promi- 
nent agrostologists as to possible taxonomic relationships among these Festuca 

Section Suhulatae consists of about 25 species from North and South 
America, Asia, and Africa (Aiken et al. 1997; Alexeev 1977, 1980, 1988; Lu 1992; 
Stancik ined.). Individuals of Festuca sect. Subulate are typically loosely tufted 

with extravaginal innovations lacking cataphylls, with flat blades without well- 
developed ribs, and have relatively large, open panicles with numerous branches. 
In this section the spikelets are typically lanceolate with two short, entire, and 
narrow glumes that are membranous to coriaceous/membranous. The coria- 
ceous/membranous lemmas are lanceolate, entire, and usually long-awned or 
rarel y awnless. Anthers are short (1-2 mm long) and the ovary hairy, rarely gla- 
brous. The two new species discussed here clearly are members of subfamily 
i\ioideae, tribe Poeae, and subtribe Lolimae (Soreng et al. 2001). With the addi- 
tion of these two new species m South America, Festuca sect. Suhulatae con- 
sists of the following eight species: F. cochahamhana E.B. Alexeev, F. coromotensis 
Briceiio, F. cuzcoensis, F.Jlacca Hack, ex E.B. Alexeey F. parodiana (St.-Yves) 
Nicora, F sodiroana Hack. ex E.B. Alexeev, F tovarensis, and Fulochaeta Nees ex 

Festuca cuzcoensis Stancik & P.M. Peterson, sp. nov (Fig. l). Type: PERU. 
Departamento Cuzco. Provincia Urubamba: above Ollantaytambo, in shade, high 
up ravine, 3600 m, 5 Dec 1923, AS. Hitchcock 22532 (holotype: US-1164548!). 
Festuca ulochaeta auct. non Nees ex Steud: Hitchcock 1927, 321 p.p. Macbride 1936, 123 p.p. Tovar 
1993, 90 p.p. 

multmodis (6-12 non 2-4) et aristae brevi [3-4 (-5) non 5-12 mm] differt. A Festuca ulochaeta Nees 
c \ Steud et F tochabambana E B Alexeev ligula folii 2-3 (non 0.2-2) mm longa distmguitur. 
Perennial, loosely tufted Culms 80-110 cm tall, erect, glabrous, with 6-12 nodes 
m basal half, the mternodes short. Sheaths membranous, brown, more or less 
pubescent, margins free, innovations extravaginal; auricles absent. Ligules 2-3 
mm long, membranous acuminate, dentate. Blades 12-15 cm long, 3-9 mm wide, 
flat gi ecn scabious on i ibs of abaxial surface. Panicles 20-25 cm long, 5-10 cm 
wide open blanches pendant, scabrous. Spikelets 8-10 mm long; florets 3-4 
(pcitect) lachilla 12-14 mm long, puberulent. Glumes narrowly lanceolate, 
coi laccous purplish scabrous, apex acute; lower glume (2-)2. 5-4.5 mm long, 1- 
vemcd uppei glume 4-6 mm long, inconspicuously 3-veined. Lemma 7-8.5 mm 
long lanceolate, chaitaceous to membranous, 3- or 5-veined, purplish-green; 
apex scabious, entiie awn 3-4(-5) mm long, scabrous, straight. Palea almost as 
long as lemma, 2-keeled, the keels finely scabrous, deeply 2-dentate. Lodicules 
oblong 2-dentate Stamens 3; anthers (1.2-)1.5-2 mm long. Ovary apex with a 
lew hairs Cai yopses lanceolate. 

Dist n bution and hah tat.— This species was collected in Andean forest zone 
ol Peru and northeastern Bolivia between 3200^3850 m. 

Etymology —The specific epithet refers to the type locality region, i.e., De- 
paitment Cuzco m southern Peru. 


I Sk I 

Festucacuzcoensis {Hitchcock 2253) A Habit B Inflorescence C Sheath, ligule,and portion of a blade D Spikelet 

)• 05' S), 3700-3850 n 

Festuca tovarensis Stancik & P.M. Peterson, sp. nov. (Fig. 2). Type: PERU. 
Departamento Huancavelica, Provincia Tayacaja: Chuspi-Hda. Tocas, entre 
Colchabamba y Paucarbamba, monte bajo, 2800 m, 22 Apr 1954, O. Tovar 2057 

Loosely tufted perennials. Culms 70-90 cm tall, erect, glabrous, with 2-4 nodes 
in basal half. Sheaths membranous, brown, margms free; innovations extrav- 
aginal; auricles absent. Ligules 1-2 mm long, membranous, truncate, margms 
ciliate. Blades 8-15 cm long, 1.5-4.5 mm wide, linear, flat, green, scabrous with 
ribs on abaxial surface. Panicles 15-20 cm long, 7-10 cm wide, open; branches 
pendant, scabrous. Spikelets 7.5-9.5 mm long; florets 3 (perfect); rachilla 1.1-1.4 
mm long, puberulent. Glumes narrowly lanceolate, coriaceous, purplish, gla- 
brous, apex acute (sometimes scabrous); lower glume 1.5-1.8 mm long, 1-veined; 
upper glume 3.5-4.5(-5) mm long, 1-3 vemed. Lemma 5.5-6.5 mm long, lan- 
ceolate, chartaceous to membranous, 3-veined, purplish-green, scabrous; apex 
entire; awn 5-7 mm long, terminal, scabrous, straight. Palea as long as lemma, 
2-keeled, the keels scabrous; apex hairy, deeply 2-dentate, Lodicules ca. 0.8 mm 
long, lanceolate, acuminate. Stamens 3; anthers 1.5-1.6 mm long. Ovary sparsely 
hairy near apex. Caryopses lanceolate; hilum linear, 2/5-1/2 of total length. 

Distribution and habitat.— This species is known only from the Andean 
forest zone of Peru, at 2500-3250 m. 

Etymology— The specific epithet honors the eminent Peruvian botanist and 
agrostologist, Oscar Tovar Serpa. 

3 Yumagual, 2500 m, 26 Jun 

epresented by eight native species (Table 
t morphological features that distinguish 
■ican species, E suhulata Trin. All the spe- 

/£ ^D C 

Fig. 2. Festuca tovarensis {Tovar2057). A. Habit. B. Sheath, ligule, and portion of a blade. C. Spikelet. D. Floret. E. L 
ventral view. F. Palea with stamen. G. Palea, with lodicules, pistil, and stamens. H. Lodicules. I. Pistil. J. Stamen. I 
opsis, dorsal view. L. Caryopsis, ventral view. 

F. flacca F. sodiroana 

Ecuador Ecuador, 


cies are morphologically very similar to one another. However, F.ulochaeta can 
be easily separated from the remainmg seven species by having long-awned 
lemmas with the awn f lexuous whereas the other taxa have straight awns or 
awnless lemmas. Festuca cuzcoensis and F. tovarensis are possibly most closely 
related to Fflacca since all three share lanceolate spikelets of the same size, 2- 
4 florets per spikelet, straight awns 3-12 mm long, and small anthers 1.1-2 mm 
long. Festuca cuzcoensis differs from F tovarensis by having 6-12 nodes per culm 
(2-4 nodes in F. tovarensis), acuminate ligules (verses truncate), lower glumes 
(2-)2.5-4.5 mm long (verses 1.5-1.8 mm), lemmas 7-8.5 mm long (verses 5.5-6.5 
mm), and a glabrous ovary (verses sparsely hairy). Festuca parodiana shares 
the following characteristics with F sodiroana: truncate ligules, 4-6 florets per 
spikelets, and awnless lemmas. However, F. parodiana can be separated from F. 
sodiroana by having a hairy (glabrous in F. sodiroana) ovary, lower glumes 2- 
2.5 mm long (verses 1.3-1.8 mm), upper glumes 3-4 mm long (verses 2.2-2.9 
mm), lemmas 7-8 mm long (verses 5-6.5 mm), and 5-7 nodes per culm (verses 
2-4). Festuca coromotensis is almost awnless (mucronate) but differs from F 
sodiroana and F parodiana by having large, scabrous lemmas 8-9 mm long. 
There appears to be a geographic component as well since the principal area of 
distribution for F ulochaeta is southeastern Brazil and northeastern Argentina 
with new range extensions reported from the Cordillera Oriental of Colombia 
and Cordillera de Merida m Venezuela (Stancik 2001). The remaining seven 
species occur exclusively in the montane forest zone of the Andes in small en- 
demic populations ranging from Argentina in the south to Colombia and Ven- 
ezuela in the north. Festuca elviae Bricefio, a Venezuelan species, is also a pos- 
sible member of Festuca sect. Suhulatae. However, further study is needed to 


?mmas 6.5-8 mm long.glabrous or pa 


Culms with 2-4 nodes; lower glumes 

1.2-1. 8 mm long; I 


;s 2.0-2.9 

mm long; lemmas 5-6.5 mm lonq;ov 

ary glabrous 

F. sodiroana 

Culms with 5-7 nodes; lower glumes 

2-2.5 mm long; uf 

Dper glumes 

long; lemmas 7-8 mm long; ovary ha 

mas with awn 3-1 5 mm long. 



Lemma awns straight, 3-12 mm long 

5. Sheaths glabrous, rarely scabrous. 

6. Lower glumes (2-)2.5-4.5 m, 

T1 long; lemma c 

,wns 3 4 n 

nm long 

We wish to thank the Smithsonian Institutions, Fellowships and Grants for 
supporting a research visit by the senior author to the United States National 
Herbarium. The senior author would like to thank ICETEX for financial sup- 
port in Colombia, the Instituto de Ciencias Naturales de la Universidad Nacio- 
nal de Colombia for use of their study facilities, and the Grant Agency of the 
Czech Republic (Grant GACR No. 42-201174) for financing fieldwork m Ecua- 
dor. We thank curators from the following herbaria: AAU, B, COL, LPB, MA, PR, 
PRC, QCA, QCNA, VEN, and W. Appreciation is extended to Alice R. Tangermi 
for providing the illustrations, Dan Nicolson for reviewing the Latin diagnoses, 
and Susan Pennington for technical support. Gerrit Davidse, Stephan L. Hatch, 
Robert D. Webster, and Barney Lipscomb are thanked for reviewing the manu- 
script on short notice. 


(Poaceae) in North America: Diagnostic evidence from DELTA and clustering programs, 

and an INTKEY pacl<age for interactive, illustrated identification and information re- 

trievafCanad.J.Bot, 75:1 527-1 555. 
Alexeev,E.B. 1 977.0n the systematics of Asian fescues {Festuca). 1 .Subgenera Drymanthele, 

Subulatae, Schedonoms, Leucopoa. Bjull. Moskovsk. Obsc. Isp. Prir., Otd. Biol. 82:95-1 02. 
Alexeev, E.B. 1 980. Festuca L. subgenera et sectiones novae ex America et Mexica. Novosti 

Sist.Vyss.Rast. 17:42-53. 

Aeexeev, E.B, 1986. Festuca L (Poaceae) in Venezuela, Colombia et Ecuador. Novosti Sist. 

ALEXEEV,E.B.1988.Genus festuca (Poaceae) in Japonia, Peninsula CoreanaetlnsulaTaiwan. 

Novosti Sist.Vyss.Rast. 25:5-27. 
Brako, L and J.L Zarucci. 1 993. Catalogue of the flowering plants and gymnosperms of 

Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45:1 -1 286. 
Clayton, W.D.andS.A.RENvoizE.1986.Generagraminum. Grasses of the world. Her Majesty's 

Stationery Office, London. Pp. 93-94. 
Hitchcock, A.S. 1 927.The grasses of Ecuador, Peru,and Bolivia. Contr. U.S. Natl. Herb.24:291 - 

Lu, L.S. 1 992. New taxa of Festuca L from China. Acta Phytotax. Sinica 30:529-540. 

SoRENG, R.J., G. Davidse, p.m. Peterson, P.O. Zuloaga, E.J. Judziewicz, a 
logue of New World grasses (Poaceae), supragenei 

Stancik, D. 2001 . New records of the genus Festuca L (Poaceae) 
ela.Caldasia 23:337-339. 

TovAR, 0. 1 993. Las gramineas (Poaceae) del Peru. Ruizia 1 3;1 - 

Turpe, A.M. 1 969. Las especies argentinas de Festuca (excluidas L 

s D M M sn H Roberto Kii m inc and Cari os Omoi \z \ 2002 A Cactus Odyssey. 
CISBN 0-88192-526-8, hbk ) Timber Pi ess, Inc 133 S W Second Avenue Suite 
450 Portland OR97204-3527,USA (Ordeis ww^^tImbelplesscom 800- 
327-5580 503-227-2878, 503-227-3070 fax) $39 95, 306 pp, 191 color pho- 
tos, 4 maps 6x9 


Botanical Research Institute ofTexas 
Fort Worth, TX 76102-4060, U.S.A. 
mdis, sp. nov., IS segregated from plants previously identified as E multiflorum Benth. 
.5 known from f 7 counties of the Rio Grande plains in south-central Texas. Eriogonu m 
s from E muUiJhrum in its broader leaves with auriculate-subclaspmg bases (vs. 

i-f2(-15) flower 

parently c 


largos, mvolucros esparsamente tomentosos y lobulos del perianto glabrescentes, y cordados. 
The Texas species of Eriogonum Michx. were treated in detail by Reveal (1968, 
1970), who observed (1968, p. 203) that E multiflorum Benth. and E. annuum 
Nutt. "are the most frequently collected species in the state and collectively the 
most widely distributed species." Together, they constitute subg. Micrantha 
(Benth.) Reveal (Reveal 1969). Reveal provided a set of detailed morphological 
contrasts to distinguish these two taxa, noting that their difference in perianth 
lobe shape allows easy identification. He did not call attention to significant 
infra-specific variation in either species. 

The present study finds that plants previously identified as Eriogonum 
multiflorum can be separated into two, morphologically distinct sets of popu- 
lations. Those occupying the largest part of the range m Texas, along with those 
from peripheral localities in northwestern Lousiana, southwestern Arkansas, 
and south-central Oklahoma (Fig. 2), are identified here as typical E. 
multiflorum. The type collection of E multiflorum (Bentham, Trans. Linn. Soc. 
London 17:413. 1837) was made in 1833 by Thomas Drummond (Drummond 

"between Brazoria and San Feiipe de Austin, perhaps m what is now Austin 
County (from where a modern collection of E. multijlorum also has been made). 
An illustration based on the Drummond type material, presumably at K (Hook. 
Icon. PI. 3, t. 250. 1840) shows leaves with non-clasping bases and the descrip- 
tion notes "Involucres with about 6-9 f lowers"-these features (see below) along 
with the general habit unambiguously establish the identity of the species. 
Observation of additional type material of E. multijlorum (GH photocopies!; 
isotype and syntypes fide annotations by J. Reveal m 1967) corroborate this. 
Plants identified as E. multijlorum m the Rio Grande plains of southern Texas 
represent a previously undescribed species. 

Eriogonum riograndis Nesom, sp. nov (Figs. 1, 2). TYPE U.S.A. TEXAS. SAN PATRICIO 
Co ■ 9 mi E of Sinton, Welder Foundation Research Area, near headquarters build- 
ings, grassy held along swale and m swale, 28 Oct 1958, F.W. Gould 8510 (holo- 

Annual or biennial herbs from a taproot, eglandular. Stems erect, 2.5-6(-ll, -20 
fide Reveal 1970, Richardson 1995) dm tall, usually unbranched until the inflo- 
rescence, closely but loosely gray-white and persistently wooUy-tomentose. 
Leaves mostly on the lower 3/4-4/5 of the stem, basal not persistent, alternate, 
oblong-obovate to oblong-elliptic or oblong, epetiolate and shallowly auricu- 
late-subclasping, apex rounded to obtuse, 15-40(-45) mm long, 10-22(-25) mm 
wide, margins flat or narrowly revolute, entire but often closely undulate, up- 
per surface lightly but persistently woolly-tomentose, lower surface densely 
and persistently white- to tawny-tomentose. Inflorescences cymoid, usually 
relatively compact. Involucres on peduncles 2-5 mm long, turbinate to turbi- 
nate-campanulate, 2-2.8 mm high, 2-3 mm wide, 5(-6)-lobed, externally 
sparsely but persistently tomentose to glabrate, mternally densely tomentose. 
Flowers 22-28(-36) per involucre, exserted from the involucre on filiform 
pedicels. Perianth bright white to pinkish, maturing or drying orangish, exter- 
nally glabrous, internally sparsely tomentose; perianth lobes dissimilar, outer 
lobes basally cordate and broadly oblong to oblong-elliptic, 2-2.7 mm long, 1.5- 
2.2 mm wide, midrib thick from base to apex, inner lobes linear, 0.1-0.2 mm 
wide. Fruits dark brown to red-brown, 1.9-2.2 mm long, not winged, glabrous. 

Habitat and phenology.— Fields, pastures, roadsides, mesquite prairies, oak 
woodlands, and other open habitats, sandy soil; (Aug-)Sep-Nov(-Dec). 

Distribution.-The distribution of Eriogonum riograndis (Fig. 2) corre- 
sponds to phytogeographic pattern #17 of Sorrie and Weakley (2001): South 
Texas-Northeast Mexico Mesquital. They note that "The northern portion is 
commonly denoted as the South Texas Plains, but ecologically the whole area 

tand details of Eriogonum riograndis (from holotype, BRIT). 

is more properly termed the Tamaulipan Scrub region, or Tamaulipan Mesquital, 
due to the dominant shrubby vegetation, especially Prosopisglandulosa Torr. ..." 
(Sorrie & Weakley 2001, p. 61). The distribution of the recently described 
Pseudognaphalium austrotexanumNesom (Nesom 2001) also fits the same phy- 
togeographic pattern. Reveal (1970) noted that E multiflorum occurs m Mexico; 
presumably this would be E. riograndis in the interpretation here, but a speci- 
men documenting the Mexican distribution has not been seen in the present 

Etymology-The appositive epithet refers to the central geographic feature 
in the range of the nev/ species, the Rio Grande (river). 

:ory 20336 (TAES), Aransas ReCuj; 
5 beyond back beach, 7 Nov 196- 
mient Shed, 18 Sep 1968 Flectwoc 

woodland, 29 Dec 1982 Fryxt 
irks and Cory MHiniAFSl I 
a few mi E of Jim Hogg Co hr 

(TEX); 4 mi SE of King Ranch, Encmo Division Headquarters, in deep, loose sand blow-out area, 18 

Pleistocene gravel, 25 Nov 1962, Dohnke 6 (SMU, TEX); 12 mi E of Hebbronville, Hwy 285, 10 Nov 

gate, SE of Falfurias, open hillside to edge of oak mott with deep sandy soil, 31 Oct 1990, 5, & G.Jones 
6081 (VDB), Calhoun Co.: Matagorda Island, 1955, Balls, n. (TAES); Matagorda Island, access road from 
air base HQ to beach at end of island, ca 2 mi E of base, 22 Jul 1973 [not flowering], Hartman 3722 
(TEX). Duval Co.: Benavitas, 340 ft, 12 Aug 1941, Fisher 41101 (TEX). Hidalgo Co.: N of Edinburg, 11 
Nov 1942, Walker 126 (LL); ca. 30 mi N of Edinburg, in sand, 23 Aug 1944, Whitchouse 44,276 (SMU); 
Bentsen Ranch, 10 mi N of Edinburg, in sand, 2 Nov 1973, Everitt s.n. (SMU); La Reforma Training 

on dry level upland, 6 Oct 1993, Carr 13177 (TEX); La Reforma Training Area (Texas National Guard), 

tered mesquite, 255 ft, 250ct 1994, Carr 14302 (TEX). Jim Hogg Co.: State Hwy 285 E of ThompsonviUe, 
in light brown loose sand, 11 Nov 1962, Ramirez, Alva, and McCart 8708 (SMU, TAES, TEX); 10 mi N 
of La Gloria on Texas Farm Rd 1017, 15 Oct 1990, Miller, Brant. & Noyes 5811 (VDB); E side of FM 1017, 
3.0 road mi S of jet with smaller road at Agua N ueva, ca. 450 ft, occasionally mown grassland m deep 
loose sandy soil, roadside, 7 Oct 1993, Carr 13208 (TEX). Jim Wells Co.: sandy loam m dry lake, 
Romarsid Ranch, 1 Nov 1943, Freeborn 142 (TEX). Karnes Co.: 2 mi S of Karnes City, frequent in dry, 
deep sandy soil, fallow field, 27 Oct 1952, Johnson 1010 (LL, SMU, TAES); 3 mi NE of Kenedy dry 
Escondido Creek, infrequent in dry sandy loam soil, 25Jul 1954 [very early bud], Johnson ]618(SMU). 
Kenedy Co.: 18.7 mi N of Raymondville, frequent along road in sandy soil, 2 Dec 1945, Cory 51495 
(SMU); Norias Division of King Ranch, Saltillo Pasture, 14 Sep 1953,Johnston s.n. (TAES, TEX), 17 Sep 
1953, Johnston s.n. (TAES, TEX), 24 Nov 1953.Johns(on s.n. (SMU, TAES, TEX); 18 mi S of Riviera, dune 
area, 12 Oct 1952, Morrow and Nord 19 (TAES); Nonas Division of King Ranch, SE corner of ranch, 
open sandy coastal plain, 23 Sep 1958, Lundell and Corrcll 15155 (BRIT LL); 3.4 mi N of Armstrong, 
fine gray sand, 6 Nov 1953, Shinners 17082 (SMU). Kleberg Co.: Padre Island, 17 Nov 1940, Cory 36795 
(SMU), Cory 36796 (TAES), Cory 36797 (TAES), Co ry 36798 (SMU), Cory 36799 (TEX); KingsviUe, sum- 
mer 1940, Sinclair s.n. (TEX); Laureles Division of King Ranch, loose sand, 15 Sep 1953, Johnston s.n. 
(TEX), 29 Nov 1953, Johnston s.n. (TEX); Brooks County line on Road No. 285, loose sand mesquite 

to dump area, small stabilized dunes near margin on brackish back island pond, 14 Oct 1989, Lemke 
3009 (TEX). Nueces Co.: Copano Bay 5 Oct 1922, Tharpl552 (TEX); Corpus Christi, 18 Dec 1935 [past 
fruit], Parhs 17437 (TAES), 17438 (TAES); Mustang Island, loose sand, 26 Oct 1954, Hildehrand 82 (TEX); 
Mustang Island, sand dunes 300 yds from Gulf, 11 Nov 1965, Crutchfield 1027 (LL); Mustang Island 
cemetery, Port Aransas, sandy soil near fence, 14 Oct 1967, Gi I iespie 110 (TEX); Mustang Island, Sham- 
rock Bay area, west side of island, 29JuI 1967, Gillespie 200 (TEX); Mustang Island State Park, NW of 
Park Rd 53, 500 ft SW of Corpus Christi Pass, dry sand along elevated margin in tidal flat, 500 ft, 20 
Oct 1989, Carr 10158 (TEX). Refugio Co.: Blackjacks below Austell, 4 Dec 1928, Phipps s.n. (TEX). San 
Patricio Co.: 1.8 mi SW of Aransas Pass, 19 Sep 1936, Parhs and Cory 20338 (TAES); 4 mi SWof Ingleside, 
19 Sep 1936, Parks and Co ry 20339 (TA ES); 2 mi SE of Ingleside m deep fine sand, 2 Oct 1950,Jones 383 
(SMU); 1 mi S of Ingleside in deep fine sand, 8 Jun 1951 Ipre-f lowering],Jones 565 (SMU); Welder Wild- 
tight sandy loam, 29 Sep 1956, Rowel 1 5212 (SMU). Webb Co.: 3 m i S of M i rando City at Los Ojuelos, in 
grayish black loose sand, 10 Oct 1961, Magnon and Rodriguez 14 (SMU). Willacy Co.: Yturria, 50 ft, 6 
Aug 1924 [not in flower], Runyon 665 (TEX); along hwy from Yturria Station north, 24 Sep 1937, 
Runyon 1808 (TEX); Yturria, open dry sandy ground, 10 m, 22 Sep 1939, Runyon 4237 (TEX); 
Raymondville, 5Jul 1941, Shiller 759 (TAES, TEX); Sauz Ranch, sand, 23 Nov 1953,Johnston and Davis 
s.n. (SMU, TAES, TEX). Zapata Co.: 7 mi N of Zapata, US Hwy 83, 15 Nov 1961, Munoz, McCart, and 
Cahrera 39 (SMU, TEX). 

uuidtiflomm u^a^ } u\ 


s c ui be distuiguishcd bv a simple 


n leal 

moiphologv A count of tloweis f 

usually requning low r 


ification confnms their distmctic differences between 


wo are summarized m the followi 

mentose to weakly glabre 

(6-)Q 12( 16) pennvoiucre_ 

long cordate 

\oluciesol E annuumd\^o uc on shoitLi peduncles (subscssilc to 1-3 mm long) 
olten gnmg the mfloicsccnLL bi inches i sc)mc\\hat sccund appeaiance and 
each involucre produces 25 oi moic tlov\cis (similai to those of E i lograndis) 

Enogonum nograndis and E multijloi um aie primarily allopatnc m geo 
graphic range (Fig 2) butE multif/oru m has been documented withm the range 
of E riogiandi^ 

Zapata Co E n(\/c;/ic/!s indE fHii/df/oi um both have been collected here 
Plants of the lattct (Hc/ni In 1711 TTX) Ui t\ pic d m leaf morphology and flower 

A few plants with ippuuitU intLi mediate featuies ha\e been collected 
(Fig 2) 

Dimmit Co (f8 mi F of El Indio Millu et al 5775 TAES) leaves narrow 
sessile but flowers 36 per involucre This plant has leaves of E multiflorum but 
ahighnumbei of flowers hke E nog^ and is Hog/unJ sn (Cairizo Springs TEX) 
has nai row sessile lea\ es but 20-21 1 low l \ s pti in\ oluc i c 

Karnes Co (2 mi S of KinusC it\ /oil n son 1010 f I SMU TAES) leaves rela 

basally tiuncate to subaui Kulatc and sulxl \spmg mvohicies 18-27-t lowered 
(tending toward intermediate m flowei numbei) These plants aie identified 
here as E nograndis Another fiom Kaines Co {.Johnson 1618 SMU) is llorally 

immature but similar m leaf morphology tojohnson 1010, but the tendency for 
smaller number of flowers and the sessile lower leaves indicate that genes from 
E multiflorum may be present. 

Wilson Co. (Kicaster, Parks Rx 3016, TEX): upper leaves clasping to 
subclasping, flowers ca. 18-22. This plant is similar to those o[ Johnson 1010 in 
Karnes Co., apparently mtermediate. Parks and Cory 11800 (E of Floresville, 
TAES) has narrow sessile leaves but 24 flowers. Parks 5355 (Kicaster school, 
TAES) is typical E multiflorum. 

Choice of rank -Conspicuous morphological and geographic discontinuity 
between Eriogonum multiflorum and E riograndis is the predominant feature 
of their contrast. Putative morphological intermediates suggest that hybrid- 
ization may occur where they come into geographically close contact, but no 
zone of intergradation exists to suggest that significant gene flow occurs be- 
tween the two taxa. It might be argued that their apparent sister relationship 
should be recognized by treating them as varieties within a single species, but 
the degree of morphological differentiation between species in any given ge- 
nus (including Eriogonum) is variable, and the discontinuity documented here 
(with inference of genetic isolation) provides justification for treating E 
multiflorum and E riograndis at specific rank, consistent with species concepts 
in many other genera. 

I am grateful to the staff of TEX-LL for their help during a recent visit there, 
TAES for a loan of specimens, Emily Wood (GH) for sending photocopies of 
type material of E multiflorum, Amy Buthod at OKL for helpful information 
on Oklahoma distribution, staffs of CAMU and OKLA for sending photocopies 
of Oklahoma collections, and to Jim Reveal for comments on the manuscript. 
Linny Heagy provided the fine illustration; a discussion with Ted Barkley re- 
garding the choice of epithet was helpful 


MacRoberts, DT 1 989. A documented checklist and alias of the vascular flora 
Louisiana State Univ, Shreveport, BulL Mus. Life Sci. No. 9. 

cies from southeastern Texas and adjacent Mexico. Sida 1 9:507-5 1 1 


Reveal, J.L 

1 968. Notes on the Texas Enogonums. Sida 3:1 95-205. 

Reveal, J.L 

RevealJ.L. • 

1 969. The subgeneric concept in Eriogonum (Polygonaceae), In: J. Gunckel, ed. 
: topics in plant science. Academic Press, New York. Pp. 229-249. 
1 970. Eriogonum. In: D.S. Correll and M.C.Johnston. Manual of the vascular plants 
;.Texas Research Foundation, Renner. Pp. 5 10-5 16. 


,A1995.Plantsofthe Rio Grande Delta. Univ. of Texas Press, Aust 


Botany and Bacteriology, Univ.c 




Robert J. O'Kennon and Guy L Nesom 

endemica de la region de la meseta Edwards de Texas, de donde se conoce de seis condados. Se separa 
de I. costellata por su combinacion de pedunculos mas cortos y flores bianco brillante, casi rodadas 

Plants of a small-flowered, pedatisect-leaved morning glory from the Edwards 
Plateau region of Texas are identified as Ipomoea costellata Torrey but are dis- 
tinct from other plants of the species in morphology, phenology, and geogra- 
phy The Edwards Plateau plants are here recognized as a distinct variety 

Ipomoea costellata Torrey var edwardsensis O'Kennon & Nesom, var nov; (Figs. 
1, 2, 3). Type: U.S.A. TEXAS. Travis Co.: Colorado River, Montopolis Bridge, 8 Nov 
1934, B.C. Tharps.n. (holotype; TEX). 

Plants annual herbs from a filiform taproot, at first erect, becommg prostrate or 
clambering on low vegetation, slightly twining at the very tips. Stems simple or 
with 1-5 (or more) branches originating ca. 2-4 cm above the base, terete, 3-60 
cm long, 0.5-1 mm in diameter, branching, green, glabrous. Leaves pedatisect, 
2^3.5 cm wide, petioles 7-25 mm long, nearly even in length or decreasing 
slightly from base to apex of stem, glabrous, ultimate leaf segments 7-9(-ll), 
linear to linear-lanceolate, unequal, the longest 15-24 cm long, 1.5-3 mm wide, 
outer shorter than the inner and usually lobed from near the base, green above 
and beneath, glabrous, margins entire, sparsely hispid-ciliate, apices acute, api- 
culate. Flowers solitary or less commonly paired; peduncle spreading to slightly 
decurved or ascending, terete, filiform, 3-22 mm long, green, glabrous to sparsely 


hispid-pilose; pedicel 6~8 mm long, slightly thicker than the peduncle, sepa- 
rated from the peduncle by a pair of awl-shaped bracts, glabrous to sparsely 
hirsute-pilose; sepals equal, subimbricate, elliptic to elliptic-lanceolate, 4-5.2 
mm long, 1.5-2 mm wide, midrib green, low-carinate, smooth to muricate, mar- 
gins entire, hyaline, the apex rounded to retuse, apiculate; corolla tubular to 
long-campanulate, all parts bright white, glabrous, 8-11 mm long, tube 4-5 mm 
long, 2 mm m diameter, limb with 5 oblong-ovate, apiculate, spreading lobes 
2-2.5 mm long, flowers opening with lobes neariy erect, spreading at right angles 
at full anthesis, closing erect past maturity; stamens subequal, white, filaments 
pubescent along whole length with long, viscid trichomes; style white, 4.5 mm 
long (equaling the filament length), glabrous, stigma white, slightly bilobate, 
apparently in contact with the anthers. Fruits bilocular dehiscent capsules 5-6 
mm in diameter, tan upon drying, chartaceous, glabrous; seeds 4, dark brown 
to blackish, obovoid, 3-angled, ca. 3 mm long, 2 mm wide, densely and minutely 



)ns examined. United States. Texas. Bexar Co.: Governm 

ent Canyon State Natu- 



'ildcat Canyon, E edge of San Geronimo Quad or W edge 


;lotes Quad, 'guess- 

.ted' la 


?g" 33'5i" N, 98- 45'00" W, elev. 1200-1240 ft, rare, two pi; 


seen among sparse 


on in very shallow dark brown clay in natural opening c 


jtaceous limestone 

■ock ex 

posed or 

1 flattest part of ridgetop, associates include S^orohoXus 




1 var songarica. Croton monanthogynus. Senna lindheimmana, \ 

?4 0( 

zt 1995, Carr 15050 


net Co.: \ 

^ of RM Road 1431, 1.8 road mi WNW of US Rte 281, Ma 


^alls Quad, local in 

, fairly 

dry, orga 

nic-sandy soil in unshaded weather pit on low granite c 


op, ca. 1500 ft elev, 

jal, ste 


,ng only at tips of taller plants, 18 Aug 1988. Carr 9B6 


Kutac, Lynch, and 



oad 1431, 1.7 mi W of jet with US Rte 281 in Marble Falls, 


of exposed granite, 


ad across 

from large quarry, 23 Oct 2001, Nesom ¥W99 and O'Kennc 

3n (BRIT TEX). Gillespie 

e Onio 

n Creek, 

19 Oct 1990, O'Kcnnon 8125 (BRIT). Llano Co.: near the s 

lit of Dutch Moun- 

ca. 1.5 

mi N of I 

mchanted Rock, infrequent vine, 29 Sep 1976, Butterwick and 

L. mb 3303 (TEX). 

Austin, ( 

Zolorado River below Dam, 29 Sep 1929, Ecology Class 5.; 

k,15 0, 


Vhitehouse W-29-3 (TEX); McKinney Falls State Park, S o 


on Creek at mouth 


son Creek, frequent, locally abundant in thm soil in weather pits 




) ft, 29 Oct 1985, Carr 7050 (BRIT TEX). Uvalde Co.: 2.1 m 


049 or 


dry limestone ledge, 2 Nov 1985, Kcency5371 (BRIT). 



-The epithet alludes to the location of the plai 

)n the Edwards 


Distribution, habitat, and pheno Zogy.-Collections at hand indicate that Ipo- 
moea costellata var edwardsensis is distributed over the Edwards Plateau, al- 
though the plants are apparently uncommon and inconspicuous. Shinners' 
treatment of Texas Ipomoea (1970) noted only that I. costellata occurs in the 
trans-Pecos region, although in I960 he annotated the three early collections 
from Travis County (1929 and 1934, cited above) as I. costellata; McDonald (1995) 
did not map or cite any collections of I. costellata from the Edwards Plateau. We 
suspect that searches for var edwardsensis will broaden its known distribution 

but confirm that it is endemic to the Edwards Plateau. Limestone ledges and 
crevices, and thm soil of weather pits in bedrock of limestone and granite, 100- 
1500 ft elev. Flowering (August-)September-November. We observe that var. 
edwardsensis apparently is particularly sensitive to grazing. 

Ipomoea costellata (excluding var. edwardsensis) occurs in Texas in the trans- 
Pecos region and in Webb Co. (Fig. 3), continuous with its distribution in the 
Mexican states of Chihuahua and Coahuila. It is recorded from all of the north- 
ernmost states of Mexico (Sonora, Chihuahua, Coahuila, Nuevo Leon, and 
Tamaulipas) except Baja California and continues southward to Guatemala (see 
McDonald 1995, Fig. 2). Austin and Huaman (1996) reported its occurrence also 
in Venezuela but not Guatemala. 

A recent taxonomic revision (McDonald 1995) treats Ipomoea costellata and 
seven closely related species within Ipomoea sect. Leptocallis (G. Don) J.A. Mc- 
Donald, and another species has been recently added (Austin & Tapia Munoz 
2001). Ipomoea costellata is distinct withm sect. Leptocallis m its annual, 
taprooted habit and small flowers (ca. 1 cm long). Ipomoea costellata alone in 
the section is autogamous-self-pollination is facilitated by the stigma m con- 
tact with stamens. Variety edwardsensisalso has this reproductive arrangement 
and is obviousl y closely related to the rest of the species, but its geographic sepa- 
ration and set of differences (couplet below) provide a reasonable basis for for- 

Ipomoea flowers are fragile and commonly are nc 
tinctions in pressed specimens. We emphasize the short peduncle length, espe^ 
cially of mature fruits, as the most obvious and easily observed feature to dis- 
tinguish var. edwardsensis. Scattered plants of I. costellata sensu lato (var. 
costellata) may have peduncles approaching the short length of var. 
edwardsensis, but the latter (as a population system) is set apart from rest of 
the species by this feature. 

The flower color of Ipomoea costellata was described by McDonald (1995) 
as "blue, or rarely yellow throughout." Other sources note the following: "pale 
lavender" (Texas-Whitehouse 17154, SMU), "pale pmk" (Texas-Turner et al. 
53447, SMU), "reddish" (Texas-Keough 227, TEX), "purple" (Texas-Butterwicfe 
and Lamh 1759, TEX), "rose-purple" (New Mexico-Martm & Hutchins 1981), 
lavender (photo of Texas plant-Rickett 1969), "limb pale pink, throat and tube 
near white" (New Mexico-SpeHen berg 3852, LL), "lavender or purplish, tube 
paler" (Sonora-Wiggins 1964), "tube lavender, purple near tips, yellow-white 


,^_iinj_i..i_u..+f-Kj>' ■" 


at base" (Chihuahua-Hen ric/eson 1551, TEX), "yellow tube, purple-red limb" 
(Chihuahua-Henncfeson 1691, TEX), "cream with green-yellow throat" 
(Coahuila- Wendt llSly, TEX), and "pale blue" (Chiapas-Breedlove 511>51>, TEX). 
Plants in the vicinity of Edo. Mexico produce yellow flowers (according to Mc- 
Donald) and have been segregated as I. painteri House— this taxon was included 
in the synonymy of I. costellata by McDonald but treated as distinct by Rico R. 
(1985), who described the flower color of I. painteri as white or creamy Plants 
in Nuevo Leon and Tamaulipas also apparently produce white flowers and are 
further distinguished within the species by thick taproots and apparently pe- 
rennial duration; peduncles of these plants are long. 

The corolla morphology of var. edwardsensis also is distinctive from any 
we are aware of within Ipomoea costellata and certainly from populations of 
the species on the periphery of var edwardse nsis m Texas and northern Mexico. 

In var. edwardsensis, the limb is deeply and relatively narrowly lobed, and at 
full anthesis, the lobes spread at nearly right angles to the tube (Fig. 2). The Ime 
drawing (Fig. 1) shows a flower as it appears in early anthesis, just opening. Flow- 
ers of I. costellata elsewhere in its range apparently have ten, shallow lobes and 
the limb is funnelform, widening gradually toward the apex (Rickett 1969, pi. 
99; Warnock 1977, Fig. 6, p. 178)-"corolla campanulate ... limb subentire, scant- 
ily 10-lobate," as described by McDonald (1995, p. 106). 

Variety edwardsensis is cleariy a 'satellite' of the larger and variable I. 
costellata sensu lato. Recognition of var, edwardsensis emphasizes its relative 
internal consistency and its morphological and geographical distinction from 
the rest of the species. Other recognizable geographic variants, particularly as 
distinguished by flower color, have been formed within i. costellata, this pro- 
cess perhaps quickened by the apparent tendency for autogamy Formal recog- 
nition of the Edwards Plateau populations implies that other geographic seg- 
ments of I. costellata might also be justifiably recognized (McDonald pers. 
comm.), and we agree with this. 

McDonald (1995) did not provide a formal statement of his concept of va- 
rietal versus specific rank within sect. Leptoca 1 1 is, but several species in his treat- 
ment include geographically distinct varietal taxa separated by one to several 
apparently non-intergrading morphological characters. In contrast, Yatskievych 
and Mason (1984) recognized two varieties within I. tenuiloha Torrey (of sect. 
Leptoca His) that are morphologically mtergrading, this taxonomy accepted by 
McDonald. Yatskievych and Mason also observed that I. tenuiloha, I. capillacea 
(Kunth) G. Don, and I. plummerae A. Gray (including I. patens (A. Gray) House) 
constitute "a very close-knit species complex." With further consideration of 
the rationale for applying ranks within sect. Leptocallis, we believe that the 
Edwards Plateau variety of I. costel lata eventually may be treated at higher rank. 


We are gratefuf to the staff of TEX-LL for their help during rece 
drew McDonald and Dan Austin reviewed the manuscript and w 
to Andrew McDonald for a discussion of variation m I. costellata. 
provided the fine illustration. 

nt visits. An- 
e are grateful 
Linny Heagy 


Austin, D.F. and Z. EIuaman. 1996. A synopsis of Ipomoea (Convolvulaceae) ir 

ithe Americas. 

Taxon 45:3-38. 
Austin, D.F. and J.LTapia Munoz. 200]. Ipomoea sororia (Convolvu 

from Yucatan, Mexico. Sida 1 9:807-81 0. 
McDonald, A.J. 1 995. Revision of Ipomoea section Leptocalh 

Pap, Bot. 6:97-1 22. 
Martin, W.C, and C,R, Hutchins. 1 981 . A flora of New Mexico. Vol. 2, 

RicKETT, H.W. (ed.). 1 969.Wild flowers of the United 5tates:Volume 3:Texas (part 2). McGraw- 
Hill Book Company, New York. 

Rico Rodriguez, LI 985. /pomoeo.ln:Rzedowski,J.yG.C.deRzedowski,eds. Flora fanerogamica 
del Valle de Mexico. Diseno Editorial: Myriam Cerda. Pp. 250-256. 

Shinners, LH. 1 970. Convolvulaceae (excluding Cuscuta). In: D.S.Correll and M.C.Johnston. 
Manual of the vascular plants of Texas. Texas Research Foundation, Renner. Pp. 1241- 

Warnock, B.H. 1977.Wildflowers of the Davis Mountains and Marathon Basin, Texas. Sul 
Ross State University, Alpine, Texas. 

Wiggins, I.L. 1 964. Flora of the Sonoran Desert. In: F.Shreve and I.L.Wiggins.Vegetation and 
flora of the Sonoran Desert. Stanford Univ. Press, Stanford, CA. 

(Convolvulaceae), with notes on related species. Madrono 31:102-108. 


HFN Anderton 2001 Urban Sanctuaries: Peaceful Havens for the City Gardener. 

(ISBN 0-88192-502-0, hbk.). Timber Press, Inc., 133 S. W Second Avenue, Suite 
450, Portland, OR 97204-3527, U.S.A. (Orders;, 800- 
327-5680, 503-227-2878, 503-227-3070 fax). $29.95, 144 pp, 148 color photos, 
4 color plans, 4 b/w plans, 8 1/2" x 11". 

dunng the deMgn stage the fact that e\ en 
they can be adapted to create outdoor sp, 



Noel H.Holmgren 

describe como una nueva variedad, S. caespitosa var. wiUiamsiae N.H. Holmgren. 

The genus Sphaerakea, consisting of about 40 species, is found in temperate 
and warm temperate parts of Northi and South America. It is a difficuk genus 
taxonomically with poorly deUmited species. Frequent hybridization and back- 
crossing may be the culprits. In North America there are about 26 species with 
the greatest concentration in southwestern United States and northern Mexico. 
Arizona should be considered as the center of diversity for the North American 
members with about 16 species, and the concentration decreases nearly expo- 
nentially with distance in all directions. The Intermountain Region, an area 
more than two-and-a-half times larger than Arizona, is home to 12 species with 
only three extending north of the region: S.grossulariifolia (Hook. & Arn.) Rydb., 
5. munroana (Douglas ex Lindl.) Spach ex A. Gray and S. coccinea (Nutt.) Rydb. 
The latter two species extend to Canada and S. coccinea reaches the furthest 
east for the genus, extending into southern Manitoba, western Minnesota, west- 
ern Iowa, Kansas, Oklahoma, and western Texas. 

Although Kearney's (1935) revision of the North American species is now 
67 years old, it has held up relatively well, which is remarkable for such a com- 
plex genus. His concepts of the species are still accepted, but perhaps reluc- 
tantly by many flora writers, in the absence of a modern assessment of the all 
the North American species. The only modifications come from the seven new 
taxa that have since been added, five of which are in the Intermountain Region. 
Welsh, in his study of the genus for the Utah Flora (Welsh et al. 1993), added 
four of them, 5. psoraloides S.L. Welsh (1980), S.janeae (S.L, Welsh) S.L. Welsh 
(1980, 1998), 5. moorei (S.L. Welsh) S.L. Welsh (Welsh 1980; Atwood & Welsh, in 

press), and S.grossuiarii/oiiavar./umariensis S.L.Welsh (Welsh & Atwood 2001); 
and a fifth from just south of the Utah border in northwestern Arizona, within 
the Intermountam Region, S.gicrischii N.D. Atwood & S.L Welsh (In press). 
Two other new additions are species from south of the region, S. polychroma La 
Duke (1985) and 5. rejlcxa Fryxell, Valdes-Reyna & Villarreal (1991). 

In my study of the genus for IntermounLain Flora I have found the need to 
expand 5. caespitosa M. E.Jones to accommodate a disjunct population from 
Railroad Valley in Nye County, Nevada, which can be distinguished as a separate 
variety The species is closely related to S.ambigua A. Gray, differing from it most 
significantly in its shorter stature. The geographical ranges of the two varieties 
of S. caespitosa lie to the north of the northern limits of 5. amhigua m Utah and 
eastern Nevada and to the east of its most northern limits in western Nevada 
and eastern California. A historical scenario for how the two varieties evolved 
is not implicit from their geographical setting. The two exist m similar desert- 
valley habitats separated by about 160 km of four major north-south oriented 
mountain ranges alternating with three relatively broad valleys. Their close 
similarity may be indicative of recent evolution from an element that migrated 
northward during a warmer Holocene episode, and their subsequent isolation 
has been long enough (m this interpretation) for some genetic drift to occur 

The two varieties of S. caespitosa can be distinguished from each other and 
from S. amhigua by the characters m the following key. 

. Calyx 13-1 8 mm long; petals 
leaves thickish, grayish-green, 
lapping and concealing or n 

tall; endemic to the 

western Uta 

)h desert 

in southwe: 

stern Millard County 

ern Beaver C 

ountyat 1 

600-2000 m elev< 

ation var. 


X 10-14 mm long;F 

mm long; 


6-9 mm long; 

jre leaves relatively! 




It with 

305ed; plant 

c to Railroad 

>y in northeastern N 

ye County N 


1400-1600 r 

Ti elev, 

ation var.' 


ilcea caespitosa 



Perennial herb, 0.3-1.6(-2.5) dm tall, arising from a woody, branched caudex 
surmounting a taproot; herbage grayish-green with a dense pubescence or pale 
greenish with a moderately dense pubescence, the hairs stellate, the rays of the 
hairs spreading in several different directions; stems few to several, ascending 

or erect, unbranched; leaves all caulme, petiolate, the petioles l-3.5(-7) cm long, 
the blades of the leaves at midstem larger than the lower, 1.5-4,5(-5) cm long 
and about as wide, deltate, ovate, or suborbicular, the base cordate, truncate, or 
broadly cuneate, the margin coarsel y crenate or dentate-crenate, and sometimes 
obscurely 34obed, palmately veined, the veins prominent beneath and slightly 
channeled above; stipules filiform or linear; inflorescence few-flowered, the lower 
flowers racemose or solitary in leaf axils, the upper ones often in thyrsoid clus- 
ters, the rachis, pedicels, and calyces pubescent as below; pedicels 3.5-13 mm 
long, divaricately ascending, with stipular bracts at the base; involucral bracts 
filiform, pale brownish; calyx 10-18 mm long, the lobes lanceolate to ovate, acute, 
2/3-3/4 of the calyx length; petals 15-23 mm long, the short claw ciliate, the 
blade obovate, reddish-orange (grenadine); staminal column 3.5-9 mm long, 
stellate pubescent, bearing anthers at the apex; styles with capitate stigmas; 
schizocarp of 11-14 mericarps forming a ring 7.5-8.5 mm in diam., each mericarp 
2-seeded, 3-5.5 mm high, 2.2-4.6 mm wide, rounded dorsally and rounded to 
obtuse apically, coarsely reticulate on the sides of the indehiscent lower 1/3 
and smooth-sided on the upper 2/3, pubescent on back; seeds 1.9-2.3 mm long, 
reniform, dark brown to black, minutely puberulent in patches. 

Sphaeralcea caespitosa M.E. Jones var. caespitosa (Fig. lA-D). Sphaeralcea 
caespitosa M.E.Jones, Contr. W. Bot. 12:4. 1908. Type: UTAH. Beaver Co.: "Wa Wa ..., 
west of Frisco, at Dry Station, ... on very poor volcanic soil covering lava with a 
shallow coat, and m crevices of the rocks where there is a little soil," 6000 ft, Bea- 
ver Co., Utah, 25 Jun 1906, M.E.Jones s.n. (holotype: POM!; isotypes: NY!, US!) 
Perennial herb 0.3-1.7 dm tall; herbage grayish-green, densely pubescent, the 
stellate hairs on mature leaves usually overlapping and concealing the leaf sur- 
face; leaves with petioles 1-3.3 (-5) cm long, the blades thick, 1.5-4.5 cm long 
and about as wide, deltate, ovate, or suborbicular, the base truncate to broadly 
cuneate, the margin coarsely crenate and sometimes obscurely 3-lobed; calyx 
13-18 mm long; petals 16-23 mm long; staminal column 3.5-6 mm long; 
mericarps each 4.5-5.5 mm high and 3.2-4.6 mm wide, rounded apically 

Habitaf.-Shallow, gravelly alluvial soils, mainly from the Sevy Dolomite 
Formation (Welsh 1993), and also from calcareous formations, often growing 
in shadscale, rabbitbrush, or matchweed associations, ranging from (1400) 1600 
to 2000 m elevation. 

Distribution.- West-central Utah in the valleys and foothills of southwest- 
ern Millard County and adjacent Beaver County, in and around the Desert Range 
Experimental Station (Fig. 2). 

Phenology.— Flowering from late May to June, fruiting from mid June to 


^xr^l?| '"^^ 

DSp/jflera/ceflfflwp/fosaME Jones var.cflesp/fos(7.A.Habit,B. Leaves (showing variation),C.Calyx and bractlets, 
irp E-H 5 caespitosa var. williamsiae N.H. Holmgren. E Habit, F. Leaves, G. Calyx and bractlets, H. Mericarp. 
ons by Bobbi Angell (all at NY), based on:A,«67,B.{leftto uqhi) R.C.Holmgren 265, B.Maguire 
L Welsh & M. Chotterley 19530, D. R.C Holmgren 265, E, G. 5.L Welsh 20579, F, H . K.H. Thome & B.T. Welsh 957. 

calyce breviore. 10-14 mm (non 13-18 mm) longa diflert. 

Perennial herb 0.7-2.5 dm tall; herbage greenish, moderately pubescent; leaves 
with petioles 1.5-3.7(-7) cm long, the blades relatively thm, 1.5-3.5(-5) cm long, 
1.5-3(-4) cm wide, deltate or ovate, the base cordate to truncate, the margin 
coarsely toothed or dentate-crenate and sometimes obscurely 3-lobed; calyx 10- 
14 mm long; petals 15-20 mm long; staminal column 6-9 mm long; mericarps 
3-5.5 mm high, 2.2-3.5 mm wide, obtuse apically 

Habitat.-Shallow gravelly soils on alluvium and valley fill, m greasewood 
and shadscale communities, growing with Artemisia spinescens, Atriplex 

caespitosa and 5. caeipitosa ve 

americana, ranging in elevation from 1400 to 1600 m elevation. 

Distrit)ution.— Railroad Valley, northeastern Nye County, Nevada. 

Phenolog}/.— Flowering from May to June, fruiting from mid May to July. 

Etymology. -\i gives me special pleasure to name this lovely desert mallow 
in honor of Margaret Jensen Williams (1917-2000). Sphaeralcea ranked among 
the top of the many native Nevada genera for which Margaret had great fond- 
ness, and it is apparent from her many collections of this taxon that she re- 
garded it as being particularly special. As seen by the tributes to her (Tiehm 
2000), Margaret touched many lives. 

Her sense of adventure lured her to many parts of the world, but she had a 
special bond with the Great Basin, especially Nevada. She always lived in Ne- 
vada, born in Gardnerville to ranchers,John and Wilhelmina Jensen. She earned 
her bachelor's degree at the University of Nevada, Reno, in chemistry and a 
master's degree in mathematics. She married her Chemistry Professor, Loring 
Williams, and together raised to two children. She was a math instructor at the 

University until her appomti 
rules. From 1959 until retirer 
Reno elementary school. 

At home, in the early 1950s, she began to concentrate her enormous energy 
on gardening, at first growing the standard nursery-trade plants. As time passed, 
she became more and more sophisticated in her choice of garden subjects, es- 
pecially alpine rock garden plants. It was not long before she was an active 
member of American, English, and Scottish rock garden societies. Her interests 
in the Nevada flora were spurred on by the writings in the society journals of 
Dwight Ripley and Rupert Barneby on their botanical excursions throughout 
the Great Basin. This deepening interest led her to take classes in plant tax- 
onomy and field botany. At first she was just collecting seeds for exchange with 
rock garden friends in the U.S. and abroad. Then in the early 1970s, John Tho- 
mas Howell of the California Academy of Sciences inspired her to collect plant 

In 1975 she founded the Northern Nevada Native Plant Society, which she 
presided over for several years, first as President, then as Executive Director Her 
charisma, strong will, passion for plants, and contagious enthusiasm drew in a 
large and active membership, and the Society continues to thrive. 

Her legacy lives on through the many research collections filed through- 
out many herbaria. Her collections filed at the New York Botanical Garden are 
especially valuable in our research on the Intermountain Flora (Cronquist et al. 
1972-1997). She was involved m one way or another in collecting 12 specimens 
that are now types for species or varieties. Where she was not listed as the prin- 
cipal collector, it can be assumed that Margaret was the one who enabled the 
collection to be made. This attractive new variety joins Polyctenium williamsiae 
Rollins, Eriogonum ovalijolium van williamsiae Reveal, and Astragalus 
convallarius var. margaretae Barneby as honorific eponyms. 

On a personal note, my wife Pat and I will always treasure our many memo- 
ries of Margaret. She supported our Intermountain Flora research m so many 
ways, from providing food and lodging in her home, transporting us to and from 
the airport and the RENO herbarium, to being a prompt and delightful corre- 

Parat^ PEb Nevada Nye Co all in Railroad Valley low volcanic hills 1 mi NE of Currant 23 May 1982, 
RC Barneby 17821 (NYO 20 mi SW of Currant 14 May 1941 A Eastwood &JT Howell 9426 (US!, 
UTCi) TRNRiiFS2l(NVvl/4) 49001t 4 Jun 199] L Gwxu sn (RFNOO 50 2 km road distance S of 

8N R57hbl 4800 It 12 Ma\ 1987 / Knight 1557 
aguue&AH Holmg> en 25128 WV UTO) US Hw 
Z 94320 (BRYi) Cherry Creek Road 1 6 mi to a ^m 

road, T5N R56E S22, 5200 ft, 2 Jun 1980, K.K Thome & B.T. Welsh 942 (BRY!); 1 mi NE of Currant, 
white knoll on N side of road, TION R58E S9. 5250 ft, 3 Jun 1980, KM. Thorne & B.T. Welsh 957 (BRY!, 
NY!); ca. 22 ml SW of Currant, T8N R56E SI, 4800 ft, 1 Jul 1980, B.I Welsh, KH. Thorne &5.L Welsh 
413 (BRY!, RENO!); 1 mi NE of Currant, TION R58E S9 (NW/NW), 5300 ft, Sarcobatus-Hilaria- 
Chrysothamnus community, gravelly substrate, 23 May 1981, S.L Welsh 20579 (BRY!, NY!, RENO!); 
near Hwy 6, SW of Lockes, T8N R55E S21, 5300 ft, 20 Jun 1980, M.J. Williams 80-164-1 (NY!, RENO!); 
ca. 1 mi NE of Currant, near Hwy 6, TION R58E S4, 5300 ft, 29 Jun 1980, M.J. Williams 80-166-1 (RENO!); 
3.4 mi SW of Lockes, NW of Hwy 6, T8N R55E S19, 5300 ft, 9 May 1980, MJ Williams.S. Cochrane & 
A. Tieh m 80-8-3 (RENO!); 3.5 mi SW of Lockes on U.S. Hwy 6, then NW for 1.4 mi, T8N R54E S25, 5350 
ft, 9 May 1980, MJ. Williams, S. Cochrane & A. Tiehm 80-9-3 (RENO!); near U.S. Hwy 6, 0.4 mi NE of 
Currant, TION R58ES4, 5150 ft, 26 May 1979, MJ. Williams&J.S.Holland 79-54-7 (BRY!, RENO!); U.S. 
Hwy 6, near Mile 119, Nof Currant. TION R58ES4, 5300ft, 23 May 1981, MJ.WilHams& A. TiehmSJ- 
J8- J (RENO!); S of Lockes, 1.6 mi S of U.S. Hwy 6, T8N R55E S27, 4770 ft, 23 May 1981, MJ Williams& 
A. Tiehm 81-19-7 (RENO!); ca. 3 mi SW of Lockes, T8N R55E S30, 5300 ft, 28 May 1978, MJ. Williams 
& M. WUUams 78-75-1 (RENO!, UTGj. 

My deepest gratitude goes to Patricia K. Holmgren for her careful reading of the 
manuscript, assistance in field work, dealing with collecting permits, and in 
procuring loans. I am grateful to Arnold Tiehm for his prompt and informative 
responses to my queries, to Bobbi Angell for her expert illustrations in Fig. 1, 
and to Patricia Eckel for her rendering of the Latin diagnosis, and to Paul Fryxell 
and Steven Hill for their thoughtful and constructive reviews. I thank the cura- 
tors of the herbaria at BRY, RENO, US, UT. and UTC for being gracious hosts in 
their herbaria and, in some instances, for lending me their specimens. The re- 
search for this paper was in conjunction with my ongoing research on the In- 
termountain Flora, which has, during my research on the Malvaceae, been sup- 
ported by The New York Botanical Garden with additional funding from the 
Mary Flagler Gary Gharitable Trust, the George S. and Dolores Dore Eccles Foun- 
dation, the Harriet Ford Dickenson Foundation, The Andrew W. Mellon Foun- 
dation, and Mrs. Nicholas J. Sakellariadis. 


ATwooD,N.D.and S.L.Welsh (In press). Sp/ioera/ceo g/er/sc/i/7 N.D.Atwood 

& S.L.Welsh. 

Cronquist, a. et al. 1 972-1 997. Intermountain flora: vascular plants of th 

e Intermountain 

West, U.S.A. vols. 1 , 3a, 3b, 4, 5, 6. New York Bot. Garden Press. 

Fryxell, P. A., J. Valdes Reyna, and J.A. Villarreai Q. 1 991 . A new specie 

5 of Sphaeralcea 

(Malvaceae) from Coahuila, Mexico. Southw. Naturalist 36:358-360. 

Kearney,T H. 1 935. The North American species of Sphaeralcea subgeni 

js Eu sphaeralcea. 


La Duke, J. C. 1 985. A new species of Sphaeralcea (Malvaceae). Southw. N 

aturalist 30:433- 

1 flora. Ed. 2. Print Services, Brigham Young Univ 
S.L. 1998. An undescribed Astragalus (Legumin 
section of the genus,and validation of the comb 
sh. Great Basin Naturalist 58:386-389. 


Manager, PNG National Forest 

PNG National Forest Service 

P.O. Box 5055, Boroko,NCD 




Of all the exploratory surveys conducted in New Guinea during the last 100 
years, the German Kaiserm-Augusta Expedition of 1912-13 stands out as being 
particularly significant. During that expedition, botanist Carl Ledermann trav- 
eled extensively through the south Sepik basin with a multidisciplinary con- 
tingent, eventually collecting a total of 6,639 angiosperm numbers over a pe- 
riod of 18-19 months. Originating at an early point in the exploration of the 
Papuasian flora, a disproportionate number of Ledermann's collections were 
designated as type specimens. Many of his localities have never been revisited. 
The unfortunate circumstances of Ledermann's labors are an enormous obstacle 
to modern scientific inquiry, for the main German sets were destroyed in the 

1943 lire at Berlin Herbarium, effectively erasing the primary basis for the iden- 
tification of numerous plant species. Veldkamp et al. (1988) have provided an 
informative discussion of the relevant issues surroundmg the loss of such a criti- 
cal botanical corpus. 

One of the most productive parts of the Kaiserin-Augusta itinerary was 
the series of collecting trips made by Ledermann to the April River (Aprilfluss). 
The Aprilfluss yielded a bountiful harvest of new species, many of which have 
unfortunately proven impossible to evaluate without the original collections. 
With modern assessment indicating the apparent existence of a zone ol f loris- 
tic endemism centered on the Hunstein-Apnl region, it has become particu- 
larly desirable to revisit the classical localities and secure new material. Such 
efforts would enable neotypification of many species whose types were lost at 
Berlin (e.g., Tapeinochilos hollrungii K. Schum.; cf. Gideon 1998). Although a 
list of plants requiring new collections is reportedly being compiled (Veldkamp 

The first serious reexamination of Ledermann s sites occurred m 1966, with 
the CSIRO-sponsored expedition to Mt Hunstein by R. Hoogland and L. Cra- 
ven. In 1989 a multi-institutional survey returned to the Hunstein Range under 
the auspices of the National Geographic Society, obtaining 1,200 more botani- 
cal numbers (cf . Bakker 1994 for a general account). The first author, one of five 
botanists on that expedition, independently returned to the Hunstein summit 
in July-August 1990 and to the nearby Waskuk Hills m August 1994 andjune 
1995. Additional collections from the AprihSitipa drainages were made in Sep- 
tember 1990, and from the April River serpentine zone on the south side of Mt 
Hunstein, m July 1995 (Fig. 1). The most recent trip to the subdistrict occurred 
in August 2001 (cf . Takeuchi & Golman 2002; Takeuchi & Renner 2002), from 
which much of the information in the following narrative has been taken. 

The indigenous population of the Hunstein subdistrict is spread diffusely 
through the region in small clan-based villages. Ambunti is the largest settle- 
ment; with a district headquarters, tradestores, an aid post, two schools, several 
churches, and a grass airstrip. Further upriver and within the April catchment, 
the average settlements are much smaller, often consisting of a single extended 
family. At Garuka (Waskuk Hills) for example, the total adult population was 
exactly ten during the time of the 1995 survey. The larger villages like Baglam 
and Melawei have about a hundred inhabitants. 

In the Sepik region, river-borne traffic is the major means of communica- 
tion, trade, and transport. Most of the population are thus concentrated near 
the water. This also affords the people convenient access to fish and saksak (made 
from the starchy pith of the Metroxylon palm) which together comprise the 
bulk of the average villager's diet. A limited amount of subsistence slash and 

burn gardening provides dietary variety. Sweet potatoes, tare, bananas, and 
coconuts constitute the mam crops, but plantings are generally not a principal 
source of village sustenance. For the most part, the April River populace are 
semi-nomadic hunter-gatherers. 

Like the Sepik River, the April is a meandering silt-laden waterway which 
often changes course after severe floods. A recently introduced fish from South 
America (the pacu, erroneously identified as a piranha in the local media) has 
now entered the April drainage and is rapidly becoming the most commonly 
caught fish for consumption. Villagers report that the appearance and prolifera- 
tion of the pacu have been accompanied by disappearance of many native fish 
species, particularly the endemic species of Sepik catfish (Anus spp., cf. Allen 
1991). The pacu has reportedly been involved in several fatal attacks on bathers, 
but such anecdotal reports are often exaggerated and their veracity is uncertain. 

Socioeconomic development in the April region is virtually nonexistent. 
The local economy is primarily subsistence in nature, and cash incomes are 
opportunistic. Money is occasionally earned from the sale of wood carvings, 
crocodile skins, birdwing butterflies, cacao, and from seasonal sources in the 

volvingGyrinops/ederman nil Domke(Thymelaeaceae), with Indonesian buy- 
ers paying up to PNGK 800-1,000 (USD 265-335) for each kg of black wood. 
The arborescent species is endemic to the upper Sepik (Ding Hou 1960), and 
has become a significant income source for landowners due to consumer de- 
mand for the wood's pleasing fragrance, a development reminiscent of the san- 
dalwood trade. Known as 'may-hasei' in the Parenemo language, G. Udermannii 
is common on the southwest side of Kamelsrucken and is also distributed on 
slopes facing the April River, particularly near Etappenberg (Takeuchi & Gol- 
man 2002). For many decades the species was known only by Ledermann s type 
gathering from the May River, but current indications suggest the tree prob- 
ably ranges even into West Papua (ibid.). 

In spite of income from such incidental sources, by Western standards the 
overall status of the people would be regarded as impoverished. Due to the in- 
accessibility of health services, mortality rates are high, especially from ma- 
laria and infections, and the local inhabitants frequently seek palliatives and 
remedies from whatever natural resources are at hand. Skm diseases like ring- 
worm and scabies often affect entire villages. 

April River culture is an improbable assortment of incongruities and ap- 
parent contradictions. Villagers have adopted western standards of dress, yet 
still retain their traditional lifestyle and customs. Many of the residents speak 
standard English, often freely intermixed with their tribal tokples (Baihenemo, 
Parenemo, Waskukil) in ad hoc combination. Although Christianity in its many 
variants is dominant throughout the region, belief in sorcery is closely inter- 
woven through the Christian ethic. Sudden deaths are nearly always attributed 

to black magic or to secret murders (sanguma) facilitated by plant poisons. 
'Ukhop pok' (Anodendron oblongifolium Hemsl.; Apocynaceae) is the most com- 
monly mentioned agent in sanguma killings. There is also a widespread con- 
viction in the existence of spirits (masalai), believed to pervade all aspects of 
the natural world and which form an inseparable part of the villagers' meta- 
physical world-view. 

The population growth rate for East Sepik Province is currently 2.9% per 
annum, a relatively high figure, but below the Mamose^ regional average of 3.3% 
(National Statistical Office 2001). Most of the provincial population is concen- 
trated in the built-up areas near the coast, and the interior sections such as April 
River consist primarily of wilderness habitat. Due to the low population densi- 
ties m this part of PNG, there are no subsistence-based threats to the environ- 
ment. However, because the entire region along the April-Salumei is one of PNG's 
most economically depressed areas, landowners are understandably attracted by 
the inducements offered by logging and mining interests. A major portion of the 
district has been proposed as a timber concessional area under the Forest Man- 
agement Agreement protocols (cf. Papua New Guinea Forest Authority 1998). 
Reconciling the conflicting desire for socioeconomic development on one hand, 
with conservation concerns on the other, will be a major challenge for commu- 
nity planners. Many of the relevant issues are presently being addressed by the 
World Wildlife Fund ( W WF), through the establishment of an ICAD (Integrated 
Conservation and Development) program in the Hunstein area and the related 
demarcation of a Wildlife Management Area (WMA; Fig. 1). The Village Devel- 
opment Trust is also contemplating start-up of a comparable program in the 
Bugabugi territory and the Waskuk Hills (Aung & Kisokau pers. comm.). In the 
past, investigators moved freely throughout the area, subject only to approval 
from the customary landowners. With the recent and impending developments, 
this is likely to change. Intervention by nongovernment organizations (NGOs) 
invariably results in new requirements which can complicate scientific access 
and research. The complications are offset however, by the real and perceived 
benefits which NGO association can deliver to affected communities. 

Land ownership and the subsequent control of access to forest resources 
are traditional and patrilineal. Nearly all territories fall under clan jurisdiction, 
so discretion must be exercised during plant collecting m order not to trespass 
on land unapproved for gathering. A gratuity paid to the landowner(s) is often 
necessary to ensure approval for plant collecting. 

In contrast to the many obstacles which confronted the first explorers, access to 
the upper Sepik has improved to such an extent that the region is now a major 

cLOtouiist distmuion Lhitil icecnth o 

ecan gomg lomist MsstK m kL iLgu 

UiU scludukdluxuu ciuistsasfatup 

,ttL-imasAmbunti Akhough tiaxeleis 

cm still txpcet numuousdcla\saiisn> 

^ I torn conditions Lndtmie to develop 

m^eountiRs cntn totlu Apiil basm b 

as ken eonsideiabh cased b> the new 

aiiliclds at Bugabugi ^nd NiksUv ' Botl 

1 mhelds ncsmrllgiassstiipsmam 

tamed by New Tribes Mission and em 

iLccpt onh the smgk engine Cessna 

206 with maximmn lift eapaeit) ol S passe iigets oi 400 kg Otherwise access 

can also be made via the an field at Am 

bunti which can accommodate laiger 

twin engmedlslandeis capable ol cam 

mglQpasscngcisoi 1 600 kgs of freight 

(Missionary An Fello\^shipJ Fiom Amb 

unti iMsitoi would oidmaril) pioceed 

to the Apiil Ruei w ith dugout c moes 

powcicd b\ \amaha outboaid motors 

(15 30 01 40 hpmotoi sate the usual lat 

ingsj The lucl consumption of a 30 hp 

motoi on a loundttip bet\\een Ambuni 

ti and Bugabugi will be at least 140 h 

teis In the uppci '^epik fuel is problematic being eithei unavailable or pi ohibi 

The buttress iidges m the vicmit) of Bugabugi (Pmape Natawe and 
Okahsa) lead dnectly to Mt Hunstem and lepresent the easiest path to the top 
In 1989 the National Geogiaphic expedition appioached the summit fiom the 
north which is actually the longest and most logisticalh difficult land route 
kedermann s two ti ips to Hunstemspitzc had come the same w av fiom Ambunti 
and thiough Wasui Wagu kagoon (cf Steenis Kiuscman 1950 317-18) The 
cnri \ \ n W i^u w \s similail) taken b> Hoogland tnd Cia\en m 1966 (Ciaven 
pels comm J 1 he quickest access howe\ei is to lollow the south descending 
iidgcshomthc Apnl Ri\er piefeiabh starting at Natawe (Fig 2) Withthenewly 
opeiational an held U Bugabugi the lattei loute can now be used to reach the 
Hunstem ciestlme withm two da>s oi departure fiom the provincial seat at 
Wewak The south side appioaeh piovides the added advantage of passage 
through unexploied ulttabasic habitats 

Field conditions in the Apnl drainage are oneious The heat and humidity 
aie stultifying dmmg dry periods and the wet season is accompanied by toi 
rential downpouis aftei which the ri\ei is hazardous at man) places along its 
course Piolonged \ isits uc a health nsk The ph\sicil difhculties are aggra- 
vated by the ibsence ol m\ set\ice mil istiuctuic thioughout the legion ex 
ceptfoi basic he ihties umissonu\ stuions 0\ei muchol the \eai the mos 
quito and leech popul itions iic exceptionalh tioublesomc e\en by the 
standaids ol an alkuial tiopieal en\uonment When it is not taming horse- 
flies are anno)ingl\ plentiful especialh m areas where pigs ate present m the 
forest Poisonous snikcs ha\e been lesponsible for a substantial number of 
deaths m the \ illage communities 



I ° 5 1- - - - 

The vegetation along the April River consists of three principal formations. The 
swamp forest (with several facies including levee forest, meander scrolls, swales, 
etc.) IS the dommant community on the flood plain. Typically waterlogged or 
mundated during rainy periods, the swampy habitat is dominated by Metroxylon 
sagu Rottb., Campnosperma hrevipetiolata Volk., or herbaceous vegetation (espe- 
cially Saccharum rohustum Brandes &Jeswiet ex Grassl; cf . Warner & Grassl 1958). 
On better drained alluvium the forest is more structurally developed, giving rise 
to a second association with many macrophyllous taxa (e.g., Artocarpus commu- 
nis JR. & G. Forst., Caryota rumphiana Mart., Pangium edule Reinw., Sterculia 
macrophylla Vent., and Nauclea orientalis L). These communities have charac- 
teristically high proportions of robust climbers along the edges, particularly Ca- 
lamus, Korthalsia, and to a lesser extent Freycinetia, and the interior sections 
often support a luxuriant shrub layer. On the Hammermaster and Saunders 
(1995a, b) forest classification system, the riverine vegetation is described as an 
open forest on low elevation plains and fans, or as swamp woodland (forest code 
To' and 'Ws' on overlays SB 54-4 and SB 54-3 to the Ambunti and Mianmin 
1:250,000 Australian Survey Corps topographic sheets). Paijmans (1976) provides 
a concise overview of the vegetation in such environments. 

At several places along its course, the April River skirts the base of serpen- 
tine/ultramafic foothills and there is a sharp f loristic break as the swamp and 
alluvial associations are replaced by foothill forest. The contrasting features of 
the forest facies on ultramafics are: 1) a pronounced reduction in climbers (e.g.. 
Calamus is virtually absent), 2) a more varied canopy structure, including a 
noticeable reduction m large-leaved taxa, 3) increased floristic richness, most 
apparent m an understory which is not subject to the periodic flooding charac- 
teristic of the alluvial flats, 4) a 'clean' visual aspect to the community with 
good visibility through the forest because of the absence of climbers and com- 
parative lack of epiphytic growth on tree boles, 5) a very obvious change in 
taxonomic composition, many taxa being common on the ultramafic slopes 
but absent from immediately adjacent alluvial forest. When observed from the 

a montane forest, except for the paucity of epiphytes. Based on air photo inter- 
pretation, the April foothill zone is a medmm-crowned low altitude upland for- 
est (code 'Hm' in Hammermaster & Saunders 1995a), a vegetation type which 
is probably Papuasia's richest forest structural category (Louman & Nicholls 
1995). Although an unpublished report (Sohmer et al. 1991) provides a brief 
botanical description of the Hunstein Range, there is still nothing in the way of 
a comprehensive account for the April basin. 

The forest on ultramafic foothills is easily accessed near BugabugP (Pmape, 
Natawe and Okahsa: in sequential order upriver), though the best section is situ- 

ated at the base of the Hunstein Range between Gahom and Okahsa, close to 
the junction of the April and Sitipa rivers (Figs. 3, 4). A conspicuous feature of 
the hill community is the presence of Agathis labillardieri Warb. emergents on 
the ridge buttresses and crestlines. Agathis attains heights exceeding 50 m, and 
the April provenances can be seen from the river towering over the surround- 
ing canopy. The populations are of commercial significance, so not unexpect- 
edly, has attracted the attention of logging and ecoforestry operators. Due to 
the steep and rocky slopes, Bugabugi villagers do not establish subsistence gar- 
dens in the foothill habitat, and the slopes are thus remarkably free of human 
disturbance. However, many understories near the river are being disrupted by 
domestic and wild pigs. Large serai gaps are also scattered through the forest as 
a result of natural disturbances from landslides, lightning strikes, and 
windthrows. For a lowland environment, the frequency of lightning-induced 
gaps is surprisingly high, an indication of the intense electrical and thunder- 
storm activity in the Hunstein Range. 

During the 1998 el Niiio disturbance, the Hunstein subdistrict and the en- 
tire basin to the south was severely affected by drought. The Sepik became dry 
in many sections even though it represents the largest catchment system among 
PNG's rivers. At Yambon gate, the Sepik current normally accelerates through 
a narrow channel renowned for treacherous whirlpools (cf. Townsend 1968), 
yet villagers were able to walk across to the opposite bank during the 1998 
drought. In spite of such conditions, and unlike many other forested sites in 
PNG, April River environments did not experience any fires during the last dis- 
turbance. However a large tract downstream of Bitara has been recently flat- 
tened by cyclonic-force winds, and the forest canopy in a swath ca. 1 km wide 
completely removed. From current events, it is apparent that vegetational his- 
tories in this basin are at least partly determined by catastrophic events (cf. 
Johns 1986). 

Since the time of its initial exploration, the Hunstein subdistrict has been 
judged a particularly significant hotspot for diversification and endemism. The 
biodiversity status of the area has assumed legendary proportions, even though 
the perception of unusual richness can be regarded as a natural outcome of 
physical and historical considerations. 

The geological complexity of the April-Hunstein territory is a major con- 
tributing factor in the diversification of its flora. Alluvium, colluvium, sand- 
stone, shale, mafic/ultramafic metamorphics, igneous, clay and coralline lime 


Stone substrates occur throughout the area m patches of varying size (Davies 
& Hutchison 1980a, b). There is thus a highly fragmented pattern of contrast- 
ing substrates over the terrain, reflecting the manifold geological origins of the 
present territory and capable ot supportmg specialist taxa in a complex mo- 
saic. Supenm posed on the toundation of edaphic environments is an excep- 
tional annual ramfall estimated at 7,000-9,000 mm (ibid.). High rainfalls are 
directly related to floristic diversification (Gentry 1988) so perhumid condi- 
tions m the April basm have probably played a major role in the development of 
a speciose flora. When the dynamic fragmentation of the ecosystem is then 
considered, over presently observable scales ranging Irom hghtning strikes to 

species can be accommodated in ^ide b\ side lai^hion over relatively small dis- 


Pigram & Davies 1987). The Hunstein flora is probably best understood as an 
extension of the West Papuan flora. During the time of Davies and Hutchison 
(1980b) the Hunstein- April-Salumei petrology was already known to extend 

leterogeneity ol April 

habitats absonccd.^ 

. to be vle^^'ed m terms of 

■regarding correlation 

s between plant du 

itributions and phases of 

retionlcl Welzenl997. 


t s reputation as a zone of 


d. 1988) is due to It: 

5 easternmost position in 

complex which consi 

tructed most of n. 

Drthern West Papua (cf. 

westwards rather than eastwards. If the division between PNG and West Papua 
were to be based on geophysical rather than pohtical criteria, the border would 
be at Apnl-Salumei rather than Sandaun. 

Based on the relationship to tectonics, future distributional records of 
Hunstein/April endemics will occur to the west, as the Papuan affinity clari- 
fies through further exploration of West New Guinea (Indonesian West Papua). 
At present, the Indonesian side is even more inadequately surveyed than PNG, 
and the historical disparity in documentation between the two halves is partly 
responsible for the perception of unusually high endemism in the Hunstein/ 
April region. The newly discerned disjunction of Myrmephytum in the Vogelkop 
and Hunstein areas (Jebb pers. comm.), of Faika in West Papua and the April 
River (Takeuchi & Renner 2002), and the probable occurrence of Gyrinops 
ledermannii injayapura (Takeuchi & Golman 2002) are suggestive of the phy- 
togeographic connections, and of the effect which historical patterns m f loris- 
tic documentation have on assumed distributions (Figs. 5, 6). 

Nearly 90 years after the Kaiserin Augusta Expedition, many of the environ- 
ments along the April River are still unexplored. The localities at the 
Expedition's farthest penetration into the south basin, near the headwaters of 
the Bamali tributary, have never been revisited. Of these, Lordberg is probably 
the most intriguing, but any effort focused on the Divide south of Niksak will 
undoubtedly reap huge rewards. The recent discoveries from Bugabugi exem- 
plify the possibilities. Although the 6,639 collections taken by Ledermann rep- 
resent an unprecedented achievement for his time, even a personal effort of that 
intensity cannot provide the desired level of sampling saturation for such a f lo- 
ristically rich region. A collections program delivering at least 10,000 new num- 
bers is much needed. 

The factors militating in favor of future survey success in the April drain- 
age can be recanted; 1) forest communities in the region are in an exceptional 
state of preservation, 2) traditional lifestyles and ethnobotanical competences 
are still retained by local villagers, who are thus a valuable source of the folk- 
lore knowledge being rapidly lost m other districts, 3) improved site access to 
the interior basin will enable adoption of cost-effective logistical schedules, 4) 
research conducted by various investigators since 1989, has created local aware- 
ness of the work done by scientists, which will facilitate future inventories. 

No collections effort conducted under normal constraints, can achieve 100% 
coverage of a targeted flora. There is always something which will be overlooked 
on account of rarity, phenology, localized distribution, logistical limitations, 
etc. Even the collections obtained by Ledermann during his Sepik travels are 
not such a superlative effort as may appear at first glance. The 6,600+ numbers 
from the Kaiserin Augusta survey amount to approximately 400 numbers per 


month on a time-averaged basis (excluding Ledermann's 2 months of mactiv- 
ity due to sickness). By the productivity standards of contemporary collectors 
working under rapid assessment protocols, this is a below-average result. The 
significance of the Ledermann collections are another matter altogether how- 
ever. And in fairness, the logistical environment of his time was undoubtedly 
much more limiting than now. The difficulties of prolonged expeditionary 
schedules also should not be underestimated. Anyone with knowledge of 
Papuasian forest environments can appreciate the hardships in being exposed 
to tropical conditions for the duration which Ledermann was subjected to. 

Some indication of the spottiness of Ledermann s sampling coverage can 
be gauged by results from the 1966 and 1989 expeditions to Mt. Hunstein. Sev- 
eral scores of new taxa have been described from the later visits, so clearly the 
original effort was not very comprehensive. To what extent does that situation 
apply to even more remote sites, such as Niksak-Lordberg? On the basis of ac- 
cumulating evidence showing that the richest floristic interval in Papuasia is 
the premontane zone (Takeuchi & Golman 2001), the best environments within 
the April region are probably the ones at the Central Divide, and these have 
never been seen since Ledermann's time. The diversity potential of this flora 
has yet to be plumbed. 

We thank parataxonomist A. Towati of Ambunti, the Glei villagers of Waskuk, 
and the Galiase and Kokomo clans, for their assistance during our visits to the 
upper Sepik. Hitofumi Abe (Ecosystem Research Group, University of Western 
Australia) wrote the Japanese translation. Figure 1 was adapted from Geographic 
Information System downloads from the PNG Forest Authority Reviewers 
David Middleton (Harvard University Herbaria) and an anonymous referee, 
made helpful comments on the draft. 

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Institute Publ, 9:1 -268. 
Davies, H.L and D.S. Hutchison. 1 980a. Papua New Guinea 1 :250,000 geological series map. 

Ambunti, sheet SB 54-4, Geological Survey of Papua New Guinea, Dept. Minerals and 

Davies, H.L. and D.S. Hutchison. 1980b. Explanatory notes to accompany the Ambunti 

1:250,000 geological map. Report 80/1. Geological Survey of Papua New Guinea, Dept. 

Minerals and Energy. Pp 1 -25. 
Ding Hou. 1 960.Thymelaeaceae, Flora Malesiana ser. 1, 6 (1 ):1 -48. 
Gentry, A. 1 988. Changes in plant community diversity and floristic composition on envi- 
ronmental and geographical gradients. Ann. Miss. Bot. Card. 75: 1 -34. 
Gideon, O.G. 1 998. Systematics and evolution of the genus Tapeinochilos Miq. (Costaceae- 

Zingiberales). Ph.D. thesis. James Cook University of North Queensland, Australia. 
Hammermaster, E.T.and J.C.Saunders. 1995a. Forest resources and vegetation mapping of 

Papua New Guinea. PNGRIS Publ. 4, Canberra, CSIRO and AIDAB. 
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Papua New Guinea. 1:250,000 vegetation map overlays separately issued as working 

copies to PNGRIS Publ. 4, Canberra, CSIRO and AIDAB. 

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eds. Papua New Guinea country study on biological diversity. Colorcraft Ltd, Hong 

Kong.Pp 155-167. 
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to the year 1 950. Fl. Male. ser. 1, 1 :1 -639. 
Takeuchi, W. and M.Golman. 2002. The identity of eaglewood (Gyr/nops, Thymelaeaceae) 

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Guinea. Fl. Male. Bull. 13:54-55. 

46. Pacific Publications, Sydney 1 -271 . 
VELDKAMP,J.R,W.ViNK,and D.G.Frodin. 1988.XI.Ledermann'sand some other German loc£ 

ties in Papua New Guinea. Fl. Male. Bull. 10:32-38. 
Warner, J.N. and C.O.Grassl. 1 958.The 1 957 sugar cane expedition to Melanesia. Hawaii 

Planters' Record 55:209-236. 
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M.J.F. Coode, and D.A. Simpson, eds. Plant diversity in Malesia III. Proceedings of t 

Third International Flora Malesiana Symposium 1995.Pp 363-387. 


Curtis J. Hansen^ and Loreen Allphin^ Michael D.Windham 

Department of Botany and Range Science Qarrett Herbanum 

Provo, UT 84602, U.S.A. UniversitvofUtah 

Chromosome studies and ph ylogenet ic analyses of morphology, allozymes, and nrDNA-irS sequel 

I pubescens, and I subnudum var.alpinum. The first two rare taxa yield only diploid (n=12) chron 
some counts; I subnudum van alpinum is consistently tetraploid (n=24). Theksperma suhnud 
'• Phylogenetic ana 

;ely related to T. pubesccns and I cmipiiosum than to I subnudum var. subnudum. Co 
lyses give the strongest support (bootstrap 100%) for the clade of I pubescens, T. caespitosi 

)s: Theksperma, Asteraceae, nrDNA-JTS sequences, phylogenetics, and systematics. 

s de nrDNA ITS paia li u u de rsoInu I is iclaciones taxonomicas dentio'del compl, 

complejo: ThekspcTmacc 

88%-99%) el agrupamiento de T subnudum var 
(bootstrap 100%) para el clado compuesto por T si 

Thdcspcrma Lessing (Asteraceae: Heliantheae, Coreopsidinae) comprises about 
15 species, mostly herbaceous perennials (annuals and subshrubs rare), native 
to south-central and western North America and warm-temperate South 
America (Melchert 1963; Bremer 1994a). The genus is thought to be monophyl- 
etic based on shared characters such as strongly dimorphic mvolucral bracts, 
scarious-margined inner involucral bracts that are connate from 1/5 to 1/2 their 
lengths, opposite leaves, and pappi that are absent or composed of hispid or 
serrulate awns (Bremer 1994a). Previous taxonomic studies of Theksperma have 
focused on morphology (Shinners 1950a, b; Alexander 1955) and cytology 
(Melchert 1963; Greer 1997; Greer & Powell 1999) and were not explicitly phylo- 
genetic. Nevertheless, past work on the genus has identified some species groups 
that may be monophyletic. One of these is the Theksperma subnudum com- 
plex, which is restricted to the Colorado Plateau and areas adjacent to the Rocky 
Mountains in western North America (Fig. 1). This group includes five com- 
monly accepted taxa; T. suhnudum A. Gray var. suhnudum, T. suhnudum A. Gray 
var. alpinum S.L. Welsh, I. pubescens Dorn, T. caespitosum Dorn and T. 
marginatum Rydb. Features used to distinguish members of the I subnudum 
complex are given in Table 1. 

Delimitation of taxa within the Theksperma suhnudum complex has var- 
ied, and at least three classifications have appeared (Table 2). Dorn (1990), who 
discovered and named T. pubescens and T caespitosum, recognized each as dis- 
tinct species. He treated T. marginatum as a separate species and accepted 
alpinum as a variety of I subnudum. In the 2nd edition of A Utah Flora, Welsh 
et al. (1993) classified both T caespitosum and T. subnudum var. alpinum as va- 
rieties of T subnudum; T. marginatum and T. pubescens were not treated because 
the former is not known from Utah and discovery of the latter in Utah post- 
dates publication of the flora. Cronquist et al. (1994) recognized only I 
subnudum and T. pubescens at the species level; T caespitosum and I suhnudum 
var. alpinum were included within T pubescens, and T. marginatum was treated 
as a variety of T. suhnudum. 

These divergent classifications result from differing interpretations of the 
available morphological and cytological information. Additional genetic data 
(e.g., allozyme and DNA) are needed to allow an informed choice between com- 
peting taxonomies. The need to pursue these studies has been heightened re- 
cently by conservation concerns. Theksperma pubescens, T. caespitosum, and I 
subnudum var. alpinum are rare taxa restricted to a few localized populations 
(Fig. 1). All three have been listed as potentially endangered (category 2) by the 
U.S. Fish and Wildlife Service (U.S. Dept. of Interior 1985, 1993). The accumula- 
tion of genetic data is an important step m determining their eligibility for fed- 
eral protection under the Endangered Species Act. 

Our objectives m this study are threefold; 1) to develop baseline genetic data 
for members of the Theksperma subnudum complex, 2) to analyze the data in 

Table 1. Selected mor|)holoqical ecological, and distributional ff 
Taxa Leaf segments Pubescence Heads Rootstoc 

concert with morphology to produce an expUcit phylogenetic hypothesis for 
these taxa, and 3) to compare that hypothesis to existing classifications and 
propose any desirable taxonomic changes. Through this process, we hope to 
shed light on the relationships and possible origin of I suhnudum var a Ipinum, 
the rarest and most enigmatic member of the group. 
Determination of Chromosome Numbers/Ploidy Level 

Ploidy levels were determined for all taxa m the I suhnudum complex using a 
combination of chromosome counts and analyses of allozyme banding pat- 
terns. For chromosomal observations, capitula (m bud) were obtained from field 
populations and fixed m Farmer's solution (3 parts ethanolT part glacial acetic 
acid). Chromosome counts were made from meiotic figures obtained using stan- 
dard squash techniques and acetocarmme staining (Turner & Johnston 1961; 
Strother 1972). Preparations were examined under phase contrast on a Zeiss 
Axioplanl Microscope. Images of chromosome squashes were saved electroni- 
cally using Zeiss Image® software from Carl Zeiss, Inc. 

In populations for which flower buds were unavailable, ploidy level was 
determined by careful examination of allozyme banding intensities across a 
variety of enzymes (Danzmann & Bogart 1982a; Dessauer & Cole 1984; Pryer 
& Haufler 1993). Because allozyme markers are additive, codommant, and 

inherited in a Mendelian fashion, the expressions of alleles at various ploidy 
levels are expected to be proportional to their gene dosages. In our study, chro- 
mosomally documented diploids in Thelesperma always showed balanced band 
patterns at heterozygous loci. If a heterozygote showed unequal band intensi- 
ties of 3:1 (in a monomeric enzyme) or 9:4:1 (in a dimeric enzyme), the particu- 
lar individual always proved to be tetraploid (the only type of polyploid en- 
countered during our study). Thus, chromosomally unknown populations 
could be assigned to a specific ploidy level based on the presence or absence of 
unbalanced heterozygous allozyme patterns. 
Morphological Data 

Our coding of morphological character states is based on observation of ap- 
proximately 300 herbarium sheets from ASC, BRY, GH, MONTU, NMU, NY, RM, 
UX and UTC (herbarium designators follow Holmgren et al. 1990). These data 
were supplemented by information obtained from Melchert (1963), Welsh (1983), 
Dorn (1983, 1990), Jansen et al. (1991), Ryding & Bremer (1992), Welsh et al. (1993), 
Cronquist et al. (1994), and Karis & Ryding (1994). Ingroup taxa included I 
suhnudum var subnudum, T. suhnudumY3iLalpinum,T.puhescens, T. caespitosum, 
and T. marginatum, plus the related species I.JiliJolium (Hook.) A. Gray, T. longipes 
A. Gray, and T. megapotamicum (Spreng.) Kuntze. Bidens has been identified as 
a possible sister genus to Thelesperma (Rydmg & Bremer 1992) and two species 
of that diverse genus, B. cernua L. and B.frondosa L., were chosen as outgroup 
taxa. A total of 16 characters (14 binary and 2 multi-state) was included in the 
morphological analysis (Tables 3, 4). 
nr DNA-ITS Sequence Data 

Samples used in the DNA study are identified by their GenBank accession num- 
bers in Table 5. ITS sequences for the outgroups Bidens cernua and B.frondosa 
were obtained from GenBank; voucher data for these collections can be found 
in Ganders et al. (2000). The ITS sequences for all Thelesperma taxa were ob- 
tained directly by extracting total DNA from the leaf tissue of dried specimens 
using a basic CTAB extraction protocol (Hillis et al. 1996). Two different indi- 
viduals from each taxon were sampled in order to check for intraspecific differ- 
ences. The IT5-1 & 2 and 5.85 regions were amplified using the polymerase chain 

t basally (1). 

lately or ternatelylobed (■ 

13, Cypselae straight (( 
15. Pappus present (0); 

16. St 

ems pubescent (0); 

stems glabrous 






and Thelesperr 

7,a. Chare 






;9Trini 1 

Thelesperma caespitoiur 
Thelesperma filifolium 
Thelesperma longipes 
Thelesperma marginatui 
Thelesperma megapotar 
Thelesperma pubescens 
Thelesperma subnudum 
Thelesperma subnudum 

reaction (PCR) atid two primers m a 1:1 ratio. Primers used m PCR amplification 
and DNA sequencing were 115-4 (White et al. 1990) and a modified sequence of 
lTS-5 (White et al. 1990). The latter, designated IT5-1, has the sequence: GTC CAC 
TGA ACC TTA TCA TTT AG (L. Urbatsch, pers. comm.). The following PCR cy- 
chng protocol was used m amplification: a 95°C hot start for 12 min.; 40 cycles of: 
95°C for 20 sec, 54°C for 30 sec, and 7 JC for 1 mm.; a 4 mm. final extension at 

7 JC; and ending with a hold at 4"C. To confirm base positions, the ITS-1 & 2 
and intervening 5.8S regions of Jheksperma were sequenced in both directions. 
The contigs, or unidirectional sequences, were assembled using Sequencher 
(Gene-Codes, Ann Arbor, MI) and visually checked. Base positions that gave 
equally strong, contrasting signals on both strands were coded as ambiguous 
according to the lUPAC-IUB ambiguity code set included in the program 
Sequencher. All characters were coded as unordered, and gap characters ("-") 
were treated as missing data rather than a fifth character state (Baldwin 1993). 
Allozyme Data 

An electrophoretic survey was performed to analyze allozyme variation at a 
variety of enzyme loci. A total of 765 plants were sampled from 25 natural popu- 
lations (about 30 plants/population) representing all taxa in the Thehsperma 
suhnudum complex, I longipes A. Gray, and one outgroup population of Bidens 
cernua L. (Table 5). Fresh leaf material was collected in the field and kept on ice 
until returning to the laboratory (2-3 days maximum). Tissue was then ground 
using a mortar and pestle in the phosphate-PVP extraction buffer of Soltis et al. 
(1983). Extracts were absorbed onto wicks cut from Whatman 3MM filter pa- 
per and stored at -80°C until electrophoresis. 

A total of 19 enzymes was surveyed electrophoretically for variability. Six 
enzymes, representing 11 putative loci, provided consistent, interpretable re- 
sults. Two buffer systems were used to resolve these enzymes in 11.5% (w/v) 
starch gels. A tris-citrate/borate buffer system (System 6 of Soltis et al. 1983) 
provided good resolution for leucine aminopeptidase (LAP), phospho- 
glucoisomerase (PGI), phosphoglucomutase (PGM), and triosephosphate 
isomerase (TPI). A pH 7.5 modification of the morpholine-citrate buffer system 
of Odrzykoski and Gottlieb (1984) was used to resolve malate dehydrogenase 
(MDH) and shikimate dehydrogenase (SkDH). Staining schedules and proto- 
cols followed Soltis et al. (1983) and Murphy et al. (1996). 

Genotypes were inferred directly from electromorphs observed on the 
stained gels, based on the assumption that enzyme substructure and compart- 
mentalization parallel those observed in other flowering plants (Gottlieb 1981). 
When enzymes showed more than one allozyme locus, the most anodal (fast- 
est migrating) locus was designated number 1, the next fastest number 2, etc. If 
more than one allele was present at a locus, the most anodal allele was desig- 
nated a, the next fastest b, etc. 

In a departure from common practice, the allozyme alleles detected at each 
locus were coded as present or absent for each taxon and included in a phyloge- 
netic analysis. Although advocated by some (Mickevich & Johnson 1976; 
Mickevich & Mitter 1983; Buth 1984), this approach has been criticized by 
Swofford et al. (1996) because it violates the assumption of character indepen- 
dence. Similar treatment of allozyme data from Mimulus sect. Erythranthe 

ABLh 5. Populations used in morphology, allozyme, and nrDNA-ITS studies of Thelespermo. I 
efore collection numbers identify the following collectors: H = C.J. Hansen; S = CJ. Stubbe 
1. D. Windham; Wo = L Allphin Woolstenhulme. All vouchers are deposited at the Univer 

Geographic origins voucher 



Nelson 25622 (RM) 




Tex: Culberson Co.: H/gg/ns 


Wyo: Fremont Co.: H 97-/34 


Wyo: Natrona Co.: f^orfman 





Utah: San Juan Co.: /^fwood 

22534 (BRY) 

Wyo: Sweetwater Co.: 


Wyo: Uinta Co.: HondS 97- /I 


Wyo: Uinta Co.: HondS 97-// 

Ariz: Coconino Co.: 14/0 

Ganders etal. 2000 
Utah:UtahCo.:W/e/s/7 6 

(Windham unpubl.) produced a phylogenetic tree highly concordant with in- 
formation from other sources, suggesting that these data may contain a strong 
phylogenetic signal in spite of their perceived limitations. The allozyme analy- 
sis undertaken here is presented as an experiment to further assess the value of 
enzyme data in phylogenetic reconstruction. 
Phylogenetic Analyses 

All phylogenetic analyses were performed using the computer program PAUP 
(Phj'/ogenetic Ana/j'sistJsing Parsimony, version3.J;Swofford 1991) utilizing ran- 
dom stepwise addition. Only the shortest trees were retained in each search. 
All characters were considered unordered and given equal weights, with multi- 
tracked, organized, and manipulated using the computer program MacClade 
3.0 (Maddison & Maddison 1992). Nodal support in each topology was deter- 
mined bylOO bootstrap replicates (BS; Felsenstein 1985), as well as by calculat- 
ing Bremer support values (BV; Bremer 1988, 1994b). 

Separate and combined phylogenetic analyses were conducted on three 
data sets: morphology, nrDNA-ITS sequences, and allozymes. Examining all 
relevant data in a combined fashion can produce a more robust estimate of 
phylogeny than separate analyses by maximizing congruence among different 
sources of data m phylogenetic mference (Hillis 1987; Kluge 1989; de Queiroz et 
al. 1995; Nixon & Carpenter 1996). To estimate levels of congruence among data 
sets, incongruence length differences (ILDs) were calculated (Mason-Gamer & 

Kel logg 1996; Johnson & Soltis 1998). This mdex measures the amount of extra 

scribed by Mickevich & Farris (1981) and Farris et al. (1994, 1995). The ARNIE 
program m the Random Cladistics software package (Siddall 1995) was used to 
determine the significance of 1 IDs. a-values less than 0.05 were considered suf- 
ficient evidence to reject the null hypothesis of data set homogeneity Com- 
bined analyses included only those taxa common to all phylogenetic data sets. 

Chromosome Data 

All individuals sampled from populations of I puhescens and I caespitosum 
proved to be diploid with chromosome counts of n = 12 (Table 6; Figs. 2A-B). 
Individuals sampled from populations of I suhnudum var. a/pinum consistently 
were tetraploid with n = 24 (Fig. 2C). Our analyses revealed that some Colorado 
Plateau populations of T. suhnudum var suhnudum are diploid (n=12; Fig. 2D) 
whereas others are exclusively tetraploid (n=24; Fig. 2E). Although the geo- 
graphic ranges of these cytotypes overlap and both appear to be common 
(Windham, Hansen, & AUphm unpubl. data), we have yet to identify a locality 
where they occur together. Both ploidy levels of T. suhnudum Yd.r. suhnudum are 
represented in the morphological, DNA, and allozyme analyses that follow. 

A chromosome number was not obtained directly for T. marginatum, and 
no previously published counts were found in the Index to Plant Chromosome 
Numhers. However, an analysis of allozyme banding intensities in our collec- 
tion from Fremont Co., Wyoming, provided strong evidence that the plants at 
this locality are diploid. This sample of I marginatum showed only balanced 
heterozygote banding patterns, as is expected in diploid organisms (Danzmann 
& Bogart 1982b; Wendel & Weeden 1989) and observed m all chromosomally 
verified diploid populations of Thelesperma. 
Morphological Data 

Parsimony analysis of 16 morphological characters yielded two most-parsimo- 
nious trees, the strict consensus of which is shown in Figure 3. Relative to the 
outgroup species chosen, Thelesperma was strongly supported as a monophyl- 
etic group (BV=6, BS=100%). The first branch withm the Thelesperma clade pro- 
duces a trichotomy that separates T. megapotamicum and I/i H/oHum (both with 
unequally lobed disc florets and a well developed pappus) from the other spe- 
cies. The clade encompassing the remaining taxa is weakly supported (BV=1, 
BS=70%), branching to form a polytomy in which each taxon (with the excep- 
tion noted below) occupies its own unresolved branch. The only deviation from 
this pattern is the grouping of T. caespitosum, T puhescens, and T. suhnudum var. 
alpinum in a single clade with strong support (BV=2, BS=88%). Within this clade, 
the last taxon is moderately supported as sister to I caespitosum and T. puhescens. 

Thelesperma caespitosum Dorr 
n=12 Utah Duchesne Co. 

: T.pubescens Dorn 

om Chokecherry 

?0 (UT) 

3 Scott's Bottom 

Wyo Sweetwater ( 

Thelesperma subnudum C 

WSW of Teasdale near the base of Boulder Mtr 
(T29S,R4E,S20);W 93- /44(UT) 
ca.0.6 mi W of State St. in Teasdale on road to 
Bullberry Creek (T29S,R4E,S21);H 97-73 (UT) 
ca.0.8 mi S of SR 24 on dirt road toGovernmei 
Creek (T29S, R4E, S 1 8); H 97-74 (UT) 

ITS Sequence Data 

Of 663 characters (aligned length), 61 were variable, and 6 were phylogeneti- 
cally informative. The sequence of I caespitosum wa.s incomplete with approxi- 
mately 87 bp missing compared to the other sequences. Possible non-specific 
amplification of the ITS region in I caespitosum resulted in a double signal on 
the chromatogram. Multiple attempts to re-extract DNA and obtain a clearer 
signal failed. To determme if there was a loss of phylogenetically informative 
characters m the missmg region of T. caespitosum, two different sequence length 
scenarios were analyzed. The first scenario involved aligning all sequence 
lengths equal to that of I caespitosum (i.e., no gaps in I. caespitosum but with 
an 87 bp truncation in all other taxa). The second scenario was to align full 
sequence lengths in all taxa except T. caespitosum (i.e., 87 bp gap in I. caespitosum 
only). In both analyses, the same single most-parsimonious tree was obtained 




2. Photomicrographs 

in morophologkal data. Bootstrap values are above the line, 

(Fig. 4). Furthermore, bootstrap support only differed by 1-2 percent, being 
slightly higher in the second scenario. These results suggest that very few phy- 
logenetically important characters are located m the 87 bp gap of T. caespitosum 
and subsequent combined analyses utiHzed the second scenario alignment. 

The topology of the single most-parsimonious tree from the IIS sequence 
analysis was similar to that derived from morphology ( Figs. 3, 4). The mono- 
phyly of Thelesperma is again supported by a 100% bootstrap value; Bremer 
support in the ITS data is significantly higher (BV=16). The only topological 
difference between the morphological and ITS trees involves the placement of 
T.filifolium. In the /TSanalysis, this species forms a clade with all Thelesperma 
other than T. megapotamicum on a relatively well supported branch (BV=2; 
BS=86%). As in the morphological tree, all taxa on this branch form an unre- 
solved polytomy with the exception of T. puhescens, T. caespitosum, and T. 
suhnudum var. alpinum. Support for this clade is significantly higher in the ITS 
tree (BV=5; BS=99) and, once again, I suhnudum var alpinum receives moder- 
ate support as the sister taxon of I caespitosum and I. puhescens. A comparison 
of pair-wise distances based on the aligned ITS sequences shows very little di- 
vergence among these rare taxa (Table 7). 
Allozyme Data 

Missing allozyme data made up 5.3% of the total data matrix due to poor stain- 
ing resolution of SKDH and PGI in populations of I marginatum and T. longipes. 
The results of a phylogenetic analysis based on presence/absence data yielded 

3 I 

a single most-pai simonious tree (Fig 5) Relative to the outgioup speeies Bide /li 
cernua, the subset of Thdcspei ma ta\a included m the allozyme analysis iorm 
two distinct clades One consists ol the three i are endemics, which show a strong 
association (BV= 5 BS=95%) com pai able to that observed m the moi phological 
and ITS trees (Figs 3, 4) Once again T sulmudum \ai aJpinum is sistei to T 
caespitosum 'dud T pubc scon, and theic isinenabcd suppoit toi this topology 
(BV=2; BS=85%) Theothei Thclcsponid clade tecoM ud m the allozymeanaly- 
sis(Fig.5)consistsof InuHgnuTtuni T suhnudum and T lon<^'i/Hs which group 
together with moderate suppoit (BV=2, BS= 79%) Neaily identical suppoit 
(BV=3; BS=79%) exists tor the placement of T mayginatum as the sister taxon to 
T. suhnudum and T longipes 
Combined Data Set Analysis 

Statistical compaiisonsol trees lesultingiiom the indi\ idual data sets levealed 
that they were highly congiuent and amenable to being combined m a single 
analysis. This combined analysis of moiphologv allozymes and mDNA-lTS 
the topology of which 
cs alone (Fig 6) Boot- 
mi the allozyme analy- 
) clade is increased by 
reases to 12 Statistical 

equences resulted ma single m 


A^as identical to that ol the ana 

lysis based o 

trap and Bremei suppoi t valu 

sioi thu/m; 

lade are not signilicantly diffei e 

nttiom those 

is. Support for the (alpinum ([ 


ombining data sets The associa 

tion ol these 

bootstrap estimate, and the Bre 

mei support 

- i 

5.5 i 

Th5u2 Thsu4 Thiol Thlo2 Thmal Thma2 Thpul Thpu; 

33 35 34 

n allozyme data only. Bootstrap v; 

We gathered molecular and morphological data, which were analyzed sepa- 
rately and in combmation to obtain the best estimate of the phylogeny of the T. 
suhnudum complex. Tree topologies were highly concordant, and the single 
most-parsimonious tree from the combined analysis was more resolved and 
exhibited improved nodal support over any of the individual analyses. 

The level of congruence among data sets in this study indicates that 
allozymes contam valuable phylogenetic information that can be recovered 
through parsimony analysis. This suggests that concerns regarding the inde- 
pendence of characters (Swoflord et al. 1996) should not disqualify allozymes 
from playing at least a limited role m phylogenetic studies. Because this ap- 
proach is relatively untested, however, we will refrain from placing undue em- 
phasis on the allozyme tree m the following taxonomic discussion. This applies 
to the combined analysis as well, because the topology of that tree may be un- 
duly influenced by the relative abundance of informative allozyme characters. 
The taxonomy outlined below addresses only those patterns independently 
observed in all data sets. 

The relationships portrayed in our phylogenetic trees can be summarized 
as follows. Thelesperma pubescens and T. caespitosum are closest relatives, form- 
ing a clade in all analyses and showing very little divergence from one another 

y, and /rS-DNA data se 

(Tables 4 & 7). Another relationship consistent across all analyses is the place- 
ment of T. suhnudum var. alpinum as the sister taxon to the I pubescens/ 
caespitosum clade. Bootstrap support for this topology ranges from 88% in the 
morphological analysis to 100% m the com bined analysis. Relationships among 
other taxa of the Thehspcrma suhnudum complex are poorly resolved, with the 
aforementioned clade forming a polytomy with I suhnudum, T marginatum, 
and T. longipes in the morphology and ITS analyses (Fig. 3). In the allozyme and 
combined analyses (Fig. 4), those three taxa form a separate, moderately sup- 
ported clade with I marginatum basal to T. suhnudum and T. longipes. These 
results are not wholly concordant with any of the proposed classifications of 
the T. suhnudum complex summarized in Table 2, suggesting that nomencla- 
tural changes are warranted. 
The Thelesperma pubescens/T. caespitosum clade 

Dorn (1990) recognized both taxa in this clade at the species level; Cronquist et 
al. (1994) combined them (and T. suhnudum var. alpinum) under the name I 
pubescens. Welsh et al. (1993) were mute on the matter of T. pubescens (which 
was not known to occur m Utah at the time the flora was published), but prob- 
ably would consider it a variety of I. suhnudum, the treatment accorded to I 
caespitosum. The last classification is the least practicable in light of the data 
presented in this study To treat T. caespitosum as a variety of T. suhnudum and 
maintain monophyly, our data (Figs. 3, 4) indicate that other species would have 
to be subsumed within T. suhnudum as well. These include T. marginatum, T. 

longipes, and possibly even T.filiJoJium. We consider such a species concept, 
encompassing taxa not previously included in the T. suhnudum complex, to be 
unacceptably broad. 

Our data indicate that Thelesperma puhescens and T. caespitosum are sister 
taxa showing minimal genetic divergence. They differ by a single morphologi- 
cal character involving the distribution of pubescence on the leaves (Tables 3 
&r 4; Dorn 1990). Recent collections from near Anthro Mountain on the West 
Tavaputs Plateau in Duchesne Co., Utah (BRY^ Goodrich et al. 25159) provide 
additional insight on their relationship. In that population, individual plants 
range from having strictly petiolar pubescence (I caespitosum type) to com- 
plete leaf blade pubescence (T. puhescens, type). Whether a result of incomplete 
primary divergence or secondary convergence resulting from hybridization or 
selection, there is complete mtergradation. It appears that distribution of pu- 
bescence, m the absence of correlated characteristics, is probably a tenuous foun- 
dation for recognizing species in the T. suhnudum complex. 

Under the Biological Species Concept (Mayr 1942), T. caespitosum and I 
puhescens are "groups of actually or potentially interbreeding populations" that 
probably should be recognized as conspecific. Invoking the Cohesion Species 
Concept (Templeton 1989), T. puhescens and I. caespitosum also qualify as con- 
specific based on phenotypic similarities (cohesion) due to gene flow by means 
of presumed interbreeding. Because these two taxa lack unique, diagnosable sets 
of morphological and molecular characteristics and don't appear to have sepa- 
rate evolutionary trajectories, they also would be considered a single species 
under the Phylogenetic Species Concept (Cracraf 1 1983; Davis & Nixon 1992). 

With regard to the caespitosum/ puhescens clade, our data are most con- 
gruent with the classification presented by Cronquist et al. (1994). In that treat- 
ment, T. puhescens is considered specifically distinct from I suhnudum, with I 
caespitosum included within T. puhescens. We propose two modifications to this 
classification. First, we exclude T. suhnudum var. alpinum from synonymy with 
T. puhescens for reasons that will be discussed below. Second, we propose to rec- 
ognize I caespitosum as a variety of T. puhescens. Excluding the intergradient 
population on Anthro Mountain, Utah, there is a definite correlation among 
pubescence, substrate, and geography (Table 1; Fig. 1). We feel that the incipient 
divergence in this clade is best recognized at the varietal level. 
Origin and relationships of Thelespenna suhnudum var. alpinum 
This taxon appears to be exclusively tetraploid (Table 6), a point that is central to 
hypotheses regarding its evolutionary origin and to its classification. Tetraploids 
typically are derived from diploids through the incorporation of additional sets 
of chromosomes (see Harlan & DeWet 1975), and they are informally grouped 
according to the similarity of their constituent genomes (Crawford 1989). Polyp- 
loids containing genomes that are very similar chromosomally and genetically 


(usually derived from within a single species) are considered autopolyploids. 
Because of genetic similarity to their progenitor diploids, autopolyploids are 
rarely given species status (e.g., Mosquin 1966). Polyploids that contain v/ell- 
differentiated genomes (often obtained through hybridization betvv^een differ- 
ent species) are considered allopolyploids and usually treated as distinct spe- 
cies. The proper classification of T. suhnudum van alpinum thus hinges on 
whether it proves to be an autopolyploid or an allopolyploid. 

Classifications of this taxon by Dorn (1990) and Welsh et al. (1993) imply 
that its closest relative is Theksperma suhnudum, and suggest that it may be an 
autopolyploid derived from within that species. Two lines of evidence refute 
that putative relationship. Although the taxon shows some morphological simi- 
larities to I suhnudum (Dorn 1990), the ITS sequence data clearly indicate a 
closer relationship to I puhescens and T. caespitosum. In fact, all data sets devel- 
oped during this study support the placement of I suhnudum var alpinum as 
sister to these taxa, not I suhnudum var. suhnudum (Figs. 3 & 4). Further evi- 
dence that var alpinum is not an autopolyploid derivative of T. suhnudum comes 
from the fact that known autopolyploids abound in this species and do not re- 
semble var. alpinum. These undisputed autopolyploids in T. suhnudum appeared 
identical to diploid I suhnudum in morphology and ITS sequences (Table 7). In 
our analyses, they were recognizable only through chromosome studies or the 
detection of unbalanced heterozygosity in allozyme markers shared exclusively 
with diploid T. suhnudum. 

Despite significant genetic si m i larity, it also seems unlikely that Theksperma 
suhnudum var. alpinum is an autopolyploid derivative of either I puhescens or I 
caespitosum. It varies toward I suhnudum m some morphological characters, most 
notably the somewhat creeping rootstock. The two taxa are similar enough to 
convince Dorn (1990) to maintain them as varieties of a single species. The pres- 
ence, in most individuals, of a PGI-1 allele otherwise found only in I suhnudum 
and I marginatum provides further evidence of genetic links to taxa outside the 
T. puhescens/ caespitosum clade. Variety alpinum also shows several 
autapomorphic traits that distinguish it from I. puhescens and I caespitosum, 
including pubescent flowering stems and unique allozyme variants. 

We suspect that T. suhnudum var alpinum may be an allotetraploid result- 
ing from hybridization between diploid I suhnudum and either I caespitosum 
or I puhescens. This would explain the pattern of shared characters and appar- 
ent morphological intermediacy that has led to such divergent classifications 
(Dorn 1990 vs. Cronquist et al. 1994). It also would explain allozyme banding 
patterns at the PGM locus, where most individuals of var. a Ipinum are heterozy- 
gous for alleles derived from the puhescens/ caespitosum and suhnudum/ 
marginatum clades respectively. At this one locus, var. alpinum approaches fixed 
heterozygosity, one of the genetic hallmarks of allopolyploidy. The absence of 
fixed heterozygosity at other allozyme loci may be due to homoeologous chro- 

mosome pairing or extensive gene silencing (Wnidham 1988). Over time, al- 
lopolyploids lose expression of duplicated parental genes through various ge- 
netic processes, especially null mutations (Roose & Gottlieb 1976; Werth & 
Windham 1991). Given enough time, the polyploid taxon becomes genetically 
"diploidized" (Grant 1981). 

A paral lei process may explain the lack of diagnostic T. suhnudum ITS se- 
quences in var. alpinum. Recently formed allopolyploids would be expected to 
show the ITS sequences of both diploid parents (Soltis et al. 1995; Cook & Soltis 
1999; 2000; Gernandt & Liston 1999). With time, however, ITS loci often experi- 
ence concerted evolution, which randomly eliminates one of the parental se^ 
quences (Sang et al. 1995; Wendel et al. 1995; Polanco et al. 1999). Under this sce- 
nario, var. alpinum still could be an allopolyploid hybrid in which the original 
I suhnudum sequence has been lost to concerted evolution. 

Each of the possible evolutionary origins (two autopolyploid and one al- 
lopolyploid) of Theksperma suhnudum var. alpinum discussed above would 
support a different classification for this taxon. The hypothesis that var. alpinum 
is an autopolyploid derived from I suhnudum, the only scenario congruent with 
the classifications of Dorn (1990) and Welsh et al. (1993), can be rejected. To 
uphold alpi num as a variety under I sub?iudum and still maintain monophyly, 
we would have to expand the species definition of T. suhnudum to include the 
entire complex plus T. longipes and, possibly, TjiUJolium (Figs. 3 & 4). In our 
opinion, lumping half of the species in the genus Theksperma into one species 
is not a desirable solution. 

Our data are not sufficiently robust to distinguish between an autopolyp- 
loid origin of var. alpinum from within the T puhescens/caespitosum clade or 
an allopolyploid origin through hybridization between members of that clade 
and T. suhnudum. Nevertheless, we can propose a classification that would be 
phylogenetically congruent regardless of which scenario proves more plausible. 
To include var alpinum within T puhescens (as done by Cronquist et al. 1994) 
would be cladistically indefensible if alpinum subsequently is shown to be an 
allopolyploid. However, if we anticipate that the latter hypothesis is more plau- 
sible and recognize var. alpinum as a distinct species, such a treatment remains 
valid in the event that alpinum maintains its current position as the basal 
branch of the T puhescens/caespitosum clade. Variety alpinum is morphologi- 
cally distinct from the other members of this clade and, because of its polyp- 
loid chromosome number, is probably genetically isolated from the strictly dip- 
loid taxa. Thus, it can be recognized as a species under the Biological Species 
Concept (Mayr 1942) as well as the Phylogenetic Species Concept (Cracraf 1 1983; 
Davis & Nixon 1992; Davis 1996). 

and I marginatum. Dora (1990) treated them as distinct species; Cronquist et 
al. (1994) considered the latter a variety of T. suhnudum. These taxa are part of 
an unresolved polytomy in the morphological and 115 data sets (Fig. 3), but 
form a moderately supported clade with T. longipes in the allozyme and com- 
bined trees (Fig. 4). The placement of I suhnudum as sister to T. longipes in- 
stead of T. marginatum in the latter trees suggests a relationship at odds v^/ith 
previous classifications, which exclude I longipes from the I suhnudum com- 
plex. This result should be confirmed by additional studies before taxonomic 
revisions are proposed. Even if the allozyme data are discounted, there still is 
no support for expanding I. suhnudum to include T. marginatum as proposed 
by Cronquist et al. (1994). Recognition of T. marginatum as a variety of I. 
suhnudum would require its placement as sister to T. suhnudum in a phyloge- 
netic analysis. That these two taxa do not form a clade in any of our analyses 
suggests that they should continue to be treated as separate species. Each has a 
unique, diagnosable set of morphological and molecular characteristics indica- 
tive of a distinct evolutionary trajectory, thus satisfying the definition of a phy- 
logenetic species (Davis & Nixon 1992). 

In order to implement the classification outlined above, two nomenclatural 

Thelesperma pubescens Dorn var caespitosum (Dorn) C.J. Hansen, stat. nov 
Basionym: Thelesperma caespitosuni Dorn, Madrono 37: 293. 1990. Type: U.S.A. 
Wyoming. Sweetwater Co.: T18N, R106W, SEl/4 of SEl/4 of Sect. 31 and SWl/4 of 
SWl/4 of Sect. 32, 5 km SE of Green River, barren white shale ridge, 1890 m, 22 
Jun 1988, Dorn 4948 (holotype: RM!). 

Thelesperma windhamii C.J. Hansen, nom. et stat. nov. Basionym: Thelesperma 
suhnudum A. Gray var. alpinum Welsh, Great Basin Naturalist 43: 369. 1983. Type: 
U.S.A. Utah. Wayne Co.: T28S, R4E, S13 (NEl/4), 3 mi due N of Bicknell, bristle- 
cone pine forest on multicolored clay hills, 2745 m, 20 Jul 1980, Atwood and Th- 

This new name for T. suhnudum var alpinum honors the junior author, M.D. 
Windham, who proposed and co-directed the study We chose not to raise the 
epithet alpinum to species rank because the taxon never occupies truly alpine 
habitats and most populations occur at moderate elevations (ca. 2200 m) in 
semi-desert regions. 

1. Leaves and stems glabrous or essentially so; leaves mostly 3-9 cm long;stems mainly 

9-35(-50) cm tall, scattered along a somewhat creeping rhizomatous rootstock; 

blade;plants appearing green T.pubescens var.caespitosum 

It is hoped that this revision of the Theksperma mhnudum complex will re- 
solve some of the taxonomic confusion in the group. Although the treatment of 
T. puhescens and T. caespitosum as conspecific makes the aggregate taxon less 
rare, long-term monitoring and land management still will be necessary. All 
three rare taxa are restricted to specific substrates, and populations of T. 
puhescens near Green River and Hickey Mountain in Wyoming are endangered 
due to off -road vehicle use and oil and gas development. Populations of T. 
windhamii located west of Teasdale, Utah, are similarly threatened by off -road 

To improve phylogenetic resolution, future studies should sequence more 
rapidly evolving regions of the Thelesperma genome, such as the external tran- 
scribed spacer regions of ribosomal DNA (Baldwin & Markos 1998). Artificial 
hybridization studies also might prove useful for revealing reproductive barri- 
ers and species boundaries. Ultimately research should be expanded to include 
all taxa of Thelespernu;. Deciphering rcUuionships withm Thelesperma will aid 
in identifying possible sister genera and thereby contribute to our knowledge 
of higher level relationships m the Coreopsidmae and Heliantheae. 

We thank Chris J. Stubben for help m locating and collecting populations of 
rare Thelesperma and for collecting samples of T. longipes from New Mexico. 
We also thank Lowell Urbatsch for providing ITS primer sequences. We are 
grateful to John Strother and Leigh Johnson for their reviews and helpful com- 
ments on earlier versions of the manuscript. We also give special thanks to 
Marcos Losada for his Spanish translation of the abstract. This study is based 
on a M.S. thesis by the senior author done at Brigham Young University 

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origin and genetic d 


of polyploid! 


wrlghtiona complex { 


lissertation. University 


;ansas, Lawrence. 




Loran C.Anderson 

Department of Biological Science 

Tallahassee, FL 32306-4370, U.S.A. 


sumably its closest relative is L gracilis, from which it differs in morphology, chemistry, and ecolog; 
The species is diploid (n = 10). 

ecologia. La especie es diploide in = 10). 

The upper Apalachicola River region in the Florida panhandle has long been 
known for its diverse and unique flora. Asa Gray (1875) visited the area to ob- 
serve Torreya taxifolia Arn. E.E. Callaway (1966), theologian, amateur arche- 
ologist, and one-time gubenatorial candidate in Florida, cited the unusual flora 
in support of his claim that the original Garden of Eden was located there. Fur- 
ther, he suggested the Torreya tree provided the Biblical "gopherwood" used in 
constructing Noah's ark. Although his ideas did not gain wide acceptance, many 
endemic taxa are nonetheless known from the area (Clewell 1977; Myers & Ewel 
1990). It is the center of one of five "hot spots" for biotic rarity and richness in 
the continental United States (Stein et al. 2000). Now, a new species of Liatris 
can be added to the list of rare endemics found there. 

For many years, local botanists, such as R.K. Godfrey and A.K. Gholson (pers. 
com.), have been aware of some unusual populations of blazing star in ravines 
and along the bluffs of the Apalachicola River in the Florida panhandle. The 
plants would most probably "key out" to Liatris gracilis Pursh, but they do not 
fit well. 

Herz et al. (1984a) studied the chemistry of Liatris gracilis. Later, his team 
(Herz et al. 1984b) studied the "bluffs blazing star" and found that, in contrast 
to authentic L. graci lis, which contained an unusual group of germacrenolides, 
it possessed a distinctive group of heliangolides. 

The Apalachicola bluffs plants are distinctive morphologically and chemi- 

mary of the genus (King & Robinson 1987) makes no mention of this taxon; 
neither does Wunderhn (1998) in his gmde to the flora of Florida. It is here named 
Liatris gholsonii to honor Angus K. Gholson of Chattahoochee, Florida, an in- 
defatigable field botanist and astute student of the local flora. Together, he and 
Godfrey collected an extensive population series of this new species. 



etative and floral features of the new species and of Liatris gracilis were 

isured from the extensive population samples vouchered at FSU. Flowering 


ds were softened m a weak solution of NaOH in 50% ethanol to return tis- 


s to their hydrated size and to facihtate dissection. Meiotic chromosome 


nts were determined from fresh flower buds preserved in Carnoy's solution 


:1 parts alcohol: chloroform: acetic acid). Gholsons personal herbarium is 


z termed "AKG"; it will eventually become part of FLAS. 

Liatris gholsonii L.C. Anderson, sp. nov (Fig. l). Type: UNITED STATES. Florida. 
Liberty Co.: shaded upper slopes of No Name Ravine on the Nature Conservancy's 
Apalachicola Bluffs and Ravines Preserve, TIN, R7W, SWl/4 Sec 7, 13 Sep 2001, 
L.C. Anderson 19932 (hoi.OTYPF.: BRIT; 1.S0TYPE.S: FSU, MO, NY). 

Capitula f 

Perennial herbs, with spherical corm-like rootstocks 2-2.5 cm wide; radical 
leaves (usually absent at anthesis) long-petiolate, 22-25 cm long, blades entire, 
narrowly elliptic-oblong, 20-26 mm wide, glabrate, finely glandular-punctate 
on both surfaces. Stems (4.5-)5.5-f 2(-18.5) dm long, terete, striate, unbranched 
below the inflorescence, pubescent with hairs appressed or spreading. Cauline 
leaves alternate, entire, numerous, much longer than internodes, punctate; lower 
leaves (8-)13-22(-25) cm long, the petioles (2.5-)4-7(-l 0) mm long, ciliate, blades 
elliptic to oblong-lanceolate, tapering at base and tip, (8-)ll-22(-27) mm wide, 
midveins raised and sparsely villosulous; leaves gradually reduced in size and 
becoming sessile toward the inflorescence, grading to the leafy floral bracts 
with little change in size or shape. Inflorescence elongate, determinate, race- 
mose (rarely paniculate in damaged plants or frequently so in garden plants), 
pedicels spreading or reflexed at anthesis, pubescent, (2-)5-12(-26) mm long 
on heads toward base of inflorescence. Lowest floral bracts sessile, narrowly 

elliptic to lanceolate, (2-)2.5-4.5(-5.9) cm long, (1.8-)2-6(-10) mm wide. Heads 
turbinate-cylindric, (6-)6.5-8.5(-10) mm long, phyllaries (8-)9-12(-15), green 
to dark purple, ovate to oblong, acute to acuminate (rarely obtuse), glandular- 
punctate, ± ciliate, margins hyaline, white or roseate. Disk flowers 3-5(-6); co- 
rollas (6.7-)7-8.5(-10) mm long, narrowly funnelform, sparsely glandular punc- 
tate, throat pubescent internally the tube ochroleucous or pinkish, the lobes 
pink to purple, slightly spreading to ref lexed, (2.1-)2.5-3.6(-4,6) mm long; bifid 
styles pmk, (If-)f 2-13(-14) mm long, anthers (2.5-)3.5-4(-4.4) mm long, with 
retuse appendages 0.2 mm long; mature achenes 3.5-5 mm long, 10-ribbed, gray 
except for raised, dull green ribs, narrowly turbinate, densely hispidulous, the 

hairs 0.4-0.5 mm long, pappus strongly barbellate, 5^6 mm long, white, tmged 
with purple to give a dull rosy appearance, n = 10. 

Phenology-Vumanly flowering from September to mid-October; a very few 
precocious plants blooming in early July and a few blooming in early November 

Habitat.-The species occurs in loamy sand or gravelly sand mostly m deep 
shade of deciduous woodlands on upper slopes of bayhead ravines or along the 
less shady ecotone between the woodland at tops of ravines adjoining more open 
scrub oak woodland. It also occurs m xeric to sub-mesic woodland and more or 
less open sites on bluffs facing the Apalachicola River or on sandy ridge tops 

Associated woody species include: Asimina parvijlora (Michx.) Dunal, 
Callicarpa americana L., Carya pallida (Ashe) Engelm. & Graebn., C. tomentosa 
(Poir. in Lam.) Nutt., Cornus florida L. , Diospyros virginiana L., Forestiera 
godjreyi L.C. Anderson, Fraxinus americana L,, Ilex opaca Ak.Juniperus 
virginiana L , Magnolia grandiflora L., Osmanthus americanus (L) Benth. & 
Hook, ex A. Gray, Ostrya virginiana (Mill.) K. Koch, Persea horhonia (L.) Spreng., 
Pinus glabra Walt., Prunus alahamensis Mohr, Quercus alba L, Q. geminata 
Small, Q. hcmisphacuca Bartr, Q nigra L, Rhus copallina L, Seba'^tiania 
j) utkosa t,Bartr)rcin,Siderox3/bn lanuginosum Michx , Vaccinium arhoreum 
Marsh , and Vibumum dcntatumL A few herbaceous species are associates m 
the shaded sites such as Cha^manthium scssi/i/lofum (Poir) Yates, Muhclla 

:amp torre>a ' 21 Sep 1983 RK Godf}c_ 
(FSU), ApaUchiLola River bluff W o 

'egetativeX R.K. Godjrey 83776 (FSU); in open, 2 Oct 1983 A 
Oct 1983, A.K. Gholson 10644 (AKG, FSUJ; ridge by Kelley B 


Gaiser (1946) recognized 10 series of species in her monograph of Liatris. Series 
Graminifoliae (with obtuse, ciliolate phyllanes as in L. gracilis) and series 
Pauciflorae (which has generally larger heads with acuminate phyllaries that 
lack cilia) are relevant to this discussion. The phyllanes of L. ghohonii are cili- 
ate (as in Graminifoliae) but usually acuminate (as in Pauciflorae). The dis- 
tinctiveness of the two series was already diminished when Godfrey (1961) de- 
scribed L. provincialise that Florida panhandle endemic has strongly acuminate 
but ciliolate phyllaries. 

Gaiser (1950) studied the cytology in these series and found the species were 
all diploids (n = 10). The chromosomes were small with uniform karyotypes. 
For Liatris ghohonii, meiotic chromosome counts from buds of Anderson 19932 
were n = 10; the chromosomes were also very small, so chromosome numbers 
and karyotypes are not distinctive features taxonomically 

The new species appears most closely related to Liatris gracilis; both have 
small heads on relatively long, usually spreading pedicels. The two differ in 
their ecology, chemistry (Herz et al. 1984a, b), and morphology. Typical habitat 
for L. ghohonii is on more or less mesic, shaded ravine slopes and bluffs; only a 
few plants occur in full sun (m openings in the canopy from tree-fall or along 
the ecotone between the ravine and the adjacent sandhills), whereas L. gracilis 
typically occurs in sunnier, more xeric, open f latwoods and savannas. The two 
species are not sympatric, but L. gracilis does occur nearby in the same county. 

Foliage is perhaps the most distinguishing feature separating the two spe- 
cies. Basal leaves on young plants are broadly elhptic in Liatris ghohonii (as in 
Godfrey 84073), whereas they are narrowly lanceolate in L. gracilis (see photo- 
graph m Gaiser 1950). Flowering plants of L. ghohonii tend to retain more of 
the basal and lower cauline leaves, which are also wider (mostly 11-22 mm wide) 
than those of L. gracilis (4-13 mm wide). The transition between cauline leaves 
and floral bracts is gradual in the new species, and the floral bracts are mostly 
2-6 mm wide, whereas they are narrower (1.0-1.8 mm wide) in L. gracilis. 

One might think the differences in foliage could be products of the species' 
differing habitats. In general, plants of the same or related species grown in full 
sun have smaller, narrower leaves than those found on plants grown in the shade. 
However, the larger leaves of L. ghohonii are not due to habitat. The few collec- 
tions of L. ghohonii taken in full sun were more robust plants that had shorter 
internodes, average-sized cauline leaves for the species, but wider floral bracts 
than plants from shadier sites. Plants grown in full sun tended to have more 
phyllaries in their involucres (and thus be more like those of L gracilis) than 
those grown in shade. 

Personnel of the Apalachicola Blulls and Ravines Preserx-e established a 
wildf lower garden in full sun near their headquarters building. The garden soil 
was enriched with mushroom compost and was given supplemental water 
Seeds of Liatris gholsonii from No Name Ravine were sown in the garden m 
1997, and many plants became established. These plants are somewhat atypi- 
cal in having mostly two or three stems per rootstock and large, paniculate in- 
florescences. These "full sun" plants still had the characteristic leaves and flo- 
ral bracts of L. gholsonii. 

Several features of floral morphology tend to separate to two species, but 

lute identification. For example, phyllarics in the new species tend to be acumi- 
nate, but some plants have phyllaries that are only acute (or rarely obtuse), and 
phyllaries m Liatris gracilis are usually very obtuse (rounded apically), but a 
few specimens have phyllaries that are acute or rounded but mucronate. Sum- 
mation of some floral measurements from over 30 samples per species is in Table 
1. Plants of L. gholsonii tend to have shorter involucres (shortest m the genus) 
with fewer phyllaries, but their corollas and achenes tend to be larger than those 

Differences between the two species in floral morphology (Table 1) are 
present (though minimal is some cases). The salient distinguishing features 
between the two taxa are mostly vegetative; the following key couplet provides 

wide; phyllaries usually acuminate, rarely obtuse Liatris gholsonii 

1 2 mm wide; phyllaries usually obtuse, rarely acute Liatris gracilis 

Now the Florida panhandle has two endemic species of Liatris] L. provincialis 
is found in Franklin and Wakulla counties and is considered endangered in 
Florida (Coile 2000). The range of L. gholsonii is even more restricted; it ranges 
about 14 km from Torreya State Park to just south of Alum Bluffs and occurs on 
bluffs or in ravines only a short distance eastward from the Apalachicola River 
The most distant population from the river is found in the headwaters of 
Sweetwater Creek about 9 km from the river This species is clearly a candidate 
for listing as endangered. 

Staff of The Nature Conservancy's Apalachicola Bluffs and Ravines Preserve 
provided some logistical support. Mark Garland supplied the Latin diagnosis. 
Ken Womble scanned the specimens into Adobe Photoshop for the illustration. 
Guy Nesom, Bob Krai, and A.B. Thistle and anonymous reviewers provided help- 
ful suggestions on the manuscript. 

Corolla lobe lenc 
Achene length, n 

Callaway, E.E. 1966. In the beginning: creation, evolution, Garden of Eden, and Noah's ark. 

Carlton Press, New York. 
Clewell,A.F. 1 977. Geobotany of the Apalachicola region.Pp.6-1 5. In: R.J. Livingston and E. 

A. Joyce,Jr.,eds. Proceedings of the Conference on the Apalachicola Drainage System. 

Fla. Dept. Natural Resources, Marine Research Laboratory. Fla. Marine Res. Pubi. 26. 
CoiLE,N.C. 2000. Notes on Florida's endangered and threatened plants. Florida Dept.Agric. 

Consumer Serv, Div. Plant Industry-Bot. Sect. Contr. 38, Gainesville. 
Gaisir, L.O. 1946. The genus Liatris. Rhodora 48:165-183,216-263,273-326,331-382, 


L.0. 1950. Chrome 

5S in /./arr/5. II.Graminifoliaeand Pauciflorae.Amer.. 

51 . Liatris provincialls, sp. nov. (Compositae), endemic in western Floridc 

pilgrimage to Torreya. Amer. Agric. 34:266-267. 

mis gracilis. Phytochemistry 23:373-382. 

^NABE, R.K.GonrLY, and J.F.Blount. 1984b. 3-germacren-6,12-olides from ai 
Liatris taxon. Phytochemistry 23:599-606. 

I. Robinson. 1987. The genera of the Eupatorieae (Asteraceae). Monogr. ir 

.J. EwEL. 1 990. Ecosystems of Florida. Univ. Central Florida Press, Orlando. 
JTNER, and J.S. Adams. 2000. Precious heritage, the status of biodiversity ir 
tates. Oxford Univ. Press, New York. 
598. Guide to the vascular plants of Florida. Univ Press of Florida, Gainesville 


RGE E. Burrows and RonaldJ. Tyrl. 2001. Toxic Plants of North America. (ISBN 
0-8138-2266-1, hbk.). Iowa State Ptess 2121 State Avenue, Ames, lA 50014, 
U.S.A. (Orders: 1-800-862-6657, 1-515 292 3348 ia\ wwwiowastatepress com) 
$174.95, 1350 pp, 8 color plates, numerous line di awmgs 8 1/2' x 11' 

lightful dedication to the late Dr. J.W.Dollahiteot Texas A&MUmveisity who made impor an con 

cally arranged treatments for 75 families of plants that aie kno\^n to be poisonous The t ea me 
are divided into genera, and the treatment for each genus mcludes a consideration ot the pec es 
Nomenclature conforms to current f loristic usage in North America. There is a general desc p or 
of the plant and a statement of range and habitat that are designed to permit ready iden f ca on 

ease genesis, clmical signs, pathology, and treatment & control. The page margi ns are wide, and pr ec 
there are Une drawings and geneiahzed distribution maps lor each plant Short phiases in boldfacf 

.s detailed chapter© 

z English language literatt 




Alexander Krings 

Herbarium, Department of Botany 

Recent study of lianas and vines in southern Costa Rica resulted in a prelimi- 
nary checklist of seventy species in thirty-two families from the Las Cruces 
Biological Station (Krings 1999). Ongoing work with collections resultmg from 
this study revealed a new species of Gonolohus Michx. (Apocynaceae: 
Asclepiadeae, Gonolobinae). 

Gonolobus tenuisepalus Krings, sp. nov. (Fig. l). Type; COSTA RICA. Puntarenas: 

Coto Brus, San Vito de Java, Estacion Biologica Las Cruces, 30 m downhill from 

Trail Marker Ft 59, 1200 m elev, 11 Mar 1997, Krings 274 (holotype: F; isotypes: CR, 


Species nova Gonolobus distinguibili a Gonolobus jaliscensis B.L. Rob. & Greenm. petioli c. 4-4.3 cm 

Twining vine; indument of short and long hairs, the short hairs straight, pointed 
or sometimes capitate, to 0.4 mm long, borne primarily along the stem, midvein, 
and secondary veins, long hairs straight to f lexuously curved, (2-)2.8-3.5(-4.3) 
mm long, borne along stem, veins, and over leaf surfaces; stems densely yellow- 
ish-clear pilose or hirsute; latex white, abundant; petioles 4-4.3 cm long, densely 
yellowish pilose to hirsute; leaves opposite, 12-14 cm long, 6.8-8 cm wide, ovate, 
densely yellowish pilose to hirsute on both surfaces, the bases cordate to some- 
what auriculate, frequently oblique, the apices acuminate; inflorescences axil- 
lary umbelliform, densely 9-15-f lowered, the peduncles 0.5-1 cm long; pedicels 
7-9 mm long, densely yellowish pilose to hirsute, the hairs to 3 mm long; flow- 
ers mildly fetid; sepals 5, linear to narrowly lanceolate, 5-7.2 mm long, pilose or 
hirsute on the outer surface, glabrate on the inner surface; corolla campanulate 

hQ.1. Gonohbus tenuisepalus Krings: A, Leaves and inflorescence; B. Long and short st 
flower in bud; E. Dried, flattened flower; F. Living flower, showing erect sepals and 
flattened flower (horizontal arrow indicates glabroi 
Pollinarium. All based on Krings 274 (HT: F; IT: CR, N' 

iomes;C. Inflorescence; D. 

to suburceolate, 5-lobed, the tube 1-2.5 mm long, the lobes deltate, uniformly 
flat, dark purple to dark brownish-red, 3-4 mm long, conspicuously shorter 
than the sepals, long yellowish pilose to hirsute below, the hairs to 2.8 mm long, 
coarsely villosulous-strigillose above; faucal annulus 5-lobed, ca. 0.05-0.1 mm 
thick, glabrous; corona 1, 5-lobed, fused at the base, adnate to and surrounding 

the gynostegium, the lobes deltoid to rhomboid, essentially as high as the style- 
head, 0.4-0.5 mm high; anthers with fleshy, obdeltate, dorsal appendages; 
pollinaria horizontal, ca. 0.5 mm long; follicles unknown. 

Gonolohus comprises about 150 species limited to the New World tropics 
and subtropics (Stevens 2001). Gonolohus tenuisepalus appears to be closely re- 
lated to Gonolohus jaliscensis B.L. Rob. & Greenm., sharing indument of short 
and long hairs, densely multi-flowered, umbelliform inflorescences, small co- 
rollas (ca. 8 mm in diam.), and similar coronas and anther appendages. 
Gonolohus jaliscensis, however, exhibits petioles shorter (to 2.2 cm long), sepals 
shorter (only to 2 mm long), corollas greenish to yellowish (purple m G. 
tenuisepalus), and anther appendages mushroom-shaped. With the addition 
of G. tenuisepalus, nine species of Gonolohus are known from Costa Rica (two 
yet undescribed, see Stevens 2001). An updated key follows: 
1 . Corollas barbellate within or with barbed lobes. 

2. Abaxial and adaxial leaf surfaces glabrous, except scattered appressed-pilose 

along the principal veins and in the axils;faucal annulus glabrous;anther append^ 

ages > 1.4 mm long 

_Gonolobusedulis h 

! or Ies5;faucal annulus 0.05-0.1 mm high, 
appendages obdeltate, ca. 0.5 mm long 
Gonolobus tenuisepalus Krings 

lobes 3= 6 mm long;faucal annulus various;anther appendages lacking (G. sp.B) 
OR ^ 0.3 mm long and variously shaped, the apices entire, bifid.or with a central 

4. Outer petal surface with trichomes 0.8-1.5 mm long; anther appendages 

bifid Gonolobus heterophyllus (Hemsl.) W.D.Stevens. 

4. Outer petal surface glabrous or the trichomes =s 0.4 mm in length; anther 
appendages entire, mamillate tipped, or lacking, but not bifid. 
5. Corolla lobes 5= 15 mm long; anther appendages subquadrate, 1-2 mm 

s albomarginatus (Pittier) Woodson 

Corolla lobes =s 12 mm long;anther ap 

pendages various or lacking 

6. Corolla purple, the lobes 7.2-9.5 mr 

n long, the apices coiK.i e fducal 

annulus an erect, 5-lobed tube, 0.7- 

1.2 mm long, anther a[)|). nfi^nns 

Gonolobus sdB 

6. Corolla greenish oryellowish (someti 

mes purple in GspD th 1 >\ 

various lengths, planar; faucal annul 

i various, sometimes in iiidMiiiM 

ridge (e.g., G.c/i/r;qu;em/s); anther ap| 

pendages present. 

7. Leaf bases truncate to rounded; ar 

ither appendages rounded, with a 

short, central, mammillate tip 

Gonolobus chiriquensis 

Collections at Las Cruces Biological Station were made possible thanks to a travel 
grant and assistantship provided by the College of Forest Resources at North 
Carolina State University and fellowships in Tropical Biology from the Organi- 
zation for Tropical Studies (OTS). I thank: OTS for permission to conduct re- 
search at Las Cruces and for help with collecting permits, Luis Diego Gomez for 
support and insight during my stay, Paul Fantz for help with the Latin diagno- 
sis, Mary E. Endress and J. Francisco Morales for thoughtful reviews of the manu- 
script, and the following herbaria and their curators (particularly Robert Wilbur, 
William Burger, and Pedro Acevedo) for access to their collections: DUKE, F, US. 

Krings, A. 1 999. An annotated preliminary checl<list of the dicotyledonous lianas and vines 

from the Las Cruces Biological Station, Costa Rica.Sida 18:1247-1258. 
Stevens, W.D. 2001 . Asclepiadaceae. Mongr. Syst. Bot. Missouri Bot. Card. 85:234-270. 
Woodson, R.E., Jr., R.W. Schery, and D.L Spellman. 1 975. Flora of Panama. Part VIII. Family 1 63. 

Asclepiadaceae. Ann. Missouri Bot. Card. 62:1 03-1 56. 


N.Sasidharan and P.Sujanapal 

Kerala, INDIA 


;iatic genus is estimated to have 430 species 
jenus has a bicentric distribution pattern due to 
the movement of landmasses during the late Cretaceous period from a 
Gondwanian origin (Nayar 1972). In Asia, the distribution ranges from Indian- 
Himalaya southwards to Sri Lanka, Myanmar, Thailand, Cambodia, Vietnam, 
Malay Peninsula eastward through the islands of the Malay Archipelago, New 
Guinea down to Northern Australia, and is adapted to warm humid 
paleotropical climate. More than 300 species are reported from the Indo-Ma- 
laysian region (Nayar 1966). Malayan regions and Madagascar are the principal 
centers of diversity and endemism- Philippines: 80 species, Madagascar: 70 spe- 
cies, Borneo: 48 species (Perrier 1951; Regalado 1990, 1995). Nayar (1972) has 
analyzed the distribution pattern of Asiatic Melastomataceae and recognized 
four centers. The Western Ghats is included in the Deccan-Ceylon center (Pen- 
insular India and Sri Lanka) of species formation. Among the 9 Indo-Sri Lankan 
species, 5 are reported from Peninsular India and Sri Lanka and others are con- 
fined to the subtropical Himalayas and Assam hills. Medinilla heddomei Clarke 
and M. malabarica Bedd. are exclusive to Southern Western Ghats and the third 
species, M.fuchsioides Card., formerly considered endemic to Sri Lanka, was 
recently reported from Southern Western Ghats (Shetty & Karthikeyan 1976). 
During explorations in the windward region of Southern Western Ghats, 
interesting specimens of Medinilla were collected from three localities of Anamalai 
Hills. The specimens did not agree with the species known from India and Sri Lanka. 
Further study with literature on Medinilla (Bremer 1987; Perrier 1951; Regalado 
1990, 1995) shows that the specimens have more affinity to Indo-Sri Lankan 

:cies and is allied to Medinilla malaharica Bedd. The specimens were further 
npared with the collections from Indo-Malaysia in the Kew Herbarimn and 
matches were found. It is described and illustrated here as a new species. 

Epiphytic suhshrubs; branches fleshy, acutely quadrangular or 
subquadrangular, sparingly branched, sometimes rooting from lower nodes, 
smooth, on drying forms a winged appearance, stem 20-35 cm long, young 
shoots reddish-brown. Leaves membranous, opposite pairs unequal, sometimes 
equal, larger one 7-11 5 x 3-5 cm, smaller 4-6 x 2.5-3.5 cm, elliptic-oblong, base 
subcordate or obtuse, apex obtusely acuminate, acumen sometimes twisted, 
margin entire, 3-nerved, lateral ribs near the margins, famt towards apex; peti- 
ole 2-4 mm, flattened, glabrous. Flowers 4-inerous, 2-5 in axils or from leafless 
nodes m horizontal row, sometimes 2 or 3 on a short stalk (1-2 mm); pedicel 
with 2 minute bracteoles at base, jointed, 2 mm at anthesis, elongating to 4 mm 

puberulous, shortly narrowed at mouth with 8 minute teeth; petals 5-7 x 4 mm, 
rose-pink, obliquely obovate, minutely apiculate; stamens 8, filaments 3 mm 
long; anthers 2 mm long; ovary 1.5 mm across, obovoid, 4-celled, connected to 
hypanthium with 8 membranous appendages; style to 7 mm long; stigma mi- 
nutely capitate, papillate. Berries 4 mm across, globose, glabrous; crowned by a 
small hypanthial limb;seedsiTiany yellowish-brown, 1.5 x 1mm, smooth, ovoid, 

Med in i Ua a namalaiana is aWied toM. malaharica Bedd. in the general ap- 
pearance of the leaves and reddish-brown tender shoots. However, it can be dis- 
tinguished I rom the latter by the succulent quadrangular or subquadrangular 
branches, 3-nerved membranous leaves with obtusely acuminate apex and 
short pedicelled flowers in axils or from leafless nodes in horizontal rows. 
Medinilla malaharica Bedd. has terete woody branches, 5-nerved thickly co- 
riaceous leaves with acute or obtuse apex and flowers in pedunculate cymes. 

Note.— The floral structure of Medinilla appears to be uniformly similar 
in most of the species and is not much diagnostic value. Because of the high 
rate of endemism in Medinilla most of the workers followed geographic distri- 
bution m addition to morphological or vegetative characters in their regional 
revisions or f loristic works. Our taxonomic concept for describing M. anamalaiana 
also follows the same approach in addition to morphological characters. 

Flowering and Fru iting.-January-June. 

Etymology— The specific epithet of the new species is derived after its type 
locahty, the Anamalai hills. 

Distribution and Status— The species is so far restricted to the evergreen 
forests, with an altitude ranges from 400-1300 m in the windward side of 
Anamalai hills. 

Habitat and Ecology— An epiphyte in the crevices of trees in the middle 
stratum of evergreen forests. This species is often found associated with epi- 
phytes like Aeschynanthus perrottetii A. DC. Common tree associates are 
Gymnanthemum arborea (Buch.-Ham.) H. Rob, Elaeocarpus glandulosus Wall, 
ex Merr., Turpi nzama/flbaricfl Gamble, Drypeteswightii (Hook, f.) Pax &Hoffm., 
Palaquium ellipticum (Dalz.) BailL, Calophyllum polyanthum Wall, ex Choisy, 
Mcsua thwaitesii Planch. & Triana, etc. 

Paratypes: INDIA. Kerala. Palakkad District: Parambikulam Vv/ildlife Sanctuary, Puliyala 1000 m, 3 
Apr 2000, P Sujanapal 30043 (KFRl). Pathanamthitta District: Goodrical Range, Moozhiyar 500 m, 
23 May 1984, N Sasidharan 3142 (KFRI). Idukki District: Periyar Tiger Reserve, Vallakkadavu 900 m, 


. Flowers in pedut 

ateful to Wildlife Wmg of Kerala Forest Department for the fi- 
ctor, Kerala Forest Research Institute, for providing fa- 
cilities and Wildlife Wardens and Staff of the Parambikulam Wildlife Sanc- 
tuary and Periyar Tiger Reserve. Sincere thanks are also due to J.F Veldkamp, 
Rijksherbarium, Leiden for critical comments and Latin diagnosis; VB. Sajeev 
for illustrations. The first author is thankful to E. Nic Laughadha, Kew Her- 
barium for the help rendered in referring the relevant Medinilla specimens. 


Bremfr, K. 1987. Melastomataceae In: M.D. Dassanayake and F.R. Fosberg, eds. A revised 
handbook for the flora of Ceylon. Amerind Publishing Co. New Delhi.Vol, 6, pp. 

YAR, M.P. 1972. Centres of development and pattern o 

f distnbution ofthefamil 

Melastomataceaeinlndo-Malesia.Bull.Bot.Surv.lndia 14 


YAR, M.P. 1966. Contributions to the knowledge of Indo 

Malaysian and other Asiati 

Melastomataceae. Kew Bull. 20:235-244. 

RRiER DE LA Bathie, H. 1 951 . Melastomatacees In: H. Number 

ed. Flore de Madagascar e 

ComoresFamilie 153:1-326. 

oAi ADO, J.C. Jr. 1990. Revision of Medinilla (Melastomatace 

e) Borneo. Blumea 35:5-70. 

GALADO, J.C. Jr. 1995. Revision of Philippine Medinilla (Me 

lastomataceae). Blumea 40 


ETTY, B.V. and S. Karthikfyan. 1976. Medinilla fuchsioides Ga 

ertn. (Melastomataceae)— /^ 

new record for India. Bull. Bot. Surv. India 1 8:21 5. 

Dennis W. Woodland. 2000. Contemporary Plant Systematics. Third Edition. (ISBN 
1-883925-25-8, hbk ) Andrews University Press, 213 Information Services 
Building, Berrien Springs, MI 49104-1700, 616-471-6134 (Orders 
aupress@andrews,edu http//wwwandrevv^sedu/press) Price not given, 
569 pp , numerous Ime dt awmgs, 7 ' 10 1/2" 

^serving specimens and the 
vmg plant diversity that mc 
:ome endangered genetic ci 
ints And finally an epilogue 

ilasol 4 700 color 

Kathryn Bernick. 1998. Basketry & Cordage from Hesquiat Harbour. (ISBN 0- 
7718-0525-9, pbk.). Royal British Columbia Museum, RO. Box 9815 Stn Prov 
Govt, 675 Belleville Street, Victoria, British Columbia, VSW 9W2, Canada. 
$14.95, 152 pp., 114 figures, 6" x 9". 




iversity of Western Ontario 'Botanical Research Institute ofTexc 

utment of Plant Sciences 509 Pecan Street 

151 Richmond St N. Fort Worth, TX 76 102-4060, U.S.A. 

Ontario, N6A 5B7, CANADA 

intish Columbia. They a 

3.nd Cokanaganensisvi 

ir wellsii, C enderbyensis, C. orbicularis and C. atrovirens 

ructi). This generates i 

I rich western North American center of diversity for 

species so far known fr 

om this newly recognized center. Four of the species de- 

demic to the northern C 

)kanagan but C castlegarensis ranges widely to northern 


V taxa are illustrated and mapped. Revised keys for the 


ystematicOkanagan, diversity center 

The northern Okanagan of British Columbia and the southwestern part of the 
adjacent Shuswap drainage is the richest area for Crataegus in North America 
west of the Rocky Mountains. In addition to the five new species described here, 
there are another three (Phipps & O'Kennon 1998) recently described species 
O'Kennon), as well as four long-ago described species, C chrysocarpa Ashe (incl. 
C. piperi Britton = C. columhiana Howell, sens, auctt.), C macracantha Lodd. ex 
Loud., C. douglasii Lindl., C suksdorfii (Sarg.) Kruschke, plus the introduced C 
mo nogyna Jacq., for a total of 13 species. This is enough to recognize a local diversity 

center for Crataegus (Fig. 1), by far the richest m western North America for an 
area of this size. This diversity center is mapped m more detail in Fig, 2 where 
many of the collection localities cited in this paper can be located. The contrast 
with earlier understanding of Crataegus in British Columbia is striking. T.M.C. 
Taylor (1973) listed only three species for the province, the native C. douglasii 
and C. columhiana and the introduced C. monogyna, although the illustration 
labelled C columhiana is clearly C. macracantha as we have noted previously 
(Phipps 1998). 

The native hawthorns of the Okanagan-Shuswap belong to a diverse range 
of series, not necessarily particulady closely related to one another, viz.: 
Douglasianae, Purpureofructi, all the species of which occur in the region, and 
Macracanthae and Rotundijoliae (sens, auctt.). The reasons lor this rich variety 
of hawthorn in the northern Okanagan and southwestern Shuswap drainage 
cannot be spelled out precisely but presumably relate to the location of former 
glacial refugia, as well as the benign climate. It is, of course, clear that wherever 
the relevant refugia were at a given time they must have been constantly relo- 
cated by changes of Pleistocene climate. In this mountainous region, hawthorns 
are virtually restricted to the valley bottoms, become rarer southwards (the 
southern Okanagan is much drier) and are nearly absent from the forest envi- 
ronment of the middle slopes. In the somewhat mesic area of the northern 
Okanagan, which is the area of greatest natural diversity, natural hedgerows 
dominated by hawthorns are a feature of the agricultural landscape and haw- 
thorns also occur along smaller watercourses and ditches as well as forming 
downslope hillside thickets in some moister sites. The annual rainfall here is 
around 500-600 mm. but falls off to about 200 mm in the southern Okanogan. 

In this paper we describe the new taxa C. castlegarensis (ser. Douglasia nae), 
C. shuswapensis (? ser Douglasianae), C. okanaganensis var. welhii, C. orbicu- 
laris^ C. enderhyensis and C. atrovirens (all ser Purpureofructi). Updated keys 
are produced for the series in question. All species are illustrated with line il- 
lustrations. Distribution maps of the new taxa were created by Range-Mapper 
(Tundra Vole Software, Fairbanks, AL) from files of latitude-longitude coordi- 
nates and then artistically improved by Susan Laurie-Bourque using Corel Draw. 
The discoveries are the result of several years of field work by the authors in the 
mid4990s and 2001-2 plus further discoveries in 2000 by the first author with 
his wife Sheila and son William. The collections of May, 2002 substantially in- 
crease the number of flowering specimens previously available for study and 
from which the cited material derives. 

In this part we set out the new taxa in the context ol others in the same series by 
providing a key to series and references to descriptions of the already known 

species in these series. In the cases of C. douglasii, which has been much con- 
fused with other hawthorns in the area, and C. suksdorfii, both of which, as far 
as we know, lack modern, detailed species descriptions, we also provide full 
English descriptions. We also provide amended series descriptions. The depth 
of lobing of leaves is referred to by term 'leaf incision index' or 'LIF ('IFF in latin). 
Series Douglasianae (Rehder ex C.K. Schneid.) Rehder 

Bushes to small trees; 1-year old wood generally tan to deep purple^brown; 
thorns mostly 1.5-2.5 cm long, dark and shiny when young. Leaves deciduous; 
petioles fairly short; blades 3-5(-7) cm long, broadly elliptic or elliptic to some- 
what obovate m general shape, shallowly (0-)2-3 lobed per side, the lobes ob- 
tuse to subacute; margins toothed; venation craspedodromous to nearly fully 
camptodromous in some unlobed forms; petioles and leaf teeth tips somewhat 
glandular Inflorescences 8-20 flowered; branches glabrous or pubescent, 
bracteolate; anthesis early. Flowers 12-18 mm diam.; calyx-lobes subentire, 
small; stamens 10 or 20, anthers pink; styles 3-5. Fruit smallish, 0.6-1.0 cm long, 
ellipsoidal to subspherical, black or dark purple at full ripeness, hairy or gla- 
brous; nutlets 3-5, dorsally furrowed, lateral faces pitted. 

This is a small series of ± widespread 'black-fruited' hawthorns of the 
northwestern USA and adjacent Canada with four species, one of which, C. 
douglasii, extends to the Great Lakes area. Series Douglasianae are notable for 
their laterally pitted nutlets, short thorns and dark purple to black fruits at full 
maturity in which they resemble Eurasian ser Nigrae. 'We add one new species 
to this group and amend the key 

1 . Stamens 20; leaf-blades often unlobed, if lobed, lobes generally obtuse to subacute; 

purple or even black by late August, very bloomy at this time 2. C. doi 

Thorns 1 .5-2 cm long; flowers 15-18 mm diam.; fruit ampulliform, chestnut 
orburgundy in late August, generally not very bloomy B.C.okei 

onally paired or in triads;fruit subglobose to depressed, passing through crim- 
3n-lake (purplish-crimson) to burgundy in late August 4.C.castlegarensis, ; 

. Public 

Bushes, often 3-5 m tall, perhaps larger; one-year old twigs tan, older grayish; 
thorns 1.5-2.5 cm long, recurved or straight, glossy dark brown when young. 

Leaves deciduous; petioles 0.75-1.5 cm long, glabrous, eglandular westwards, in 
Montana glandular; blades ± rhomb-elliptic to ellipt-obovate in general shape, 
3-6 cm long, tapered to the base, unlobed to irregulariy sharply to very bluntly 
2-3 lobed per side; margins toothed, the teeth in Montana glandular; venation 
craspedodromous to semi-campdodromous in blunt-lobed western specimens; 
appressed hairy above, ± glabrate below except pubescent near the veins in 
Oregon. Inflorescences 7-15 flowered, branches glabrous, punctate, bearing 
small, early caducous, linear, membranous, gland-margined bracteoles. Flow- 
ers ca. 15-17 mm diam.; hypanthium glabrous externally; calyx-lobes ca. 4 mm 
long, triangular, abaxially glabrous, margins minutely and sparingly glandu- 
lar-serrate; petals ± circular, white; stamens 20, anthers pink; styles 4-5. Fruit 1 
cm long, subglobose to broad-ovoid in the few specimens seen, black at matu- 
rity, dull vinous purple a month before; calyx-lobes ref lexed; nutlets 4-5, dor- 
sally grooved, sides erose. 

Crataegus suksdorfii occurs mainly along the coast and ranges from California 
northwards to Vancouver Island, the Queen Charlotte Islands and coastal Alaska 
to around Anchorage. It is also scattered sporadically inland east to Montana. 
Crataegus suksdoifi is by far the most northerly occurring Crataegus in North 
America at 61° N lat. It is diploid according to Brunsfeld and Johnson (1990). 

In the relatively few specimens 1 have studied, the fruit is smaller and more 
globose than in C. douglasii. It is most convincingly differentiated from C. 
douglasii by stamen number and perhaps by fruit shape and early color, but 
like C douglasii, it also has a variety of leaf shapes and a venation type which 
lie generally outside the range of that species. The glandular leaf-teeth and 
petiole make the Montana specimens distinct. Some of the deeply dissected, 
pointed lobed, ± delt to rhomb-ovate leaved forms from the western US appear 

2. Crataegus douglasii Lmdl, Bot. Reg. 21: pi. 1810. 1835. 

Bushes 4-6 m tall, thorny; thorns ± stout, 1.5-3.5 cm long, straight or slightly 
recurved, deep brown and shiny when young; one-year old twigs deep tan to 
dark mahogany, somewhat shiny when young, older branches gray to dark gray, 
often showing brown or alternatively tan-brown eastwards. Leaves deciduous; 
petioles 0.75-1.5 cm long, slightly glandular and pubescent in youth; blades 4- 
7 cm long x 2-3 cm wide, somewhat variable in shape but generally elliptic to 
broad-elliptic or sub-rhombic, generally with 2-4 coarse lobes per side, alter- 
natively elliptic-obvate with much smaller lobes; margins coarsely and sharply 
toothed, the teeth tipped with small glands when young; venation 
craspedodromous with 4-5 lateral veins per side; quite densely appressed short- 
pubescent above, below usually glabrous except on the veins. Inflorescences 
10-25 flowered; branches usually glabrous, bearing caducous, membranous. 

linear, gland-margmed bracteoles. Flowers 12-15 mm diam.; hypanthium ex- 
ternally glabrous; the calyx-lobes very short, 3-4 mm long, fairly broad trian- 
gular, adaxially glabrous, the margins with a few tiny glands distally; petals ± 
circular, white; stamens 10, anthers pink; styles 3-4. Fruit 6-8 mm diam., elhp- 
soid, dull black when fully ripe, though vinous when younger, generally very 
bloomy until maturity; calyx-lobe remnants short, blunt, ref lexed; nutlets 3-4, 
dorsally grooved, laterally excavated. 

Douglas hawthorn is a common, somewhat variable bush of southern and 
western British Columbia up to the panhandle of Alaska which ranges south 
to northern California (Shasta Co.) and inland to the Rocky Mountains of Idaho, 
Montana and sw Alberta. Further east, disjunct populations occur in the Cy- 
press Hills (AB & SK), Black Hills (WY & SD), and around the northern Great 
Lakes. In western North America Crataegus douglasii is found mainly near 
water in drier areas but in more mesic localities in a good variety of open wood- 
land. In the Great Lakes area it mainly occurs m old pastures and on fence lines. 

Crataegus douglasii is distinguished from C suksdorfii which has 20 sta- 
mens, semi-camptodromous venation and little lobed, ± obtuse leaves, and from 
the taller C. okennonii which has generally shorter thorns, usually straight 
single trunk, ampuUiform-orbicular fruit and crimson-lake autumnal foliage 
color Crataegus douglasii is mainly tetraploid (Brunsfeld et al. 1990). 

3. Crataegus okennonii J.B, Phipps, Sida 18:170. 1998. 

This species commenced flowering 10-14 days later than C douglasii in the 
northern Okanagan in 2002. 

4. Crataegus castlegarensis J.B Phipps & O'Kennon, sp. nov (Fig. 3). Type; UNITED 

STATES. Idaho. Lemhi Co.: US 93, E side, 14.5 rd. mi N of Salmon, roadside ditch next to 

irrigated field, alt. ca. 3950 ft, bush, 5 m tall, foliage dull, dark, coriaceous, fruit strongly 

clustered, red-burgundy with short pedicels, 20 Aug 1996,J.B. Phipps & O'Kennon 7396 

(holotype: UWO; isotypes: BRIT CAN, MO, MONTU, TRT UBC, Y WS). 

Frutices, 2.5-5.0 m alti, uno vel pluribus truncis erectis; rami pa rentes; ramuli unius anni brunnei; spinae 

2-3 cm longae, mediana crassitudine, rectae vel leviter recurvatae, nitenter atrobrunneae, apice atro in 

juventute. Folia decidua; petioli 0.75-1.5 cm longi, varie pilosi, eglandulosi; laminae 3.5-6.0 cm longae, 

oblanceolatae vel ovato-rhombeae in forma generali; apex late triangularis, basis late cuneata vel 

cuneata; vi.x lobatae vel 3-4 lobatae per latus; aliquantum coriaceae; venatio craspedodroma, 4-5 

Shrubs, 2.5-5.0 m tall with one or more erect stems; branches spreading; 1-year 
old twigs brown; thorns 2-3 cm long, of medium stoutness, straight to slightly 

recurved, shmy dark or reddish-brown with blackish tips when young. Leaves 
deciduous; petioles 0.75-1.5 cm long, eglandular, variably hairy; blades 3.5-6.0 
cm long, oblanceolate to ovate-rhombic in general shape; the tip broad-trian- 
gular, base broad-cuneate to cuneate; scarcely lobed to 3-4 lobed; margins 
toothed, teeth with fine gland-tips when young; venation craspedodromous, 
each side 4-5 nerved; conspicuously appressed hairy above when young, some- 
what glabrescent except on the mid-nerve above, thinly pubescent to glabrous 
below except on the veins, somewhat coriaceous. Inflorescences 8-20 flowered; 
branches thinly to densely pubescent, bearing caducous, linear, membranous, 
gland-margined bracteoles. Flowers ca. 12-15 mm diam.; hypanthium gener- 
ally glabrous externally; calyx lobes 3 mm long, triangular, remotely glandu- 
lar-serrate; petals ± circular; stamens 10, anthers pink; styles 3-4. Fruit 10-12 
mm diam., ± orbicular, often somewhat oblate, recessed at junction with pedicel, 
thinly pilose, crimson turning to reddish-plum or ultimately blackish purple; 
calyx-lobes patento-ref lexed, + erose; nutlets 3-4, dorsally grooved, lateral faces 
usually pitted. 

Crataegus castlegarensis has a wide range (Fig. 4) from the John Day (South 
Fork) R. basin of Oregon and the Okanagan of British Columbia eastwards to 
the Flathead drainage of northwestern Montana, thence south down the moun- 
tains to northeastern Utah and west-central Wyoming. We have also recently 
recorded it from the Cypress Hills of Alberta and Saskatchewan but those speci- 
mens are not cited. It is most abundant north of Kelowna, British Columbia 
and in the Flathead drainage m Montana and is found in brushy places in the 
valleys, particularly near ditches and natural waterways. Fine specimens of this 
species occur on the grounds of Selkirk College, Castlegar, British Columbia, 
where several were collected for us by Peter Wood, then a professor at that insti- 
tution. It was here that the authors first became aware of the distinctness of 
this taxon and we are pleased to name it after the town of Castlegar. 

Like a good number of Crataegus species from the Pacific Northwest C. 
castlegarensis has been collected but ovedooked and the few specimens that we 
have encountered other than our own col lections have been variously assigned 
to C. douglasii and C. columhiana. Crataegus castlegarensis is apparently the 
most closely related of ser Douglasianae to C. douglasii. It differs in the fruit 
shape (± orbicular/turbinate in C. castlegarensis; ellipsoid in C douglasii), fruit 
color at comparable period of ripening (often dull plum blotched green early, 
then burgundy to crimson, finally deep purple and not very bloomy in C 
castlegarensis; starting about the third week of August as for the former species, 
usually already full black and remaining very bloomy unless abraded in C 
douglasii); inflorescence branches and hypanthium, also, often fruit, proximally 
(pubescent to dense pubescent in C castlegarensis; glabrous in C douglasii). 
Pubescence of parts may abrade through the season and one may have to look 

very carefully to discern it on fruiting pedicels. The junction of tfie fruit to the 
pedicel, often recessed like an apple, is also a characteristic of the new species. 
It came into flower about a week later than C. douglasii in the northern 
Okanagan in 2002. 

A very interesting feature that we record is branched thorns (double to 
quadruple) on the young twigs. This feature is not known to us in other haw- 
thorns except for the mutation of C monogyna, cv Hornda and occasional speci- 
mens of C enderhyensis. On some of the bushes this multiple thorniness was 
abundant, in others more sporadic. We did not deliberately collect for this fea- 
ture, indeed were not conscious of its relationship with C. castlegarensis early 
on, so it is possible that a more thorough check m the field of many of the col- 
lected bushes may have yielded more material like this. Branched thorn speci- 
mens are asterisked in the list of e 

Drive, ca. 10 ft down the embankmt 

' (BRIT, CAN, ID, MONTU, RM, UBC, UT, UWO, V). Pleasant Valley Rd. NNE of Vernon, E side of 

forked thorns, 25 Sep 1993J.B. Phipps & R.]. O'Kennon 6818' (BRIT, UWO). Castlegar, Selkirk College, 
top of wooded steep slope near Peter Wood's office, alt. 1000 ft, collection of different leaves from 4 m 
bush, 27 Sep 1993, J.B. Phipps&RJ. O'Kennon 6825 (UWO). Okanagan Valley, ca. 8 km NNE of center 
of Vernon, Pleasant Valley Rd., behmd Vowle's residence, hedge at S boundary of Burke's PYO, alt. 380 

94, J.B. Phipps 6881* (BRIT, UBC, UWO). Kalamalka L. Provincial Park, along road ca. 25 m SW of 
d gate, brushy roadside, alt. ca. 1450 ft, bush, 5 m tall, fruit burgundy turning black, foliage large 

ilamalka Provincial Park, roadside near red gate, bush, 5 m tall, fruit burgundy ± spherical to short 
rbinate, locally abundant, 26 Aug 2000, J.B. & S.K Phipps 8187* (UWO). Castlegar, top of Frank 

ish. 5 m stamens 5 anthers small pmk {-jBP & RO K 6823) 06 May 1994 J B Phipps 6925 (UBC 
vVO) Castlegar top of Frank Bemder Rd to Selkirk College opposite airport bush 2 m tall half 
ly up slope before curve road sign stamens 5 antheispmk 06 May 1994 Phipps ]B 6926 (UWO) 
' rd mi SW of Princeton on BC3 edge of Typha marsh on SE side of hwy alt 2800 ft bush 3 5 m 
11, fruit orbicular burgundy beginning to be taken by waxwmgs Ivs thm coriaceous shiny i=JBP 
RO'K6774) 17 Aug 1994 J B Phipps & RJ Ktnnon 6972 (BRIT CAN MONTU TRT UBC UWO 
.Castlegar Frank Beinder Drive scrubb) slopes with aspens ca 250 m E of campus gate alt Ti50 
tree 6 m tall fruit ± orbicular reddish plum to deep pm pie 23 Aug ] 994 J B Ph I pps (S R / K c n non 
53 (BRIT UBC UWO) Castlegar Fiank Bcindei Drue scrubby slopes ^Mth isptns t i / ^ m F ot 
mpus gate & behind pine alt 1550 ft bush 2 5m tall with purple black globost liuit ^3 \u,,IW4 
.Phippi&RJOKennon 7055 (BRIT UBC UWO) Castlegar Frank Beinder Drixc opposite mpott 

-, turning burgundy from reddish plum 23 Aug 1994 J 6 Pliipps&RJ OKennon 7054 (BRIT UWO 
. Ca 8 road mi SW of Princeton on BC3 edge of large dried up marsh on E side of hwy bush 10 ft 

,sh,4mtall fruit deep burgund\ ± spherical 20 Augl994 JB Phipps&RJ OKennon7030(UWO) 
rnon Y Lake open aspen groxc ncn pnnd 10 Mn 1970 1 M Bray^ha^^ & D Laidmansn* {VVJO 

road, fenceline again 
of flowers flowers c 
(equals J6P 7396) 17 


3 Aug 1996 J B Phippi ^RJOKennon 7395 (ID UT UWO) US 93 17 4rd mi 
oadside ditch alt 3900 ft bush 2 m tall fruit dull purple red 20 Aug 199 
mon 7397 (UWO) Payette Co Hwy 53 roadside m f loodplam of Payette Rive 

Payette, alt. 2300 ft, slightly swampy fields, 27 Aug 1994, J.B. Phipps & R.J. O'Kennon 7094 (UWO). 
Washington Co.: US 95 at Monroe Creek, ca. 5.6 mi N of Weiser (MP 88.4), alt. ca. 2700 ft, brushy area, 
bush, 5 m tall, fruit large, spherical, red changing to burgundy, 27 Aug 1994,J.B. Phipps & R.J. O'Kennon 
7095 (BRIT, ID, MONTU, RM, UT, UWO). MONTANA: Flathead Co.: Hwy 35, ca. 0.5 mi SE of Creston, 
alt. 3050 ft, dense bush 3 m tall, fruit orbicular, dull red, 25 Aug 1994, j.B. Phipps & R.J. O'Kennon 7071 
(MONTU, RM, UWO). Junction of state road 35 and unpaved road to Fish Hatchery, small hawthorn 
thicket in old pasture, alt. ca. 2800 ft, bush, 3 m tall, flowers at popcorn, anthers pmk, 20 May 1997. 
J.R Phipps (SRJ. O'Kennon 7589 (MONTU UWO) Fhthead River Valley 1 mi E of 206 ca 3miSEof 
Columbia Falls center on dirt extension of S end Berne (Jensen) Rd 3100 ft large 6 m bush near 
pond, (-JBPcS-RO'K #6838) stamens 10 anthers pmk coll 8Ma> budsloiced 08Ma> 1994 JB Phipps 
6938 (BRIT ID, MONTU RM UBC UWO) Flathead River \allc> 1 mi F ot 206 3 m SE Columbia 
Falls center at extreme S end of Jensen Rd by pond alt 3100 ft 5 m bush Iruit orbicular purple 
black, 27 Sep 1993, J B Phpps&RJ Kennon 6838 (BRU ID MONTU RM UBC UWO) Lake Co 
State Road 35 E of Poison between Turtle Lake Road and Ducharmc Indi m Agenc> lands with stream 

some bi-thorns, in bud anthers pale pink 19 May 1997 JB Phipps <S RJ O Ktnnon 758J*(UWO) Old 
Freight Rd., off US 93 W of St Igmtius haw thorn thickets in creek beds between dr> grassland at 

7409 (BRIT MONTU RM UWO) Old Freight Rd oifUS93W of St ignunis haw thorn thickets m 
creek beds between dr> grasshnd at roadside \h 2900 It bush 4 m till leaves rhomb ovate little 
lobed, badly rusted, fruit broad ampulliform tosuborbiculate ledtopuiple bhck common 21 Aug 
1996,J.B.Phipps&RJ OKtnnon 7410 (UWO) ci 5 km up Old Freight Rd lromUS93 hawthorn 
thickets along creek ah 3000 ft 17 May 1997 J B Phipps & RJ Km non 7552 (MONTU UWO) Old 
Freight Road off Hwy 93 withm sight of Hw) 93 milepost 30 4 3 mi N ol Ravalli 10 pmk anthers 
15-22 flowers per inflorescence pedicels pubescent calyx glabrous medium thorns 18 May 1995 
RJ. O'Kennon 13162 (UWO) Old Freight Road oil Hwy 93 withm sight of Hwy 93 milepost 30 4 3 

of Hwy 93, milepost 30 4 3mNof Ravalh 17-22 flowers per inflorescence 10 pmk anthers full flower 

tween Roosville & Eureka W side ol rd ca I mi N ol lunction with 37 seepage slopes below in iga 
tion channel, on hillside ah 2850 ft bush 4 m cius galh like toliage lit orbicular crimson chang 
ing to burgundy 24 Aug 1Q04 J B Phipps &RJ0 Kennon 7062 (BRIT MONTU UWO) Sanders Co 

of center of Camas Prriric B^sin on road trom Little Bitterroot Valley close to Schmilz Lookout ac 
cess road, seepage zone m pi airie alt 3000 ft bush 4 m tall anthers 10 pmk 18 Ma) 1997 JB Phpps 
& R.J. O'Kennon 7576 (BRIT MONTU UWO) at Little Bitterroot R at eastbound road crossing ex 
actly20kmSEofHotSprmgs E\ m Melton ranch thickets at edge o( watet alt ca 2800 ft bush 4 m 
tall, anthers 10, pink 18Ma\ 1997 J B Phipps fv RJ OKtnnon 757KBRIT MONTU RM UWO) Ca 24 
km (direct) SE of Hot Springs hawthorn thickets along banks ot Little Bitterroot R alt 2800 ft 

UWO); ca. 24 km (direct) SE of Hot Springs hawthorn thickets along banks of Little Bitterioot R 

MONTU, UT, UWO) hawthoin thickets with pasture along Little Bitterroot R ca 24 km SE ol Hot 
Springs, Melton Sr. (grandfather) ranch, alt 2750 ft, bush 3 m tall, leaves broad, fruit burgundy oi 
bicular, Aug 21 1996, J.B. Phipps & RJ. O'Ke n no n 7423* (UWO); hawthorn thickets with pasture along 
Little Bitterroot R., ca. 24 km SE of Hot Springs, Melton Sr (grandfather) ranch, alt 2750 ft, bush 3.5 m 
tall, foliage intermediate, fruit burgundy orbicular, 21 Aug 1996, J.B. Phipps & R.J. O'Kennon 7425 

Little Bitterroot R., abt 24 km direct SE of Hot Springs, 2800 ft, bush, 3 m tall, 18 May 1997 J.B. Phipps 
& RJ. O'Kennon 7566 (BRIT, MONTU, UWO); hawthorn thickets along bank of Little Bitterroot R., 
abt 24 km direct SE of Hot Springs, 2800 ft, bush, 3 m tall, 18 May 1997, J.B. Phipps & RJ. O'Kennon 
7568 (BRIT, MONTU, UWO); crossing of Little Bitterroot R. exactly 20 km SE of Hot Springs, thickets 
at edge of water, Evan Melton ranch, 2800 ft, bush 4 m tall, 18 May I997,J..B. Phipps & RJ. O'Kennon 
7573 (BRIT, MONTU, RM, UWO); mam road SE from Hot Springs along Little Bitterroot River at 5 mi 
due N of major NS/EW bend (near location of 7420) at Malmak mailbox, alt 2800 ft, two hawthorns 
along ditch on west side of road, healthy bush, 4 m tall, 21 Aug 1996,J.B. Phipps & R.J. O'Kennon 7427 
(UWO); Hot Springs-Perma Rd., Camas Prairie Basin, ca. 4 mi S of Camas Prairie, alt. 2850 ft, large 
bush 6 m tall, fruit burgundy-red, 21 Aug 1996, J.B. Phipps & RJ. O'Kennon 7429 (UWO); Little Bitter- 
Evan Melton ranch, alt. 2790 ft, open thicket of hawthorns in pasture near creek, bush 3 m tall, leaves 
blue-green, fruit deep burgundy orbicular, with recessed pedicel, locally frequent, 21 Aug 1996, J.B. 
Phipps & RJ. O'Kennon 7420 (MONTU, RM, UWO). OREGON: Grant Co.: John Day Valley S fork, 
Humphreys, small tree, 30 Apr 1925, LF. Henderson 5146* (CAS, DS, photo UWO). UTAH: Cache Co.: 
S of Hyrum Reservoir at 7800, off 400S, fenceline, S side of dirt road, alt. 4800 ft, bush, 5 m tall, small 
flowers, 10 pink anthers, 15 May 1997, J.B. Phipps & RJ. O'Kennon 7528 (BR1T,1D,UT,UW0); E side of 
south end of Hyrum Reservoir across field from intersection of 7000W and 400S, hedgeline, alt. 4800 
ft, bush, 4 m tall, flowers small, anthers 10, deep pmk, 15 May 1997, J.B. Phipps & RJ. O'Kennon 7527 
(BRIX ID, MONTU, NY, RM, UBC, US, UT UWO). Weber Co.: Huntsville, on N side of UT 39 ca. 0.5 mi 

C rivularis, 15 May 1997, J.B. Phipps & RJ. O'Kennon 7526 (UWO); UT 39 ca. 1/2 mi after it turns E, E 
of Pineview Reservoir, roadside with ditch (water flowing) backing on to irrigated field, 2 adjacent 

orbicular, recessed at pedicel contact, green turning light to dull red to vinous to black, on same tree, 
18 Aug 1996, J.B Phipps & RJ. O'Kennon 7389 (BRIT, ID, MONTU, RM, UT UWO); ravine to E of S.R. 
167 (Trappers' Loop Rd.), along creek, alt. 5200 ft. bush 4 m tall, abundant, 22 May 1997, J.B. Phipps & 
RJ. O'Kennon 7608 (BRIT MONTU, UT). WASHINGTON: Okanogan Co.: Chopaka Rd., 2.75 mi from 
Loomis-Nighthawk Rd. junction, 28 Aug 1994, J.B. Phipps 7106 &RJ O'Kennon (UWO). Spokane Co.: 
S of Deer Park, 0.5 mi S of milepost 179 on US 395, 30 Sep 1993,J.R Phipps 6861 & RJ. O'Kennon (UWO). 
Whitman Co.: Staley Rd., 2 km NE of Chambers, f ioodplain of small creek, dense small tree 4 m tall, 
fruit orbicular, red changing to burgundy 27 Aug 1994, J.B. Phipps 7097 & RJ. O'Kennon* (BRIT, UBC, 
UWO, WS, WTU). WYOMING: Lincoln Co.: US 89, S of Thayne and 0.4 mi N of Salt River, f Ioodplain 
pastures with hawthorn thickets, alt. 5950 ft, bush, 4 m tall, inflorescence pubescent, anthers 10, 
pink, m green bud, forced, 21 May 1997, J.B Phipps & RJ. O'Kennon 7597* (BRIT MONTU, RM, UWO); 
US 89 S of Thayne at 0.4 mi N of Salt River crossing, fenceline in flood plain, alt. 5950 ft, bush, 3.5 m 
tall, (= JBP 7462), in tight bud, forced, 16 May 1997,J.B Phipps & RJ. O'Kennon 7531* (RM, UWO); US 
89, S of Thayne and 0.4 mi N of Salt R. crossing, 2.5 mi N of junction with 238, fencehnes in flood- 
held erect, 25 Aug 1996, J.B, Phipps & RJ. O'Kennon 7462 (BRIT. UWO). Teton Co.: Targhee National 
Forest. US 89. 5.4 mi S of Hoback Jet., flats of Snake River, alt. 6200 ft, dense brushy area with tall 

plum to burgundy 25 Aug 1996, J.B. Phipps & RJ. O'Kennon 7460 (BRIT MONTU, RM, UT UWO). 
Targhee National Forest, flats of Snake R., 5.4 mi S of Hoback Jet., alt. 6200 ft, 21 May 1997,J.B Phipps 
7593a* & RJ. O'Kennon (UWO). Washakie Co.: US 16, 1 mi E of Tensleep, river flats of Tensleep R., in 

fruit suborbicular, plum-colored, 24 Aug 1996, JB. Phipps G'RJ. O'Kennon 7456 (UWO). 

Incertae Sedis 

Crataegus shus wapensis J B Phipps & O'Kennon, sp nov (Fig. 3) T\pf CANADA 
British Coi umbia Northern Okanagan Endeib), ca 200 m E of budge over 
Shuswap Rivei on Mabel Lake Rd bush, 3 5 m tall Iruit deep purple ^=JBP 69101 
20 Aug 1994JB Phipps & RJ O'Ktnnon 7009 [\\o\o\\\'\ UWO, iS0T\rES BRIT, 

dorsahter sulcatae, m latenbus planae 

Bushes to 5.5 m tall; one year old twigs tan to deep brown; thorns 2-3 cm long, 
stoutish, straight or slightly recurved, very dark brown when young. Leaves 
deciduous; petioles 1.5-2.25 cm long, sparsely gland-dotted, slightly pubescent; 
laminas 5-7 cm long, rhombic in general shape; ca. 4-lobed per side, LIl to 25%; 
venation craspedodromous, with 4-5 pau's of secondary veins; lobes sharp; 
margins remotely serrate; somewhat sparsely appressed-pubescent above, gla- 
brous below. Inflorescences 12-18 flowered, the branches glabrous, some linear 
caducous gland-margined bracteoles present. Flowers ca. 15 mm diam.; hy- 
panthium glabrous externally; calyx-lobes 4-5 mm long, triangular, minutely 
gland-margined; petals chcular; stamens 10(15-18), anthers pale to bright pink 
or rarely puce; carpels and styles 2-3. Fruit ± broadly ellipsoidal, 12 mm long, 
claret color at first, deep purple at maturity; calyx remnants present, patento- 
recurved; nutlets 2-3, grooved dorsally, smooth on sides. 

Crataegus shuswapensis is locally common near Enderby in the northern 
Okanagan and southwestern Shuswap drainage, British Columbia. Collection 
localities may be identified, at least approximately on figure 2. 

This is a very distinctive hawthorn on account of its rhombic, wide and 
quite deeply and sharply lobed leaf-blades, stamens with anthers pink or some- 
times puce and fairly large suborbicular to broad-ellipsoid purple fruit. The 
flowers, and to some extent the fruit, are much like C douglasii but other char- 
acteristics, the rather long stout thorns, leaf shape, fruit color about the third 
week in August, 2-3 styles and nutlets point m a different direction. Flowering 
material with 10 pink stamens from near Poison, Montana, may be the same 
and fruiting specimens are required to verify this. This species flowers synchro- 
nously with C.castlegarensism the northern Okanagan. 

Fig. 5. Line drawing of Crataegus shuswapensis. Flower details 

of Enderby, little to E of Brash Creek, road-edge, woodland S side of road, shade, alt. 1270 ft, 4 m tall 
large bush in shade of balsam poplar, black fruit, 20 Aug 1995, J.B. Phipps 7172 (UWO); Northern 
Okanagan. Spallumcheen Municipality, Marshall Road (dead end gravel drive off Mcleery Rd.), alt. 
1300 ft, bush, 2.5 m tall, fruit deep plum-purple, ellipsoid, 20 Aug I995,i.B. Phipps 7169 (BRIT, UBC, 
UWO, V); Mabel L. Rd., N side, ca. 150 m E from Shuswap River bridge, 3-3.5 m tall, anthers exhibit 
slight pmk tinge, 15 May 1995, IS. & D. Donovan 2755 (UWO); Near Enderby S end of 97 A, E side of 
Hwy 97A, ca. 150 m Irom Mabel L. Rd., forest margin, alt, ca. 1250 ft a.s.l., ca. 2.5 m tall, 15-18 pink 
anthers, spindly leaves larger, lighter green (shady location?), 15 May 1995, LS. & D. Donovan 2754 
(UWO); Near Enderby S end of 97A, E side of Hwy 97A, ca. 150 m from Mabel L. Rd., forest margin, 

2753 (UWO); S end of 97A, near Enderby, E side of Hwy 97A, ca. 150 m from Mabel L Rd., forest 
margin, alt. ca. 1250 ft a.s.l., ca. 1.75 m tall, 15-18 stamens, deeply lobed leaves, anthers pmk, 15 May 
1995, LS. & D. Donovan 2752 (UWO); Rd. to Mabel L. ca. 1.2 mi E of Enderby and between 25 m and 
200 m W of "Autobody collision and repairs" sign, hedgerow, alt. 1275 ft, bush 3 m tall, fruit bur- 
gundy-red, 20 Aug f 994,J.B. Phipps & R.]. Okennon 7015 (BRIT, UWO); 1.25 mi E of Enderby on Mabel 
L. Rd., ca. 200 m E of "Autobody Collision, Repairs & Painting" sign, 1275 ft, hedgerow, bush, 5.5 m 
tall, 20 Aug 1994, J.B. Phipps & R.]. O'Kennon 7018a (BRIT, UWO); Shuswap Valley, ca. 8 km E of cen- 
ter of Enderby on road to Kingfisher, S side of road, 1270 ft, hedge backing small woodlot, bush, 4 m 
tall, stamens 10, anthers pmk, (-]BP & RO'K 6815\ 05 May 1994, J.B. Phipps 6922 (BRIT CAN, UBC, 
UWO, V); Shuswap \ alley road to Kmgl isher, ca. 2 km E of Enderby opposite sign on N, "Autobody 
Collision, Repairs .'v ramting', alt 1275 It, hedgerow S side of road, bush 5 m, ca, 18 puce anthers, 

medium, dark green foliage, stamens 15, anthers pale pink, 05 May ig94,J,B Phipps 6910 (BRIT, UWO); 
Northern Okanagan, Spalumcheen Municipality Stepney Cross Rd„ lowest point, large bush, 4 m 
tall, 26 Aug 2000, J,B. Phipps 8182 & S.R. Phipps (UWO), 

ser, Purpureofructi J.B. Phipps & O'Kennon. 

Bushes or small trees, generally 3-6 m tall; thorns 2-5 cm long, slender to quite 
stout, shiny dark brown to purple or black at 2 yrs, usually quite numerous; 
extending twigs thmly pubescent or glabrous; at 1 year deep reddish-brown or 
mahogany, older dark gray Leaves deciduous; petioles 1/4-1/2 length of blade, 
1-3 cm long, eglandular or with some small black glands, glabrous to thinly 
pubescent while young; blades 3-8 cm long, ± rhombic to elliptic-oblong, ovate- 
rhombic or occasionally suborbiculate in general shape; acute at the apex, cu- 
neate to somewhat rounded at the base; l-3(-4)-lobed per side, the lobes ob- 
scure and shallow to well-defined and acute and with max. Lll ca, 20%; margins 
serrate except at t he base, the teeth with small gland-dots; venation craspedodromous, 
4-6(-7) per side, ± impressed in the more coriaceous-leaved species; usually 
scabrate-pubescent on the upper surfaces young, this variably glabrescent 
(abrading with age), below glabrous on the surface, pubescent or not on the 
veins; markedly coriaceous to thin; autumnal color generally brilliant. Inflo- 
rescences 8-25 flowered; branches glabrous or thinly pubescent, bearing cadu- 
cous, membranous, linear, gland-margined bracteoles. Flowers 12-18 mm diam,; 

hypanthium externally glabrous to pubescent; calyx-lobes 2-4 mm long, nar- 
row, abaxially glabrous, margins finely glandular-serrate; petals ± circular, 
white; stamens (5)10-12, anthers white or pink; styles 2-4(-5). Fruit 8-10 mm 
diam., ± orbicular to ellipsoidal or ampulliform, mostly red in late August, deep 
purple or deep reddish-purple (often nearly black) in late September; calyx- 
lobes narrow or broad, ref lexed to erecto-patent, thinly hairy to glabrous; nut- 
lets 2-4(-5), dorsally furrowed, sides plane to quite deeply pitted. 

Six species are now recognized, found from the Okanagan Valley of British 
Columbia and Washington east to northwest Montana. They are all locally com- 
mon in suitable habitats. We add three new species and one variety to this se- 
ries and emend the key. 

These include some very handsome hawthorns, particularly in late sum- 
mer and autumn when their usually abundant fruit turns from bright red to 
deep purple and the leaves manifest brilliant coloring in bronze, burgundy yel- 
low etc. These hawthorns have very high ornamental potential but are not yet 
cultivated outside botanic gardens and arboreta. 

ion above, flattish to strongly concave, generally mid-green until fall; hy- 

lanthiunn glabrous or hairy 2.C.okanaganensis [incl.var. w 

eaf-blades 4-8 cm long, softly coriaceous, with distinctly impressed ve- 
lation above, flattish to convex,generally deep green until falfhypanthium 
llabrous S.C.enderbyensis, s 

;. Club 36:64T 1909. See also J.B. 
2. Crataegus okanaganensis J.B. Phipps & O'Kennon, Sida 18T78. 1998. 


Leaf^blades 1.2-1.6 x as long as wide, sharply lobed; fruit flask-shaped 2a.var 

Leaf-bladesl.6-2 x as long as wide,shallowly lobed in distal half only or occasionally 

2a. Crataegus okanaganensis var. okanaganensis. 

2b. Crataegus okanaganensis var. wellsii J.B. Phipps & O'Kennon, var. nov. (Fig. 
6). Type: UNITED STATES: Washington: Okanogan Co.: Palmer Lake, N side; large 
bush, 5 m tall, stamens 5, anthers ivory, (equals J.B.P & R.J.O'K. 6875), 4 May 1994, 
}B. Phipps 6905 (holotype: UWO; isotypes: BRIT, CAN, DAO, UBC, US, WS). 

abaxialis glabra venis exceptis. Inf lorescentiae 8-15 florae; ramuli sparsim pilosi, bracteolis caducis, 
linearibus, membranaceis, glandulo-marginatis. Flores ca. 15 mm diam.; hypanthium supra glabrum 
et subter villosum; lobi calycis anguste-triangulares, tenuiter glandulosi; petala ± circularia; stamina 

lares, subter pilosi, sanguinei in Augusto exeunte, in maturitate atropurpurei; lobi calycis triangu- 
lares, plerumque reflexi; pyrenae 2-3, dorsaliter sulcatae, lateribus planis vel vadositer foveolatis. 
Large shrubs to 6 m tall, often with multiple trunks; 1 year old tv/igs mahogany; 
thorns 2-5 cm long, shiny dark brown when young, stout, recurved. Leaves de- 
ciduous; petioles 1-1.5 cm long, glandular above at least when young, pubes- 
cent when young; blades 3.5-5 cm long, elliptic or ellipt-oblong to elliptic-rhombic 
or elliptic-ovate in general shape; apex acute, base often slightly tapered into 
petiole or obtuse; often ± parallel-sided at broadest part, unlobed or with 1-3 
distinct lobes, distal side of angle of lobes usually ± perpendicular to mid vem; 
margins serrate; venation craspedodromous with 4-5 main secondary veins per 
side; short-appressed serrate when young above, glabrous below, except along 
veins. Inflorescences 8-15 flowered; branches thinly hairy, with early caducous, 
glandular bracteoles. Flowers 15 mm diam.; hypanthium externally glabrous 
above and villous below; calyx-lobes narrow-triangular, finely glandular; pet- 
als ± circular; stamens (5-)10-12, anthers usually ivory; carpels and styles 2- 
3(-4). Fruit ca. 10 mm diam., ± orbicular, hairy below, dull deep red at late Au- 
gust, deep purple at full ripeness; calyx-lobes triangular, usually ref lexed; nutlets 
2-3, dorsally sulcate, sides plane or shallowly pitted. 

Crataegus okanaganensis var wellsii is found in the Okanagan valley of 
British Columbia and Washington and is common around Vernon (Fig. 7). Ab- 
original people in Washington recognized this taxon as "stemoq'w" according 
to R. May's collection notes (cited specimen May 258RM from WA). 

This variety was first collected by T.C. Wells at Kalamalka Provincial Park, 

J^ - 



■ 1.114- 
x-"\ 51i 

"\ AB 



1 r\ 

(A. ^^^ 


T7 .1 

y ) 1 \ ; 







?. Distribution map of fr 

near Vernon. In its modal form it is very distinct with its unusual leaf -shape 
(quite differently lobed from var. okanaganensis). It also differs from var. 
okanaganensis in its orbicular fruit, fewer nutlets, fewer stamens (in the type), 

okanaganensis. Fruit may be a somewhat different color than nearby var. 
okanaganensis at the same time in late August, although in the third week of 
August they are brilliant scarlet. The most extreme form, virtually thornless, was 
seen at Palmer Lake, northern Washington. In spite of the several differences 
from var okanaganensis the presence of intermediates prevents a higher rank. 

hillside, alt. ca. 474 m, young 3 m bush, purple fruit. 26 Sep 1993, J.B. Phipps & R.]. O'Kennon 6819a 
(UWO). Okanagan Valley, ca 10 km N ol Vernon oil 97A NW bound along dirt road into O'Keefe 
Ranch, at L. and A. Cross Road, alt 1 270 f t, boundary hedge along ditch, on SW side, bush, 4 m tall, 
stamens 10, anthers pale pink {=J.B Ph i /)/is & R J 01\c n non 6804), 03 May 1994,J.B. Phipps 6893 (UWO). 

gate. alt. ca. 1375 ft, large bush 6 m tall, 5 ivory stamens, {--J.B. Phipps & R.J. O'Kennon 6798), 03 May 
1994, J.B. Phipps 6885 (TRT, UBC, UWO, V). Ca.lO km N ol Vernon center on dirt road N mto O'Keefe 
Ranch off 97A at L & A. Cross Road, alt 1270 ft, boundary hedge along ditch, common component, 
3.5 m bush, fruit burgundy 25 Sep 1Q93 J B. Phipps & R J. O'Kennon 6805 (UWO). Shuswap Valley just 
E of Enderby across bridge, S side ol road, top of river bank, alt. ca. 1250 ft, bush 4 m tall, m bud, 

ens 5 or 10, anthers ivory 05 May 1 

994, /.B. PI 

npps 6914 (UWO). K; 


Park. 2 


rnon, Kidston Rd. entrance at red g 

ate, with/ 





ion, 18 Aug 1987 

: #19422, UWO) KalamalkaLPr 

ov Park. 2 

km SH of Vernon, Ki 

ce at n 

:d gate. 


carpus all 


open thickets: a 

It. 550 


shiny scarlet fruit, common. 18 Au 




E of Enderby across bridge, 5 side ( 

Df road, to] 

D of river bank, alt. c 

a. 1250 ft, bush 4 


in late 

stamens 10, anthers ivory 05 May 1 

994,7 B PI 



Park, 2 


rnon, at S terminus of Kidston Rd 

red gate, ^ 

vith Amelanchier al 




s ponderosa, open thickets, 550 m 

imy scadet frt., 

5 691 (UWO, UBC 194424 J. A us 1 « 

K)R7 Shu 

..ap \ alley just Eol 

[Enderby across 



rubby slopes rich in Crataegus, bush 6 m tall, red iruit, most abundant Crataegus here, 19 Aug 
] B Phipps & R.J. O'Kennon 6999 (BRIT, UWO, V). Ca. 1 mi S of Enderby N-trending, downslope 
1 driveway bordered with hawthorns, alt. 1300ft, bush, 4 m tall, fruit burgundy 20 Aug 1994, 
tipps&R.J. O'Kennon 7006 (BRIT, UBC, UWO). Ca. 10 km N of Armstrong, W of Indian Reserva- 
dt. ca. 1550 ft, long farm drive with Crataegus hedgerow, bush 2 m tall, fruit in various stages of 
1 Aug 1994,i.B. Phipps & R.] O'Kennon 7035 (UWO). Okanagan Valley Spallumcheen Munici- 
r, L. & A. Cross Road, extreme E end, just W of Hwy 97A, alt. 1270 ft, along driveway N into farm, 
pecies abundant here, shrub 4 m, rather dense fruit, blood-red, ± spherical.J.B. Phipps 7161 (CAN, 

:msa Pme and Douglas fir forest, 2 m tall, 
JWOJ Shuswap Vallc)',just E of Enderby 


R / O'Kennon 6982 ICAN, UBC, U 

[WO, X'J.JustE of Enderby after 

bridge over Shuswap R., rough grass 

near water, S side of road, alt. 12' 

50 ft, edge of water near oxbow 

of Shuswap R, bush 5 m tall, burgi 

mdy fruit, 25 Sep 1993, i.B.Phip 

ps & R.j. O'Kennon 6809 (UBC, 


ersection with 97, scrubby slope; 

5 with long cleared area, bush 4 

m tall, frmti spherical, plum-colore 

;d, common here. 19 Aug 1994, J. 

B.Phipps& R.J. O'Kennon 6997 


, of Enderby gravel driveway to f 

alt. 1300 ft, bush, 3.5 m tall, fruit b, 

right red, ellipsoid, scarce here, 

20 Aug 199^ J.B. Phipps &rI 

O'Kennon 7007 (UWO). Ca.l mi S of 

Enderby gravel driveway to NE s 

.tarting at Indian Cemetery alt. 

I opposite house with sohc 

n em (BRIT UBC UWO) Northern O 

ogan Co US 97 1 1 km S of Flhsforde hawth 
:ed field (mobtl) C ma racanthv nit 275 m bush 4 m till den: 
g 1995 J BPIupps 7176 (UWO WS WTL) Westbide Rd ci 4 km f 

3Ktnnoif86S(BRIT UWO WS \ML ) Pnlmci I il e N s de lit 11 


erbyensis J.B. Phipps & O'Kennon, sp. nov. (Fig. 8). Type: CANADA: 

H Columbia: just E of Enderby after bridge over Shuswap R, S side of road, 

rough grass near water; alt. ca. 1250 ft, large busfr 5m tall, burgundy-red fruit, 25 

Sep 1993, J.B. Ph ipps & RJ. O'Kennon 6808 (holotype: UWO; isotypeS: BRIT, CAN, 


calycis anguste-triar 

Large bushes or small trees to 5 or 6 m tall; 1-year old twigs purple-brown, older 
dark gray; thorns 2.5-3.5 cm long, stout, recurved, shmy mahogany-brown when 
young. Leaves deciduous; petioles 1-2 cm long, ± glabrous; blades 4-8 cm long, 
ovate-rhombic to broad elliptic-rhombic in general shape; base broad-cuneate 
to ± rounded; with ca. 3 lobes per side, largest LII ca. 20%; margins finely toothed; 
venation craspedodromous, distinctly impressed, v/ith (3-)4-5 lateral veins per 
side; leathery and dark green at maturity also rather convex; appressed-pubes- 
cent above, ± glabrous below except on the ± pubescent veins, brilliantly 
colored in fall. Infloresences 12-25 flowered; branches villous, bearing cadu- 
cous, hnear, membranous, gland-margined, bracteoles. Flowers ca. 15 mm diam.; 

hypanthium glabrous externally; calyx-lobes narrow triangular, finely gland- 
margined, glabrous abaxially; petals ± circular; stamens 10, anthers cream; car- 
pels and styles 3. Fruit 8-10 mm diam., ellipsoid-urceolate, often in a large, ± 
pendant corymb, cream when young, turnmg to apricot-colored then red, at 
full maturity purple; calyx-lobe remnants narrow, ref lexed, pubescent; nutlets 
3, dorsally furrowed, their sides plane or shallowly pitted. 

Crataegus enderhyensis is presently only recorded from the Armstrong- 
Enderby region of the upper Okanagan of British Columbia where it is known 
from a good number of specimens. Collection localities may be identified, at 
least approximately, on figure 2. 

This species is easily recognised m the field bv its handsome softly leath- 
ery, somewhat drooping convex daik green foliage bnlhantly colouring m tall 
with clearly impressed venation as well as by its usuallv laige h uitmg coiymbs 
with then at fust veiy pale pendant huit which become bnght red by late 
August and finally turn purple black ItiscloseK iilucdtoC okanaganensis 

AdditiomlspecimtnbtMmmcd CANADA BRITISH COLUMBl \ NiiithtinOl m i^ui bpallumcheen 

utumnU truitpuiple 23 Sep 1993 J iJ Phipps^ RJOKtnnon 68/3 (UWO) Ca 1 7 mi E ot Enderby 

/oodland alt 1280 1 1 bush 4 m tall fruit purple lohage deep green 25 Sep 1993 ]B Phipps & R] 
)Ktnnon6812{mC UWOJ 

-. Crataegus orbicularisJ.B. Phipps 61 0'Kennon, sp. nov (Fig. 9). Type: CANADA: 
British Columbia, Northern Okanagan, Spallumcheen Municipality, Back Enderby 
Rd., ca. 4 mi SE ol Enderby, ca. 200 m N of creek crossing, alt. ca. 1450 ft, dense 

hedges E side of road, bush 4 m tall, large, orbiculate leaves, fruit pale deep red, 21 
Aug 1994 J.B. Phipps & R.J. O'Kennon 7039 (holotype: UWO; isotypes: BRIT, CAN, 

I vel plerumque 

r latus, breves, triangulares 


■ petic 




5a, ab; 


Wide-spreading bushes, 2-4 m tall; 1 year old twigs strong mid-brown, branches 
dull gray; thorns 3-5 cm long, slimmish, straight or more usually slightly re- 
curved; Leaves deciduous, petiolate; petioles 1.25-2.0 cm long, adaxially shortly 
hairy usually eglandular; blades mainly 5-8 cm long, broadly elliptic to nearly 
orbicular, the broader ones curved across the base, the narrower somewhat at- 
tenuated; venation craspedodromous, with 5-6(-7) secondary veins per side; 
with 3-4 short triangular lobes, Lll ss 10 %; margins with very small teeth ex- 
cept near the base; shortly appressed hairy above, thin-hairy below, more pu- 
bescent on veins. Inflorescences 10-18 flowered; branches sub-glabrous distally 
sparsely villous proximally bearing caducous, linear, membranous, gland-mar- 
gined bracteoles. Flowers ca. 18 mm diam.; hypanthium sub-glabrous externally; 
calyx-lobes 2-3 mm long, narrow-triangular, finely gland-margined; stamens 
10, anthers pink; carpels and styles 3-4. Fruit 8 mm diam., oblong, glabrous, 
red, later changing to plum when fully ripe, in showy infructescences; calyx- 
lobes very conspicuous, erecto-patent; nutlets 3-4, dorsally sulcate, their sides 
± plane or shallowly and smoothly pitted. 

This is a very attractive species so far only known within a few miles of 
Enderby, B.C. and easily recognized by the foliage shape, thorn-type and fruit 
(color, conspicuous calyx-lobes). Crataegus orbicukris stands out in early spring 
with its light green, tinted eau-de-nil, foliage and handsome clusters of large 
flowers. Collection localities may be identified, at least approximately, on fig- 
ure 2. 

Municipality, Back Enderby Road, hedges, E side, alt. ca. 1450 ft, same as J.B. Phipps & R.]. O'Kennon 
7039 but leaves smaller this year, bush 3.5 m tall, fruit lake. 19 Aug 1995 J.B. Phipps 7166 (BRIT, MONTU, 
UBC, UWO). Northern Okanagan, Spallumcheen Municipality, Mountain View Road, N end, E side, 
alt. ca. 1500 ft, 19 Aug 1995, J.B. Phipps 7165 (BRIT UBC, UWO, V). Ca. 10 km N of Armstrong, W of 

various shades of red, 21 Aug 1994, J.B. Phipps & R.J. O'Kennon 7036 (BRIT UBC, UWO, V). Mabel 

3. Crataegus atrovirensJ.B.Phipps & O'Kennon, sp. nov. (Fig. 10). Type: CANADA: 

British Columbia: Northern Okanagan, Spallumcheen Municipality, Otter Lake 

Cross Road, 22 Aug 2000,J.B. Phipps 8171 & S.R. Phipps (holotype: U WO; ISOTYPES: 


Frutices 2-5 m alti, caulis multiplicibus suberectis vel erectis. Rainuli elongati pubescentes, illi unius 

annus brunnei nitentes, veteriores atrocinerei; spinae 3-5 cm longae, numerosi, ± rectae vel leviter 

longitudine 15-20% illius laminarum, dense pubescentes in sulco adaxiali, in maturitate sparse 
pubescentes, eglandulosi vel cum paucis, parvis glandibus praecipue distalibus; laminae 4-6(-7) cm 
longae, anguste-ovatae ad late elliptico-rhombeas vel late ellipticas in forma generali; apex acutus 

dense patente-pubescentes. Flores 10-13 mm c 
nguste-triangulares, marginibus glandulo-se 
ntheris roseis; styli 2-3. Fructus 13-15 mm a 
Keunte,atropurpurei in mense Septembro, ± v 

Shrubs, 2-5 m tall, with multiple, mainly suberect to erect stems. Extending 
branches of the season pubescent; at 1 yr glossy brown; older dark gray; thorns 
3-5 cm long, numerous ± straight to very slightly curved, at 2 yr. glossy dark 
brown, black at the tip, older deep gray. Leaves deciduous; petioles 15-20 % length 
of blades, densely pubescent at maturity in the adaxial groove, sparsely pubes- 
cent elsewhere, eglandular to with a few small glands mainly distally; blades 
4-6(-7) cm long, narrow-ovate to broadly ellipt-rhombic or broad-elliptic in 
general shape; tip acute to subacute, base cuneate; sides with 3-4 small sub- 
acute to obtuse lobes, LII of largest sinus ca. 10%; margms finely toothed, teeth 
acute to subacute, bearing minute glands at their tips; venation 
craspedodromous, 3-5 veins per side; adaxially with scabrous appressed pu- 
bescence, abrading with age, abaxially nearly glabrous; dark green and some- 
what coriaceous at maturity, in fall turning bronze-brown. Inflorescences 8-15 
flowered; branches with a fairly dense spreading pubescence. Flowers 10-13 mm 
diam.; hypanthium externally pubescent; calyx lobes narrow-triangular, mar- 
gins glandular-serrate to glandular-laciniate; petals white; stamens 10, anthers 
pink; styles 2-3. Fruit 13-15 mm high, 10 mm wide, ± ellipsoid, burgundy in 
late August, blackish-purple a month later; variably hairy; flesh soft orange 
color; calyx-lobes very prominent, suberect; nutlets 2-3, dorsally grooved, pit- 
ted on the sides. 

FiG.10. Line drawing of Crataegus c 

Crataegus atrovirens is now known from about 20 collections and many 
more specimens on the ground from an area of about 10 miles north and south 
of Enderby, British Columbia, where it is locally common to abundant in natu- 
ral hedgerows and other brushy sites in the valley. Collection localities may be 
identified, at least approximately, on figure 2. This taxon appears quite distinct 
from other sympatric kinds and is often recognizable in habit alone. 

Of species known to date from the northern Okanagan, C shuswapensis, 
also newly described in this paper, is the one most likely to be confused with C 

atrovirens. The coloring of C. shuswapensis is similar, but it has a finer more 
wide-branching growth habit, and a glabrous mf lorescence, redder fruit in the 
fourth week of August, much smaller and more recurved calyx-lobes in fruit, 
whose margins rather than being serrato-laciniate, are merely minutely glan- 
dular. In addition, the leaves of C, shuswapensis are more sharply lobed. 
Crataegus atrovirens is probably more closely related to C okanaganensis, but 
is easily differentiated from that species m the field by the much darker color- 
ing of foliage, fruit and bark, as well as by its pink anthers. Compared to C. or- 
bicularis the fruit is conspicuously larger and more hairy in C atrovirens and is 
soft and ripe in the fourth week of August when it is hard and unripe in C. 
orbicularis. It is a dull hawthorn, sombre m appearance, contrasting markedly 
with more flashy congeners such as C okanaganensis. It appears to be more 
xeromorphic than other fairly large Okanagan hawthorns. 

Additional specimens examined: CANADA. BRITISH COLUMBIA: Okanagan Valley, NNE of Vernon, 
Pleasant Valley Rd., hedge behind Vowles residence, alt. 380 m, natural hedge, bush 3 m tall. 19 Aug 
1994, J.B. Phipps 6996 &R.]. O'Kennon (UWO). Northern Okanagan, Spallumcheen Municipality, 
Stepney Cross-Road, bush 5 m tall, 26 Aug 2000, j.B. Phipps SI84 & S.R. Phipps (UWO). Northern 
Okanagan, Spallumcheen Municipality Back Enderby Rd., ca. 1 mi N of Send, bush 5 m tall, 26 Aug 
2000,J.B. Phipps 8J81 (^ S.R. Ph (pp,s (UWO, V J. Northern Okanagan, Spallumcheen Municipality Back 
Enderby Rd., 1/4 mi from S end, bush, 2,5 m tall. 26 Aug 2000,J.B. Phipps 8179 & S.R. Ph:pps (UBC, 

tall, J-B- Phipps 8180 & S.R. Phipps (CAN, UWOJ. Northern Okanagan, Spallumcheen Municipality 
Marshall Rd., bush, 4 m tall, j.B. Phipps 8186 & S.R. Phipps (UWO). Northern Okanagan, Hwy 6 E of 
Vernon, ca. 2 mi W of Lumby, bush, 12 ft tall, 26 Aug 2000, J.B. Phipps 8188 & S.R. Phipps (UWO). 
Shuswap Valley, ca. 3 mi NNE of Enderby Anderson Rd., bush, 3 m tall, 26 Aug 2000, J.B. Phipps 8185 

tall, coriaceous leaves, deep green, fruit purple, abundant here, 20 Aug 1994,J.B. Phipps&R.J. O'Kennon 
7008 (UWO). 

6. Crataegus phippsii O'Kennon, Sida 18:185. 1998. 
The line drawings were prepared by Susan Laurie-Bourque of Hull, Quebec who 
is thanked by the authors for the beauty of her realisations and her meticulous 
attention to detail. Susan Laurie-Bourque also artistically improved the rather 
crude output of Range-Mapper by using Corel Draw. The work was largely 
funded by grants to the first author by the National Sciences and Engineering 
Research Council of Canada. We also wish to thank Laurie Donovan of 
Okanagan University College, Vernon, British Columbia, for collection of criti- 
cal flowering material at times when we were unable to be present and Antony 
Littlewood, Dept. of Classical Studies, The University of Western Ontario, for 
critically checking the Latin diagnoses. 

Brunsfeld, SJ, and F.D. Johnson. 1 990. Cytological, m 

logical support for specifc status of Crataegus 

Phipps, J.B, 1 998. Introduction to the red-fruited hawthorns [Crataegus, Rosaceae) of we 

ern North America. Canad. J. Bot.76:1863-1899. 
Phipps, J.B. and R.J. O'Kennon. 1998. Three new species o^ Crataegus (Rosaceae) from we 

ern North America: C o^ennon//, C.o/(onogc?nens/s and Cphippsii. Sida 18:169-191. 
Taylor,T.M.C. 1973. Crataegus. ln:The rose family of British Columbia. British Columbia P 

vincialMuseum,Dept.ofRecreation and Conservation. Handbook no.30:26-32. 


Guy L. Nesom 

Watson (1977) described Xylorhiza glahriuscula Nutt. var. linearifolia TJ. Wats, 
from the region of Moab in east-central Utah. Cronquist (1994) observed that it 
"appears to form a distinctive local taxon with a range more than 60 km long" and 
treated it at specific rank, but he placed it within a broadly defined Machaeranthera. 
Welsh et al. (1993) followed Watson's treatment. For the treatment of Asteraceae in 
the developing Flora of North America, this taxon is recognized at specific rank 
and the following combination is required to place it within Xylorhiza. 

Xylorhiza linearifolia (T.J. Wats.) Nesom, comb, nov Basionym: Xylorhiza 

glahriuscula Nutt. var. lineariJoliaT]. Wats., Brittonia 29:215. 1977. Machaeranthera 

linearifolia (T.J. Wats.) Cronq., Intermountain Fl. 5:274. 1994. Type: UNITED 

STATES. UTAH. Grand Co.: 6 mi NW of Moab on US Hwy 160, 21 May 1971, TJ. 

Watson 679 (holotype: TEX!; isotypes: COLO, GH, MO!, MONTU, NY, UC). 

Xylorhiza glahriuscula, X. linearifolia, and X. venusta (M.E.Jones) Heller are 

closely morphologically similar among themselves and presumably are closely 

interrelated. All three have diploid populations; X. venusta and X. linearifolia 

also have tetraploid populations (Watson 1978). Within X. linearifolia, the two 

cytotypes "grow intermixed over the small range of the taxon" (Watson 1978); 

four populations of diploids and four of tetraploids were found, all within Grand 

County, Utah. According to Stockton (1983), the diploids and tetraploids of X. 

linearifolia differ in minor ways, but the present study (following Watson) finds 

no variation that would require further nomenclatural differentiation. 

Watson (1977, p. 215) observed that Xylorhiza Hneari/olia (as var Hnean/olia) 
"overlaps morphologically" with X. glahriuscula (var. glahriuscula) but the 


Fig. 1. Geographic 

of the species ranges were derived from d; 

rilyWatson (1977)andAlbeeetal.{1988).The Arizona record forXi'enuifow 

Co., Lee's Ferry, 6 Jun 1927, Coffom C26 7 7 (LL!). 

present study notes that X. linearijolia similarly overlaps with X. venusta. 
Xylorhiza linearijolia differs from both taxa m its leaf morphology, as outlmed 
by the following key. 

1. Leaves linear-oblong, truncate or rounded-aurlculate at the base Xylorhiza linearifolia 

1. Leaves oblanceolate to oblong-spatulate, tapering at the base. 

2. Stems leafy for more than 2/3 of their length, peduncles 2-6 cm long; leaves 

2.5-8 mm wide; involucres 7-14 mm long, 12-25 mm wide Xylorhiza glabriuscula 

2. Stems leafy in the lower 1/2-2/3 of their length, peduncles 6-20 cm; leaves 2.5 

Xylorhiza glabriuscula and X. venusta have slightly overlapping ranges in 
northeastern Utah and northwestern Colorado (Fig. 1) and intermediates, pre- 
sumably of hybrid origin, occur in that region (Watson f 977). The range of X. 
linearifolia is essentially imbedded within that of X. venusta, but no evidence 
of hybridization between these two taxa has been reported. Xylorhiza linearifolia 
apparently is reproductively isolated from both of its close relatives. 

In contrast to the isolation of Xylorhiza linearifolia from X. glabriuscula 
and X. venusta, the range of X tortifoha var imherbis (Cronq.) TJ. Wats, con- 
tacts that of X. linearifolia from the south and these two taxa grow intermixed 
within a few localities. "In these zones of contact, intermediates (suggesting 
hybridization) are found" (Watson 1977, p. 215). 

Stockton (1983) noted that Xylorhiza linearifolia is intermediate in m 
phology and habitat between X. glabriuscula sensu stricto and X. tortifoha ^ 
imberbis and suggested that X. linearifolia may be "a stabilized hybrid deriva- 
tive" between them, presumably from an earlier time when the putatively pa- 
rental species were sympatric. Further evidence that would test this hypoth- 
esis is lacking. 


I am grateful to the staffs at MO and TEX-LL for help during rec 

ent visits there 

and to John Semple and Eric Lamont for review comments. 


Albee, B.J., LM. Shultz, and S. Goodrich. 1 988. Atlas of the vascular plants c 

f Utah. Utah Mu- 

seum of Natural History, Univ of Utah, Salt Lake City. 

Cronquist, A. 1 994. Intermountain flora.Vol. 5, Asterales. New York Botanic 

al Garden, Bronx. 

StocktonJ. 1983.The origin of Xylorhiza glabriuscula var. linearifolia (Asteraceae). Amer. J. 


Watson,T.J., Jr. 1 977.The taxonomy of Xylorhiza (Asteraceae-Astereae). Britt 

3nia 29:199-216. 

Watson,T.J.,Jr. 1978.Chronnosome numbers in Xy/or/i/zoNuttall (Astera 

eae - Astereae). 

Madrono 25:205-210. 

WbLSH, S.L, N.D. Atwood, S. Goodrich, and LC. Higgins (eds.). 1 993. A Utah flo 

a (ed. 2, revised). 

Print Services, Brigham Young University, Provo, Utah. 

B.G. Bowes. 2000. A Color Atlas of Plant Structure. (ISBN 0-8138-2693-4, pbk.) 
Iowa State Press, 2121 State Avenue, Ames, lA 50014, U.S.A. (Orders: 1-800- 
862-6657, 1-515-292-3348 fax; $49.95, 192 pp 
numerous color plates, 7 3/4" x 10 1/4". 

student of plant 

biology However, whh the adven 


y and physiology many students 

include a course, 

This book f 

provides a long-awaited solution t. 

tasks for instruct 

ors teaching beginning botany st 

anatomy using r 

nicroscopic cross-sections of the 

's inexperience in using microsco 

teaching many u 

Bryan Bowes' Color Atlas of Plant Structure 




Botanical Research Institute 

Pseudognaphalium biolettii A. Anderb. [nom. nov.], Opera Bot. 104:147. 1991. 
Gnaphalium hicolor Bioletti, Erythea 1:16. 1893 [non Gnaphalium (Helichrysum) 
bicolor (X-indl.) Schukz-Bip., Bot. Zeit. 3:171. 1845; nee Gnaphalium bicolor Franch., 
J. Bot. (Morot) 10:411. 1896]. Typh: U.S.A. California. Without designating a type, 
Bioletti (1893, p. 17) noted that "The description is drawn up from dried specimens 
from San Diego and from a plant growing in the Garden of Native Plants at the 
University of California. ... Common in low thickets among the coast hills about 
San Diego, according to Prof. Greene. ..." At UC, there are at least two specimens of 
Gnaphalium bico/or that might represent the material referred to by Bioletti: San 
Diego, 25 Mar 1891, G.W. Dunn sn. (UC 31964); San Diego Co, Witch Creek, 1893, 
R.D. Alderson s.n. (UC 31965). Two ND-G specimens, however, apparently more 
cleariy represent authentic type material-they are labeled "Gnaphalium bicolor, 
Bioletti, Erythea 1.16. Berkeley, Bot. Garden, 1892," without other information (lec- 
totype, designated here: ND-G 59203, photocopy!; isolectotype: ND-G 59205, pho- 
tocopy!). These two large branches were broken above the base, without roots, and 
their similarity suggests they may have been collected from a single plant. The 
morphology of the ND-G types indicates that this species consistently has been 
identified correctly 
Perennial herbs, taprooted, fragrant, the stems 2-7(-12) dm tall, much-branched 
at the base and commonly lignescent, eglandular, persistently tomentose above, 
glabrescent or not below. Leaves oblanceolate to oblong-oblanceolate, mostly 
1.5-5(-8) cm long, 4-10(-15) mm wide, the upper becoming lanceolate, widely 
to narrowly auriculate-clasping, not decurrent, strongly bicolored, bright green 
and usually densely glandular above, persistently white-tomentose beneath, 
sometimes the upper leaves glandular but those toward the base eglandular. 
Capitula 5-5.5(-6) mm high; phyllaries shiny opaque-white, sometimes slightly 
pinkish, strongly graduated, apex rounded to nearly truncate, apiculate. Pistil- 
late flowers 41-73. Bisexual flowers 5-13. Cypselae shallowly ridged, not papillate. 
Pseudognaphalium biolettii occurs in the southern/southwestern half of 
California and in adjacent Mexico (Baja California and Baja California Sur), at 
elevations of 5-1200 meters (under 600 meters in California) on rocky slopes, 
sandy plains with Larrea, coastal strand, matorral, and chaparral. 

Barbara Hellenthal provided infortnation and photocopies of the relevant ND-G 

pecimens of Gnaphalium bico/or and John Strother provided mformati 
arding material of G. hicolor at UC and JEPS. Review comments of K. G 
nd J. McNeill also are much appreciated. 


lOLETTi, FT. 1 893. Two new Californian plants. Erythea 1:16-17. 




Richard Spellenberg 

Biology Department 
Las Cruces, NM 88003-8001, U.S.A. 

;ico, is proposed for a Boerh 
)rnia to the Big Bend regie 

iende desde las Islas Canales de California hasta la 

<as, y es muy comun en el Desierto Sonorense. Las 

variedades a menudo son simpatricas, y se intergradan un poco. Se presenta una clave para distinguir 

The species of Boerhavia with spicate inflorescences have proven taxonomi- 
cally difficult for the past century, the variation presented by the complex re- 
sulting in very different taxonomic treatments. At one extreme is that of 
Standley (1911) where nine species were recognized. In 1918 Standley submerged 
B. watsoni as a synonym of B. coulteri, but maintained the other taxa he had 
recognized. At the other extreme is Reed's (1969) treatment, in which five of the 
species Standley recognized [B. coulteri (Hook, f.) S. Wats., B. rosei Standi, B. 
spicata Choisy in D.C., B. torreyana (S. Wats.) Standi., B. watsoni Standi] were 
placed as synonyms in one variable species, B. spicata. Most authors of floras 
that include all or part of the Sonoran Desert region have followed Standley 
(1918) in recognizing B. coulteri, and also B. spicata, and B. torreyana if they were 
present in the area of concern (Spellenberg 1993; Martin & Hutchins 1980; 
Kearney & Peebles 1964; Munz & Keck 1968; Wiggins 1964). In contrast, Felger 
(2000) recognized some plants included in B. coulteri, as broadly constructed 
by those authors, as B. spicata var. palmeri S. Wats. It is this taxon that is recog- 
nized as a variety of B. coulteri in this paper. 



;n (Hook. f.)S. Wats, va 


y var. pal-t 

leri S.Wats., from the VI, 

iffers from the var coulter, m US' 


near the I 

Uo Grande, and is very 


atric, and 



avia with: 

spicate mflorescences, 





y var. pain 



•rhavia que difierede la variedadc 





1 del "Big Bend" del Rio Bravo del > 

While studying the complex for an upcoming treatment for the Flora of 
North America North of Mexico, I noted that B. torreyana was conspicuously 
papillate in the sulci between the ribs of the fruit, a character previously un- 
published. None of the other Boerhavia taxa in the complex are at all papillate. 
Watson (1889) cited no specimens when he coined the name Boerhavia spicata 
var. torreyana S. Wats., but specimens from western Texas collected during the 
mid-19th century Boundary Survey, very likely seen by him, have papillate fruits 
(specimens from Torrey Herbarium, NY, have been seen; relevant specimens 
from GH on loan to ENCB have not been seen). The character of papillae on the 
fruit is consistent m populations from western Texas, New Mexico, southeastern 
Arizona at higher elevations, throughout much of Arizona north of the Mogollon 
Rim, and extreme south-central Utah (= B. spicata by Welsh et al. 1987). On very 
similar appearing plants from the Sonoran Desert region, some of which are 
identified as B. torreyana in collections, but which actually represent B. coulteri 
var palmeri (as proposed below), papillae are absent and sulci are much narrower 
and less rugose. That B. torreyana is clearly distinguishable from other spicate 
Boerhavia has helped to clarify the complex. This is particularly so in Arizona 
and Sonora where three or four very similar taxa occur within the region. 

In 1889 Watson transferred Senkenhergia coulteri Hook, f . to Boerhavia, and 
he stated that the collection upon which the name was based was Coulter M25. 
He noted that the perianth was one line in length (= 1/12", ca. 2 mm) and that 
fruits were clavate, 1-1.5 lines long, truncate, and acutely angled. Only one such 
specimen exists in the Hooker Herbarium at Kew. It is labeled as having come 
from "Aha Sonora, Mexico," and is here considered the holotype (photo and 
photocopies at NMC). Its fruits are clearly in irregularly fasciculate clusters in 
the spicate inflorescence, and they are truncate at the apex. On the same page 
of his 1889 publication, Watson stated that his new variety, Boerhavia spicata 
var. palmeri, based on the collection Palmer Ml from Guaymas, Sonora, in 1887, 
had perianths 0.5 line long, and fruits clavate-oblong, a line long, obtuse, ob- 
tusely angled. He also noted the inflorescence to be "open and very slender." 
Palmer Ml is represented by two specimens at GH! (on loan to ENCB, lectotype 
not selected; fragment at DS!). In raising this taxon to specific status, Standley 
(1909) had to adopt a new name (B. watsoni) because the binomial B. palmeri S. 
Wats, was occupied. He noted that Arizona plants were placed there somewhat 
doubtfully, suggesting that geography would place them with B. coulteri. Rela- 
tive to other plants in the complex, the plants he identified as B. watsoni have 
smaller, more widely spaced, less clustered fruits (if at all clustered), and seem 
to have had stems that were decumbent-spreading. 

It IS proposed here that the populations Standley recognized as B. watsoni, 
and that Watson described as B. spicata var palmeri, are part of the B. coulteri 
complex and can be recognized at the varietal level. The following combina- 
tion is proposed: 

.. Wats. var. palmeri (S. Wats.) Spellenb,, comb. nov. 

The differences the two varieties of B. coulteri that Watson, and later Standley, 
noted by their classifications are very apparent in the field in their extreme 
states, but intergradient plants occur Occasionally B. coulteri var coulteri and 
var palmeri are intermixed in populations (e.g., Spellenherg & Zucker 12954 = 
var palmeri, 12955 = var coulteri, AZ, Pima Co., ca. 10 air km SW of Marana, 
Anway Rd. 1 km S of Avra Valley Rd., 16 Aug. 1999; NMC). In the two mixed 
populations I have seen, the taxa differed in habit, the stems of var coulteri de- 
cumbent at the immediate base, then strongly ascending or erect, contrasting 
to the more decumbent-spreading habit of var palmeri, but habit is very likely 
to be influenced by ecological conditions and density of populations. Some 
mono- varietal populations of the var coulteri have decumbent-spreading stems. 
The var coulteri also has larger flowers, a denser inflorescence, and fruits in 
irregular fasciculate clusters, whereas the var palmeri usually has minute flow- 
ers, and well-spaced, smaller fruits, the clusters much less pronounced. Plants 
are most often present in monomorphic populations representing only one 
variety. The flower of B. coulteri var. palmeri is illustrated in Spellenberg (2000, 
fig. 1), but is labelled as B. torreyana; the complex was not understood at the 
writing of that article. 

Inflorescences of all the spicate species (B. coulteri included) I have observed 
are visited by various insects in the morning when flowers are open and pro- 
ducing nectar, the insects moving from inflorescence to inflorescence, and from 
plant to plant. Larger-flowered species are visited more frequently (Spellenberg 
2000). In mixed populations, particularly, inter-taxon pollination is very likely, 
and hybridization is presumed possible. Both varieties of B. coulteri are weedy, 
responding positively to disturbance, and with the advent of increased human 
impact in arid areas of the Southwest, may have come into increased contact. 

Boerhavia coulteri var coulteri is geographically the more restricted of the 
two varieties in the southwestern United States (ranges in Mexico have not been 
determined), occurring from the eastern to the western borders of Arizona, 
south of the Mogollon Rim. A population discovered in south-central New 
Mexico in a well-collected area and may be the result of a recent introduction 
(N.M., Doiia Ana Co., e Las Cruces along University Ave. e of Telshor Drive, 1 km 
e of IH-25, in gravel planting strip between sidewalk and road, 24 Aug 2001, 
Spellenherg and Brouillet 13267, DUKE, MX NMC). The var palmeri occurs 
throughout the Sonoran region in the United States, from southeastern Califor- 
nia to southeastern Arizona. It occurs on Santa Catalina Island in the Channel 
Islands of California (Standley 1909, cited as B. watsoni, Orcutt s.n., 12 Sep. 1889, 
US!)., where it may have been introduced. In Texas the var palmeri is known 
from a few collections on or near the Rio Grande, the eastern most being from 
the eastern side of present day Big Bend National Park (M. 5. Young, s.n., muddy 

bank of Rio Grande, near San Vicente |5 mi SW of Boquillas], 26 Aug 19f 5, TEX 
[2], LL [I]). On one of these Texas collections, all identified originally as B. 
torreyana, is pencilled, "but looks just like B. spicata Choisy Arizona," the writer 
probably alluding to its similarity to B. coulteri var palmeri of Arizona, as here 
recognized. Because this is the first report of this taxon for Texas, the two other 
collections seen are cited here: L.C. Hinckley 1505, Presidio Co, Capote Canyon 
1 mi. below falls, 28 May f94f, US; CM. MuUer8438, Presidio Co., along washes 
of rolling Rio Grande Plain, 4 mi SE of Ruidosa, 28 Jul 1945, LL. In southwest- 
ern New Mexico the var palmeri is occasional and weedy; m the vicinity of Las 
Cruces it is always in disturbed areas. It may be introduced there, but if so, it is 
long-established m the region (e.g., Wooton & Standley3905, NM, Dofia Ana Co., 
mesa W of Organ Mts., 4000 ft, 5 Oct 1907, LL, TEX, US). 

The spicate taxa of Boerhavia in the United States may be distinguished by 
the following key. In this key the range of fruit length-to-width ratios is given 
in the format (length/diam. = 1.3-2.1). 

1. Fruits usually 4-angled(rar 

ely 5-angled), bP 

oadly obovoid (length/diam. = 1,3-2.1); 

branches of inflorescence 

usually densely 

glandular villous, witi 

nout sticky intern- 


1. Fruits 5-angled, broadly to 

narrowly obovo 

id (length/diam. = 1.; 

inflorescence glabrate to 1 


lot glandular, usually 

with sticky intern- 

2. Surface of sulci of mati 

jre fruit papillose 

;, otherwise smooth 


ose.the edges of the ri( 

dges along sulci 

strongly rugose 



2. Surface of sulci not paf 

Dillose, instead sr 

rugose, the edges 

ofthe ridges along sulc 

:i not rugose to s 

trongly rugose 

3. Fruits broadly obovc 

= 1.7-2.1),usuallyov 

eriapping in inflo- 

rescence; sulci and r 

lbs slightly rugo^ 

,e; sulci usually about 

0.5 times as wide 

6. spicata 

3. Fruits narrowly obovoid or obpyramic 

iai (length/diam. = 2.1 


in inflorescence or re 

>mote; sulci and r 

lbs slightly rugose to 


0.2 times as wide as 

1 fruits often ove 


their length, corn- 


ite in groups of 2 

Tim long.truncate. 

round-truncate, b 



4. Inflorescence with 

iO% of their length, 

infrequently 2-3 ii 


ruit 2-2.4 mm long,o 

ccasionally longer, 

usually rounded e 


B. coulteri V, 

ar palmeri 


I acknowledge with gr. 

ititude the he 

Ip of Brian Stanr 

lard (K), who ser 


copies and a photograp 

hic print of tf 



m Hook. 

f. I also thank curators. 

of herbaria w 

ho loaned specimens for this stud 

iy, and to 

Richard Felger for varic 

Dus e-mail "cf 

lats" regarding tf 

ns taxon. 

Felger, R.S. 2000. Flora of the Gran Desierto and Rio Colorado of northwestern Mexico. 

Univ. Arizona Press, Tucson. 
Kearney,T.H. and R.H.Peebles. 1964. Arizona flora, 2nd. ed., with supplement. Univ. Calif Press, 

Martin, W.C and C.R.Hutchins. 1980. A flora of New Mexico, vol. 1. J. Cranner, Vaduz. 
MuNZ, PA. and D.D. Keck. 1 968. A California flora, with supplement. Univ. Calif Press, Berke- 

Reed, CF. 1 969. Nyctaginaceae. In: C.L. Lundell, ed. Flora of Texas, vol. 2(1 ).Texas Research 

Foundation, Renner. Vol. 2(1 ), pp. 1 5 1 -220. 
Spellenberg,R. 1993. Nyctaginaceae. In: J. Hickman, ed.TheJepson manual: higher plants of 

California. Univ. Calif Press, Berkeley. Pp. 768-774. 
SpELLENBERG,R.2000.Blooming"behavior"in five species of fioer/iowo (Nyctaginaceae). Sida 


Contr. U.S. Natl. Herb. 12:303-389. 

1 3:377-430. 
Standley, PC. 1 91 8. Allioniaceae. North America flora 21 :1 71 -254. New York Botanical Gar- 

Watson,S. 1889. VI. Contributions to American botany. 1. Upon a collection of plants made 
by Dr. E. Palmer, in 1 887, about Guaymas, Mexico, at Muleje and Los Angeles Bay in 
Lower California, and on the island of San Petro Martin in the Gulf of California. Proc. 
Amer. Acad. Arts 24:36-87. 

Welsh, S.L, N.D. Atwood, S. Goodrich, and LC Higgins (eds.). 1 987. A Utah flora. Great Basin 
Naturalist Mem. 9. Brigham Young University, Provo. 

Wiggins, I.L 1 964. Flora of the Sonoran Desert. In: F. Shreve and I. L.Wiggins, Vegetation and 
flora of the Sonoran Desert. Stanford Univ. Press, Stanford. Pt. 2, pp. 1 80-1 669. 


lARD L DUBLE 2001 Turfgrasses: Their Management and Use in the Southern 

Zone, Second Edition. (ISBN 1-58544-161-9, pbk.) Texas A&M University 
Press. John H. Lmdsey Building, Lewis Street, 4354 TAMU, College Station, 
TX 77843-4354. (Orders:, 979^458-3982, 800-826- 
8911, 979-847-8752 fax). $27.95, 336 pp, 51 color photos, 15b/w photos, 68 
Une drawings, 7" X 9". 
)ugh monotypic plantings of an exotic grass species might evoke fear or loathing in those con- 
d with biodiversity, few would dispute the impact turfgrasses have on our economy or our quality 
:. Gulf courses, municipal parks, residential landscapes and interstate roadways are all depen- 
n some form on turf grass culture and management. Turfgrasses capitalizes on their importance 

:ting root growth Chapter four is devoted e 
rm-season and cool-season For each group 
tion on the origin and distribution of spec 
Itural varieties propagation, managemen 

> and tieat turf disea; 
1 playing fields 

ntati ve, the line drawings of turfgrass species and turf weeds are of poor quality. The drawing of a 
aminate Buffalograss (p. 72), for instance, could easily be mistaken for Bouteloua and Kentucky 
uegrass (p.84) looks more like Johnsongrass. It is likely that the target audience would not be fa- 

t Worth, TX, 76102^4060, U.S.A.,amy_trauth@yahoc 



Plant Resources Center 
AustinJX78712,U.S.A. ' 



In their m 
genus Me, 
proposed i 
an evaluat 

onograph of the fam 
itzeiia as having 36 1 
is new. The purpose ( 
;ion of their biologic 

laxa, two of these, IV 
3f the present paper 




ypify each and provide 

Urban and Gilg in their protologue of this species cited five specimens gath- 
ered by five different collectors, three of these from Texas, two from northern 
Mexico. All of the specimens cited by these two authors were said to be from 
European herbaria, namely "Berol., Boisser-Barbey Canby Lips, Paris,J.D. Smith, 
Vindobon." I have examined duplicates of all of the sheets concerned and select 
the following from among these as adequate lectotypes: U.S.A. Texas: Uvalde 
Co.: Jun 1885, Reverchon 1650 (lectotype: MO!; isolectotype: US!). Darlington 
(1934) also listed this collection as belonging to his concept of M. lindheimeri. 
Urban and Gilg listed the locality of Reverchon 1650 as "in planitiebus arenosus 
montis Upper Guadelupe" but the locality on the lectotype is that given here. 
Ideally, a lectotype should be selected from among the specimens exam- 
ined by its original author or authors, but the European collections cited by 
Urban and Gilg, who worked out of the herbarium at Berlin (B), were unavail- 
able to me. Because of this, I find it appropriate (as permissible under Art 9.10 of 
the current Code), to designate lectotypes from American syntypes. I have also 
applied this same reasoning in the typification of the following taxon. 

Mentzelia texana Urban & Gilg, Monogr. Loas. 52. 1900. Without specific locality, 
May-Oct, 1849, Wright 210 (Lectotype, here designated: GH; ISOLECTOTYPES: GH, 

In the protologue of this taxon, which I consider to be a synonym of M. 
lindheimeri, Urban and Gilg noted that, so far as known, it was confined to 

Texas, citing three specimens: Drummond 91 (from sandy soils between Victo- 
ria and Gonzales, Texas; Trecul 1220 (dry sites between Turkey Creek and the 
San Pedro iDevils Riverl; and Wright 210 (m part, mixed with M. oligosperma 
Nutt.). They also noted that material of their new species, presumably those 
cited, were based upon specimens " in herb. Berol., Boisser-Barbey, CandoUe, 
Delessert, Kew., Paris., Vindobon." 

Darlington (1934) retained the species, citing two of the above mentioned 
type elements {Drummond 202 and Wright 202). She also cited three additional 
specimens from the state of Pueblo, Mexico. I take the latter to be 
misidentifications of the more southern M. hispida Willd., or some close rela- 
tive having relatively short petals. Correll and Johnston (1970), following 
Darlington, retained M. texana in their treatment of Mentzelia for the flora of 

In a recent study of Mentzelia for Texas 1 have examined duplicates of two 
of the three collections cited by Urban and Gilg in their protologue ( Drummond 
91 and Wright 210) and have selected Wright 210 as an appropriate element for 
lectotypification, largely because duplicate specimens of this collection are 
currently housed at GH! and US! 

The type locality of M. texana is moot. According to label data on Wright 
210 (US!) the specimen concerned was collected "May-Oct, 1849." But other than 
this time span, no other information relating to its collection is given. Accord- 
ing tojohnston (1940), on June 28, 1849, Charles Wright collected an unnamed 
species of Mentzelia (Wright's field number 541) on the hills of Turkey Creek 
in western Uvalde County. Since Gray often renumbered and/or combined 
Wright's collections (cf. Johnston 1940), it is likely that Wright 210 is the num^ 
ber assigned by Gray to Wright's field collection S'^l; at least I encountered no 
Wright collection so numbered in the present study 

The positioning of M. texana in synonymy with the simultaneously pub- 
lished M. lindheimeri is based upon the study of numerous specimens from 
ten or more herbaria, and field work over a number of years in the areas con- 
cerned. Urban and Gilg, in their key to species, distinguished M. texana from 
M.lindhei men by its shorter petals (7-9 mm vs. 12-14 mm) and sessile or shortly 
pedicellate fruits (vs always clearly pedicellate). I find these two quantitative 
characters to be quite variable within the M. lindheimeri complex and have 
little hesitancy in reducing M. texana to synonymy Finally, it should be noted 
that Darlington (1934) listed the lectotype of M. texana (Wright 210) among his 
"Specimens examined:" as a sheet of M. lindheimeri, which agrees with the 
present authors disposition. 

Foundation, Renner. 

ARLINGTON, J, 1934. A mo. 

Leop.-Carol. Germanics 

Book Receivrd/Notk 

iNE Gail Fautin, DouglasJ. Futuyma, and H. Bradley Shaffer (eds). 2001. Annual 
Review of Ecology and Systematics. Volume 32, 2000 (ISBN 8243 1432- 
8, hbk; ISSN 0066-4162). Annual Reviews Inc., 4139 El Cammo Way, P.O. 
Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders:, 
800-523-8635, 650-493-4400, 650-424-0910 fax). $140.00 hbk., 614 pp. 

he year 2001 has 19 articles followed b 


Robert W. Freckmann Michel G. Leiong 

Department of Biology Department of Biological Sciences 

urn and Dichanthelium have a 1 

ions, made for a forthcoming treatment of these two genera, are as follows: D. sect. Angustijolia; D. 
ect. Ensijolia; D. sect. Lanceariai D. sect. Lanuginosa] D. sect. Oligosantha; D. sect. Sphaerocarpa; D. 
•xiculare subsp. angusliJoUum, suhsp. fusif or me, subsp. neuranthum; D. acuminatum subsp. 
olumbianum, suhsp.Jasciculatum, subsp. implicatum, subsp. leucothrix, subsp. lindheimeri, subsp. 
ongiligulatum, subsp. sericeum, subsp. spretum, subsp. thermale; D. chamaelonche and subsp. breve; 
> commiUatum subsp. ashei, subsp. equilaterale, subsp. joori; D. dichotomum subsp. lucidum, subsp. 
nattamuskeetense, subsp. microcarpon, subsp. nitidum, subsp. roanokense, subsp. yadkinense; D. 
nsi/olium subsp. cur(i/olium;D.o!igosanthes subsp. scribnerianum; D. ovale subsp. praecocius, subsp. 
iseudopubescens, subsp. vidosissimum; D. portoricense subsp. patulum; D. strigosum subsp. 
labrescens, subsp. leucobiepharis; D.tenue;Fflmarum subsp. (3marulum;Ranceps subsp. rhizomatum; 

ihpes; R hirticaule subsp. sonorum, subsp. stramineum; P. philaddphicum subsp. gattingeri, subsp. 
ithophilum; and P rigidulum subsp. abscissum, subsp. combsii, subsp. dongatum, subsp. pubescens. 

t. Sphaerocarpa; D. sect. Angusti/oiia; D. sect. Ensi/olia; D. sect. Lancearia; D. sect. Lanuginosa; D. 
'yiigosantha; D. sect. Sphaerocarpa; D. aciculare subsp. angusti/olium, subsp. /usi/orme, subsp. 
mthum; D. acuminatum subsp. columhianum, suhsp.Jasciculatum, subsp. impiicaCum, subsp. 
thrix, subsp. lindheimeri, subsp. iongiligu latum, subsp. sericeum, subsp. spretum, subsp. 
(ale; D. chamaelonche y subsp. breve; D. commutatum subsp. ashei, subsp. equilaterale, subsp. 
D. dichotomum subsp. iucidum, subsp. mattamuskeetense, subsp. microcarpon, subsp. nitidum, 
). roanofeense, subsp. yadhinense; D. ensijolium subsp. curtijolium; D. oligosanthes subsp. 
lerianum; D. ovale subsp. praecocius, subsp. pseudopuhescens, subsp. vtllosisstmum; D. 

urn subsp. amarulum; P. anceps subsp. rhizomatum; P. capillare subsp. hilmanii; P. 
tomiflorum subsp. hartowense, subsp. puritanorum; P. hallii suhsp. Jili pes; P. hirticaule subsp. 
um,subsp. stramineum; P pbiladelpbicum subsp. gaftingeri, subsp. (ithophilum; yP rigidulum 
. abscissum, subsp. combsii, subsp. elongatum, subsp. pubescens. 

In preparing the treatment of Panicum and Dichanthelium for Volume 25 of 
The Flora of North America (Barkworth et al. [eds.], in prep.), we fmd it neces- 
sary to describe eight new infrageneric taxa and make fifty-two new combma- 
tions. Editorial policy for this volume requires that infraspecific taxa be treated 
as subspecies rather than as varieties. 

The genus Panicum, as recognized by Linnaeus (1753), included 20 species m 
two groups: Spicata and Paniculata. Most of these species have been tranferred 
subsequently to other genera, including Digitaria, Echinochloa, Oplismenus, 
Paspalum, Pennisetum, and Setaria. The segregation of Dichanthelium from 
Panicum by Gould (1974-) on characters of anatomy and photosynthetic path- 
ways is supported by a recent molecular phylogeny of the Panicoideae (Giussani 

Bentham and Hooker (f883) treated Panicum as a relatively broad genus 
and published several infrageneric taxa. Nash (1903) treated Panicum m a nar- 
rower sense. He named and described nine unranked groups of Panicum in 
Small.Flora of the Southeastern United States. Even though no infrageneric rank 
was indicated, these names were validly and effectively published, and eight of 
these names can serve as basionyms m Panicum or Dichanthelium. Hitchcock 
revised the Gramineae for Robinson and Fernald (1908) m Gray's Manual of 
Botany, ed. 7. He divided Panicum m this treatment into two sections, the rank 
indicated by the double-s sign, which Robinson and Fernald define on p. 32 as 
indicating a section. He divided section Eupanicum into six unranked names, 
presumably subsections, and added 14 names, presumably series, within the 
"subsection" Dichotoma. Although these 20 infrasectional names are unranked, 
these names were also validly published and can also serve as basionyms. 

Hitchcock and Chase (1910) added 13 additional unranked groups in Pani- 
cum and effectively published descriptions of these groups. Several subsequent 
taxonomists (e.g. Fernald 1936, 1938, 1947; Hsu 1965) have treated their group 
names as sectional or subsectional names. Hitchcock and Chase, however, ex- 
plicitly stated (page 17) that they did not consider these groups to be formal 
taxonomic entities. They used the same group names m their later publications 
(Hitchcock 1913, 1915, 1935, 1951; Hitchcock and Chase 1915) but never suggested 
that they had changed their minds with respect to the taxonomic status of the 
groups. Article 34.1 of the International Code of Botanical Nomenclature 
(Greuter et al. 2000) states that a name is not validly published when it is not 
accepted by the author in the original publication. It seems likely that, had 
Hitchcock changed his mind regarding these group names, he or Chase would 
have included that information in one of their later publications for Hitchcock 
was a very active participant in the development of the nomenclatural code 
(Hitchcock 1919; Lawrence 1951) and hence must have been aware of the need 

to make a clear statement of such a change of mind. Therefore, if the Hitchcock 
and Chase groups added in the 1910 paper are to be recognized as formal taxa, 
they must subsequently be validly published. We have indicated that we are 
deriving our section names from these group names in the 1910 paper by using 
"Hitchcock & Chase ex Freckmann & Lelong" for those groups which retain 
the Hitchcock and Chase group name and circumscription. 

Pilger (1931) named new sections in Panicum, indicating these as "nov. sect." 
meeting all requirements that apply to a 1931 publication. He also apparently 
published validly, but inadvertently, as sections two of the Hitchcock and Chase 
groups by reference to the Hitchcock and Chase names in the 1910 and 1915 
papers, even though he did not indicate these as ''nov. secC 

Hsu (1965) is often cited as validating Hitchcock and Chase names or as 
publishing new combinations based on these names, but no types were cited, 
as required after 1 January 1958 by Article 37.1 of the Code. Gould (1974) raised 
subgenus Dichanthelium to a genus, making new combinations for 17 species 
and three varieties, but he did not treat subgenera or sections. Brown m Brown 
and Smith (1975) elevated two sections of Panicum to subgenus status in 
Dichanthelium. Zuloaga (1987) returned to a broader concept of Panicum and 
published four new sections, each with a Latin description and citation of a 
type. He also changed the status of three of Hitchcock's group names to sec- 
tions, citing types and validly publishing these names. 

jlae pyramidalis, plu; 

Plants perennial, with thick rhizomes. Culms erect, robust, hard, branching at 
the middle nodes. Ligule a ciliate membrane. Panicle pyramidal, more or less 
lax, many-flowered, with spikelets on long pedicels. Spikelets ovoid, glabrous; 
first glume about 1/2 as long as the spikelet, 3- or 5-nerved; second glume and 
lower lemma 7-nerved; lower floret staminate; upper floret smooth, shiny, x = 9. 
I Hitchc. & Chase ex Freckmann & Lelong, sect. nov. 

,. Spiculae ar 

Plants perennial, sei 


or sprawling, often 

sterile. Ligule £ 

row; branches few, spiciform, erect 


larrowly ovoid 1 

Lowell lore: sramm 

ate, upper lemiT 

only near the base 

xtensive rhizomes or stolons. Culms erect 
hate or lacerate membrane. Panicle nar- 
ith subsessile toshort-pedicelled, secund 
anceolate, laterally compressed, glabrous 
hin llexible, whitish, rolled over the palea 

Plants perennial, from a thick rhizome or caudex. Culms 50-150 cm; autumnal 
culms with branches arising from the middle and upper nodes, rebranching 
and often forming dense axillary fascicles, the panicles enclosed m the sheaths. 
Basal leaves forming a winter rosette. Sheath usually swollen, mottled, papil- 
lose-hispid or pubescent and viscid, constricted at summit; basal blades large, 
ovate-lanceolate; cauline blades 6-14. Spikelets ellipsoid to ovoid, pointed, 2.2- 
3.6 mm long, with prominent veins. 
Dichanthelium sect Linearifolia Freckmann & belong sect no\ Tmii i'duunm 

Plants perennial, tufted. Culms erect to spreading or drooping, the lower inter- 
nodes very short, the upper 2-4 internodes often much elongated. Autumnal 
culms branching from the nodes near the base, producing condensed panicles 
within about 5 cm of the ground or sterile shoots. Basal blades narrow, erect or 
ascending, resembling the lower cauline ones; cauline blades usually erect, stiff, 
the upper ones 1-5 mm wide and 15-60 times as long. Spikelets ovoid, ellipsoid, 
or obovoid, 2.0-4.3 mm long, glabrous or pubescent. 

Hitchcock and Chase (1910) used the group name "Depauperata" for most 
of the species in this section. However, because Pilger (1931) published section 
Depauperata for certain Old World species of Panicum and because 
Dichanthelium is often included in Pa nicum, we prefer to avoid creating a hom- 
onym if these genera are combined. 

Plants perennial, from a knotted rhizome, a caudex, or rarely an elongated rhizome. 
Main culms 20-110 cm, ascending to erect, sparsely rebranching, not producing 
dense axillary fascicles; additional culms (sometimes arising in autumn) smaller. 
Basal leaves forming a winter rosette. Sheaths glabrous or pubescent, not viscid; 
cauline blades 3-7. Panicle at least partly exserted. Lower floret often staminate. 

Nash (1903) published the infrageneric name Latifolia within Panicum, 
indicating that this name was based on Panicum latifolium L. However, it is 
evident from his description ("much branched shrubs...outer [spikelet] scales 
woolly-tipped") that this name was based on a misapplication of Panicum 
latifolium, referring instead to Panicum (.=Lasiacis) divaricatum L. According 
to Art. 7.4 of the Code a name is typified by the type of the basionym, even 
though it may have been apphed to a taxon now considered not to include that 
type. However, m the case of an unranked infrageneric name, a modern author 
may publish a new name at a definite rank. We prefer to avoid confusion over 
the application of Latifolia by basing this section name on a different type. 

Plants perennial, tufted. Culms 20-60 cm, weakly ascending, glabrous; autum- 
nal culms branching from the base. Blades flat to stiffly involute, ascending, 
crowded at the base, the uppermost ones much reduced. Basal leaves not form- 
ing a distinct winter rosette. Panicle long-exserted, few-flowered. Spikelets nar- 
rowly ellipsoid, 2.4-2.9 mm long, glabrous. 

: Freckmann & belong, sect. 

.el obpyr 

m puberulun 

from , 

I hard, cor 

erect or spreading; autumnal culms freely divaricate-branching before the pri- 
mary panicle matures, ultimately decumbent. Basal leaves not forming a win- 
ter rosette. Ligule a ciliate membrane. Blades papillose-ciliate. Spikelets obo- 
void to obpyriform, 3.2-4.4 mm long, papillose-pubescent, attenuate at the base. 
Upper floret pointed to attenuate, puberulent at the apex. 

Dichanthelium sect. Strigosa Freckmann & Lelong, sect. nov. Typus: Panicum 
strigosum Muhl. in Elliott. 

Plants perennial, densely tufted. Main culms slender, the lower internodes short, 
the upper 3-5 internodes elongated. Autumnal culms forming a dense cushion. 
Blades soft, pale green or yellowish, usually with ciliate margins. Basal blades 
large, scarcely separable from the lower cauline blades. Spikelets broadly ellip- 
soid to obovoid, 1.1-2.3 mm long, glabrous to pubescent. 

Bentham and Hooker (1883) created the section Laxiflora to include cer- 
tain Old World species of their subgenus Eupanicum, but not including P 
laxijlorum Lam. To avoid creating a later homonym if Dichanthelium is treated 
as part of Panicum, we are basing the section name on Panicum strigosum Muhl. 
in Elliott, rather than P laxijlorum. 

Dichanthelium sect. Angustifolia (Hitchc. m B.L. Rob. & Fernald) Freckmann 

& Lelong, Stat. nov. Basionym: Pamcum (unranked) Angustifolia Hitchc. in B.L. 

Rob. & Fernald, Gray's Man. Bot. Ed. 7:106. 1908. TYPE: Panicum angustijolium Elliott. 
Dichanthelium sect. Ensifolia ( B.L. Rob. & Fernald) Freckmann & 

Lelong, Stat. nov. Basionym: Pamcum (unranked) Ensifolia Hitchc. in B.L. Rob. 

& Fernald, Gray's Man. Bot, ed. 7:113. 1908. TYPE: Panicum ensifolium Baldwin ex 

Dichanthelium sect. Lancearia (Hitchc. m B.L. Rob. & Fernald) Freckmann & 

Lelong, Stat. nov. Basionym: Panicum (unranked) Lancearia Hitchc. in B.L. Rob. 

& Fernald, Gray's Man. Bot, ed. 7:114. 1908. TYPE: Panicum lancearium Trin. 
Dichanthelium sect. Lanuginosa (Hitchc. m B.L.Rob. & Fernald) Freckmann & 

Lelong, Stat, nov Basionym: Panicum (unranked) Lanuginosa Hitchc. in B.L. Rob. 

& Fernald, Gray's Man. Bot, ed. 7:109. 1908. Type: Panicum lanuginosum EUiott. 
Dichanthelium sect. Oligosantha (Hitchc. m B.L.Rob. & Fernald) Freckmann & 

Lelong, Stat, nov Basionym: Pamcum (um-anked) OUgosanthia Hitchc. m B.L. Rob. 

& Fernald, Gray's Man. Bot, ed. 7:114. 1908. Type: Panicum oligosanthes Schult. 
Dichanthelium sect. Sphaerocarpa (Hitchc. in B.L.Rob. & Fernald) Freckmann 

& Lelong, Stat, nov Basionym: Pamcum (unranked) Sphaerocarpa Hitchc. in B.L. 

Rob. & Fernald, Gray's Man. Bot, ed. 7:113. 1908. Type: Panicum sphaerocarpon Elliott. 

Dichanthelium ; 

aciculare subsp. angustifolium (Elliott) Freckmann & Lelong, 



:129. 1816. Panicumnitidumvar. 

angustifolium (Elliott) A. Gray, N. Amer 

Gram. 2:112 

. 1835. Chaseaangustijolia (Ellic 

)tt) Nieuwl, Amer. Midi. Naturalist 2:64. 


nthelium angustifolium (Elliott 

;) Gould, Brittonia 26:59. 1974. Panicum 


ar. angustifolium (Elliott) Wipff & S.D.Jones, Phytologia 77:457. 1994. 

Type: U.S.A. 

South Carolina: May, 5. Elliot 

t s.n. (HOLOTYPE: CHARL, fragment USD. 

Dichanthelium aciculare subsp fusiforme (H 

itchc.) Freckmann & Lelong, comb. 

& Stat, no- 

V. BASIONYM: Panicum fusiform 

e Hitchc, Contr U.S. Natl. Herb. 12:222. 

1909. Dich. 

inthelium fusiforme (Hitchc.) 

Harvill, Castanea 42:177. 1977. Type: 

CUBA: wesi 

tern Cuba, 1863, C Wright 345^ 

I (holotype: GOET, fragment US!). 


ihum var. ramosum Griseb., Cat. Pi. 

Cuh. 232. 1866. Dichanthelium aciculare var. 

ramosum (Gi 

-iseb.) Davidse, Novon 2:104. 1992. 


aciculare subsp. neuranthum (Griseb.) Freckmann & Lelong, 

comb. & s 

;tat. nov. Basionym: Panicum 

neuranthum Gnseb., Cat. Pi. Cub 232. 

1866. Type: 

CUBA: eastern Cuba, 1860, C Wright 3453 (holOTYPE: US 2209346!). 


■umSwallen.Proc. Biol. Soc. Wash, f 

i3:93. 1942. 

Dichanthelium acuminatum subsp. columbianum (Scribn.) Freckmann & Lelong, 
comb. & Stat, nov Basionym: Panicum columbianum Scribn., U.S.D.A. Div. Agrost. 
Bull. 7:78, f . 6. 1897. Dichanthelium columbianum (Scribn.) Freckmann, Phytologia 
39:270. 1978. Panicum acuminatum var columbianum (Scribn.) Lelong, Brittonia 
36:270. 1984. TYPE: U.S.A. DISTRICT OE COLUMBIA: 14 Jul 1894 F.L. Scribner s.n. (ho- 

Dichanthelium acuminatum subsp. fasciculatum (Torr.) Freckmann & Lelong, 
comb & Stat., nov Basionym: Panicum dichotomum var. fasciculatum Torn, El. 
North. & Mid. U. S. L145. 1824. Panicum huachucae var. fasciculatum (Torr.) FT 
Hubb., Rhodora 14:171. 1912. Panicum lindheimeri var.fasciculatum (Torr.) Fernald, 
Rhodora 23:228. 1921. Panicum lanuginosum var.fasciculatum (Torr.) Fernald, 
Rhodora 36:77. 1934. Dichanthelium lanuginosum var.fasciculatum (Torr.) 
Spellenb. Madrono 23445. 1975. Dichanthelium acuminatum var.fasciculatum 
(Torr.) Freckmann, Phytologia 48:108. 1981. Panicum acuminatum var.fasciculatum 
(Torr) Lelong, Brittonia 36:269. 1984. TYPE: U.S.A. NEW JERSEY: sandy fields,J. Torrey 

Dichanthelium acuminatum subsp. implicatum (Scribn. ex Nash) Freckmann 
& Lelong, comb. & stat. nov Basionym: Panicum implicatum Scribn. ex Nash 
in Britton & A. Br., Illust. Fl. 3:498, f. 267a. 1898. Panicum unciphyllum var. 
implicatum (Scribn. ex Nash) Scribn. & Merr., Rhodora 3:123. 1901. Panicum 
lindheimeri var implicatum (Scribn. ex Nash) Fernald, Rhodora 23:228. 1921. Pani- 
cum lanuginosum var implicatum (Scribn. ex Nash) Fernald, Rhodora 36:77. 1934. 
Dichanthelium acuminatum var implicatum (Scribn. ex Nash) Gould & C.A. Clark, 
Ann. Missouri Bot. Card. 65:1126. 1979 [imprint date 1978]. Panicum acuminatum 
var impl icatum (Scribn. ex Nash) Beetle, Phytologia 48:192. 1981. TYPE: U.S.A. MAINE: 
low marshy ground. Cape Elizabeth, 26 Jul 1895, F.L. Scribner s.n. (holotype: US 

Dichanthelium acuminatum subsp. leucothrix (Nash) Freckmann & Lelong, 
comb. & Stat, nov Basionym: Panicum leucothrix Nash, Bull. Torrey Bot. Club 
24:41. 1897. Dichanthelium Zeucothrix (Nash) Freckmann, Phytologia 48:101. 1981. 

Dichanthelium acuminatum subsp. lindheimeri (Nash) Freckmann & Lelong, 
comb. & Stat. nov. Basionym: Panicum lindheimeri Nash, Bull. Torrey Bot. Club 
24:196. 1897. Panicum lanuginosum var. lindheimeri (Nash) Fernald, Rhodora 
36:77. 1934. Dichanthelium lindheimeri (Nash) Gould, Brittonia 26:60. 1974. 
Dichanthelium lanuginosum var. lindheimeri (Nash) Freckmann, Phytologia 
39:270. 1978. Dichanthelium acuminatum var. lindheimeri (Nash) Gould & C.A. 
Clark, Ann. Missouri Bot. Card. 65:1127. 1979 [imprint date 1978]. Panicum 
acuminatum var lindheimeri (Nash) Beetle, Phytologia 48:193. 1981. Panicum 
lindheimeri var tjypicum Fernald, Rhodora 23:227. 1921. Type: U.S.A. TEXAS: springs, 
banks of the Guadeloupe, near New Braunfels, 1846, F.J. Lindheimer 565 (holo- 
TYPE: NY!). 

acuminatum subsp. longiligulatum (Nash) Freckmann & Lelong, 
comb. & Stat. nov. Basionym: Panicum longiligulatum Nash, Bull. Torrey Bot. Club 
26:574. 1899. Dichanthdium acuminatum van !ongiiigu/atum(Nash)Gould 61 C.A. 
Clark, Ann. Missouri Bot. Card. 65:1127-1128. 1979 [imprint date 1978]. 
Dichanthelium longiligulatum (Nash) Freckmann, Phytologia 48:102. 1981. Pani- 
cum acuminatum var longiligulatum (Nash) Lelong, Brittonia 36:270. 1984. Type: 
U.S.A. Florida: Apalachicola, 1892, G.R. Vasey (holotype: NY!). 

ubsp. sericeum (SchmoU) Freckmann & Lelong, 
ONYM: Panicum ferventicola var sericeum SchmoU, 
Madrono 5:92. 1939. Dichanthelium lanuginosum var sericeum (SchmoU) Spellenb, 
Madrono 23:150. 1975. Dichanthelium acuminatum var sericeum (SchmoU) 
Freckmann, Phytologia 48:107. 1981. Type: U.S.A. WYOMING: Mammoth Hot Springs, 
Yellowstone National Park, in wet ground about hot springs, A. Nelson 6037 (ho- 
lotype: RM 20108). 

Dichanthelium acuminatum subsp. spretum (Scbult.) Freckmann & Lelong, 
comb. & Stat. nov. Basionym: Panicum spretum Schult., Mant. 2:248.1824. 
Dichanthelium spretum (Schult.) Freckmann, Phytologia 48:102. 198L TYPE: U.S.A. 
New England, moist ground, Muhlenberg's PanicumNo. 37 (holotype: PH). 

Dichanthelium acuminatum subsp. thermale (Bol) Freckmann & Lelong, comb. 
& Stat. nov. Basionym: Panicum thermale Bol, Proc. Calif. Acad. Sci. 2:18L 1862. 
Dichanthelium lanuginosum var thermale (Bol.) Spellenb., Madrono 2.3:151. 1975. 
Dichanthelium acuminatum var thermale (Bol.) Freckmann, Phytologia 48:107. 
1981. TYPE: U.S.A. California. Sonoma Co.: northern part of Sonoma County on 
hot rocks and in hot water flowmg from Geyser Springs and Geyser Mountains, 
H.N. Bo!ander 3941 (HOLOTYPE: GH). 

Dichanthelium chamaelonche (Trm.) Freckmann & Lelong, comb. nov. Basionym: 
Enslins.n. (holotypE: LE; fragment and photograph US!). 

Dichanthelium chamaelonche subsp. breve (Hitchc. & Chase) Freckmann & 
Lelong, comb. & stat. nov. Basionym: Panicum breve Hitchc. & Chase, Contr 
U.S. Natl. Herb. 15:271. 1910. Dichanthelium dichotomum var hreve (Hitchc. & 
Chase) Gould & C.A. Clark, Ann. Missouri Bot. Card. 65:1120. 1979 ]imprmt date 
1978]. Panicum chamaelonche var breve (Hitchc. & Chase) Lelong, Brittonia 36:267. 

1984. Dichanthelium ensifolium var. breve (Hitchc. & Chase) B.F. Hansen & 
Wunderlin, Ann. Missouri Bot. Gard. 75:1646. 1989 [imprint date 1988]. Pamcum 
ensifolium var hreve (Hitchc. & Chase) Wipff & S.D.Jones, Phytologia 77:460. 1994. 
TYPE: U.S.A. FLORIDA: Jensen, low pme woods between scrub hills, among pal- 
metto, 5 Apr 1906, AS- Hitchcock 734 (holotype: US 558435!). 
Dichanthelium commutatum subsp. ashei (G. Pearson ex Ashe) Freckmann & 
Lelong, comb. & stat. nov. Basionym: Pamcum ashei G. Pearson ex Ashe, J. Elisha 
Mitchell Sci. Soc. 15:35 (1898). Pamcum commutatum var. ashei (G. Pearson ex Ashe) 
Fernald, Rhodora 36:83 (1934). Type: U.S.A. New York: Ithaca.Jul 1898, W.W.Ashe 

Dichanthelium commutatum subsp. equilaterale (Scribn.) Freckmann & Lelong, 
comb. & Stat. nov. Basionym: Panicum equilaterale Scribn., U.S.D.A. Div. Agrost. 
Bull. 11:42. 1898. TYPE: U.S.A. FLORIDA: Lake Co.: scrubby hammock lands, Eustis, 
16-25 Aug 1894, G.V. Nash 1674 (lectotype: US 743929! [designated by Hitchcock 
& Chase, Contr U.S. Natl. Herb. 15:310. 1910.]) 

Dichanthelium commutatum subsp. joori (Vasey) Freckmann & Lelong, comb. 
& Stat. nov. Basionym: Pamcumpon Vasey, U.S.D.A, Div, Agrost, Bull, 8:31.1889. 
Type: U.S.A. Louisiana: East Baton Rouge Par, near Baton Rouge, creek bank, in 
dense tufts, 1 Oct. 1885, J.FJoor 39 (holotype: US!). Panicum divergens Kunth in 
H.B.K. has been used by Zuloaga et aL (1993) to replace P commutatum. However, 
Davidse 6a: Zuloaga (2000) in proposing rejection of P divergens note that the prov- 
enance of the type collection is uncertain and that the name had never been ap- 
plied previously to P commutatum. 

:hotomum subsp. lucidum (Aslie) Freckmann & Lelong, comb. 
& stat. nov Basionym: Panicum lucidum Ashe, J. Elisha Mitchell Sci. Soc. 15:47. 
1898. Panicum dichotomum var lucidum (Ashe) Lelong, Brittonia 36:265. 1984. 
Dichanthelium lucidum (Ashe) LeBlond, Sida 19(4):831. 2001. Type: U.S.A. North 
Carolina: deep shady swamps bordering Lake Mattamuskeet, 10 Jun-6Jul 1898, 
WW. Ashe s.n. (lectotype: fragment US! [designated by Hitchcock & Chase, Contr 
U.S. Natl. Herb. 15:198. 1910.]). 

Lelong, comb. & stat. nov Basion 

Mitchell Sci. Soc. 15:45. 1898. Panicum dichotomum var, mattamusneetense (,Asne; 
Lelong, Brittonia 36:265, 1984. Dichanthelium mattamuskeetense (Ashe) LeBlond, 
Sida 19:832. 2001. Type: U.S.A. North Carolina: around Lake Mattamuskeet, 10 
Jun-6 Jul 1898, WW Ashe and G. Pearson s.n. (lectotype: US 2808955! [designated 
by Hitchcock & Chase, Contr U.S. Natl. Herb. 15:186. 1910.]). 

Dichanthelium dichotomum subsp microcarpon (Muhl. ex Elliott) Freckmann 
& Lelong, comb. & stat. nov Basionym: Pamcum microcarpon Muhl. ex Elliott, 
Sketch Bot. S. Carolina 1:127. 1816. Type: U.S.A. GEORGIA, A.A. Baldwin s.n. (lecto- 
type: CHARE [designated by Hitchc. & Chase, Contr. U.S. Natl. Herb. 15:181.1910.]). 

Dichanthelium dichotomum subsp. nitidum (Lam.) Freckmann & Lelong, comb. 
& stat. nov Basionym: Panicum nitidum Lam., Tabl. Encycl. 1:172. 1791. Panicum 
dichotomum var nitidum (Lam.) Alph, Wood, Class-book bot., ed. s.n. [hi 786. 1861. 
Dichanthelium dichotomum var. nitidum (Lam.) LeBlond, Sida 19(4):829. 200L 
Type: U.S.A. "Carolina": eastern Carolina,;. Prasers.n. (holotype: P, fragment US!). 

Dichanthelium dichotomum subsp. roanokense (Ashe) Freckmann & Lelong, 
comb. & Stat, nov. Basionym: Pamcum roanokense AsheJ, Elisha Mitchell Sci. 
Soc. 15:44. 1898. Panicum dichotomum var. roanokense (Ashe) Lelong, Brittonia 
36:265. 1984. Dichanthelium dichotomum var. roanokense (Ashe) LeBlond, Sida 
19(4):831. 2001. TYPE: U.S.A. North Carolina. Dare Co.: Manteo, Roanoke Island, 
dry soil, lOJun 1898, W.W.Ashe s.n. (lectotype: US![designated by Hitchc. & Chase, 
Contr U.S. Natl. Herb. 15:196. 1910]). 

Dichanthelium dichotomum subsp. yadkinense (Ashe) Freckmann & Lelong, 
comb. & Stat. nov. Basionym: Panicum yadkinense Ashe, J. Elisha Mitchell Sci. 
Soc. 16:85. 1900. Pamcum dichotomum var yadkinense (Ashe) Lelong, Brittonia 
36:266. 1984. Dichanthelium yadkinense (Ashe) Mohlenbr., Erigenia 6:27. 1985. 
Type: U.S.A. North Carolina. Raleigh, May 1895, WW Ashe s.n. (lectotype: US 
2808938! [designated by Hitchc. & Chase. Contr U.S. Natl. Herb. 15:195. 1910]). 

Dichanthelium ensifolium subsp. curtifolium (Nash) Freckmann & Lelong, 
comb. & Stat. nov. Basionym: Panicum curtifolium Nash, Bull. Torrey Bot. Club 
26:569. 1899. Panicum ensifolium var curti/olium (Nash) Lelong, Brittonia 36:266. 
1984. Type: U.S.A. Mississippl Ocean Springs, 2 May 1898, S.M. Tracy 4598 (holo- 

Dichanthelium oligosanthes subsp. scribnerianum (Nash) Freckmann & Lelong, 
comb. & Stat, nov Basionym: Pamcum scribnerianum Nash, Bull. Torrey Bot, 
Club 22:421. 1895. Panicum oligosanthes var. scribnerianum (Nash) Fernald, 
Rhodora 36:80. 1934. DichantheZiumo!igosan£hes var. scnbnenanum(Nash)Gould, 
Brittonia 26:60. 1974. TYPE: U.S.A. Pennsylvania. Wysox, Jul 1836,J. Carey s.n. (lec- 
totype: GH [designated by Hitchc. & Chase, Contr. U.S. Natl. Herb. 15:283. 1910]). 

nee ElUott. (1816). 
Dichanthelium ovale; 

comb. & Stat, i 

8:206. 1906. Pamcum lanuginosum var. praecocius (Hitchc. & Chase) Dore in 
McNeill & Dore, Naturahste Canad. 103:562. 1977. Dichanthelium villosissimum 
var. praecocius (Hitchc. & Chase) Freckmann, Phytologia 39:270. 1978. Type: U.S.A. 
Illinois. Stark Co.: dry bank, near Wady Petra, 30 Jun 1900, V.H. Chase 649 (holo- 

Dichanthelium ovale subsp. pseudopubescens (Nash) Freckmann & Lelong, 
comb. & Stat, nov Basionym: Pamcum pseudopubescens Nash, Bull. Torrey 
Bot. Club 26:577. 1899. Pan icum villosissimum var. pseudopubescens (Nash) 
Fernald,Rhodora 36:79. 1934. Panicum ovale var. pseudopubescens (Nash) 
Lelong, Brittonia 36:271, 1984. Type: U.S.A. Alabama: Lee Co.: Auburn, 7 
May 1898, F.S. Earle and C.F. Ba ker 1537 (holotype: NY!), 

Dichanthelium ovale subsp, villosissimum (Nash) Freckmann & Lelong, comb, 
& Stat, nov Basionym: Panicum villosissimum Nash, Bull, Torrey Bot. Club 23:149, 

1974, Dichanthelium villosissimum (Nash) Freckmann, Phytologia 32:270. 1978. 
Type: U.S.A. Georgia. Ocmulgee River swamp, below Macon, 18-24 May 1895,J.K. 
SmaH s.n. (HOLOTYPE: NYD. 

absp. patulum (Scribn. & Merr.) Freckmann & 

U.S.D.A. Dept. Agrost. Circ. 27:9. 1900. Panicum patulum (Scribn. & Merr.) Hitchc, 
Rhodora 8:209. 1906. Panicum lanceanum var. patulum (Scribn. & Merr.) Fernald, 
Rhodora 36:80. 1934. Dichanthelium sahulorum var patulum (Scribn. & Merr.) 
Gould & CA. Clark, Ann Missouri Bot. Gard. 65:1113. 1979 [imprint date 1978]. Type: 
U.S.A. Florida. Manatee Co.: Braidentown [Bradenton], 3 Sep 1898, R. Comhs 1296 
(holotype: us 2808982!). 

Dichanthelium strigosum subsp. glabrescens (Griseb.) Freckmann & Lelong, 
comb. & Stat. nov. BASIONYM: Panicum dichotomum var. glabrescens Gris^h., Fl. 
Brit. W.Indies 553. 1864. Dichanthelium leucohlepharis var. glabrescens (Griseb.) 
Gould & CA. Clark, Ann. Missouri Bot. Gard. 65:1100 (1979) [imprint date 1978). 
Dichanthelium strigosum var glabrescens (Griseb.) Freckmann, Brittonia 33:457. 
1981. Panicum strigosum var. glabrescens (Gristh.) Lelong, Brittonia 36:264. 1984. 
Type: JAMAICA: W Purdie s.n. (holotype: K). 

Dichanthelium strigosum subsp. leucohlepharis (Trin.) Freckmann & Lelong, 
comb. & Stat. nov. Basionym: Panicum leucohlepharis Trin., Clav Agrost. 234. 
1822. Dichanthelium leucohlepharis (Trin.) Gould & CA. Clark, Ann. Missouri Bot. 
Gard. 65:1099. 1979 [imprint date 1978]. Dichanthelium strigosum var. 
leucohlepharis (Trin.) Freckmann, Brittonia 33:457, 1981. Panicum strigosum var 
leucob/epharis (Trin.) Lelong, Brittonia 36:264. 1984. Type: U.S.A. North Carolina, 
South Carolina, Georgia, Lousiana: Enslins.n. (lectotype: LE, fragment US! [des- 
ignated by Hitchc. & Chase, Contr. U.S. Natl. Herb. 15:162. 1910]). 

Dichanthelium tenue (Muhl.) Freckmann & Lelong, comb. nov. Basionym: Pani- 
cum tenue MuW., Descr Gram. 118 (1817). Dichanthelium dichotomum var tenue 
(Muhl.) Gould & CA. Clark, Ann. Missouri Bot. Gard. 65:1119. 1979 [imprint date 
1978]. Type: U.S.A.: MuhlenbergHerb. 192 (lectotype: PH-M [designated by Hitchc. 
& Chase, Contr U.S. Natl. Herb. 15:259. 1910]). 

Panicum amarum subsp. amarulum (Hitchc. & Chase) Freckmann & Lelong, 
comb. 61 Stat, nov Basionym: Panicum amarulum Hitchc. & Chase, Contr U.S. 
Natl. Herb. 15:96. 1910. Panicum amarum var amarulum (Hitchc. & Chase) P.G. 
Palmer, Brittonia 27:149. 1975. Type: U.S.A. Virginia. Virginia Beach, 24 Sep 1900, 
lA. Williams 3090 (holotype: US 592748!). 

Panicum anceps subsp. rhizomatum (Hitchc. & Chase) Freckmann & Lelong, 
comb. & Stat, nov Basionym: Panicum rhizomatum Hitchc. & Chase, Contr. U.S. 
Natl. Herb. 15:109, f . 104. 1910. Panicum anceps var rhizomatum (Hitchc. & Chase) 
Fernald, Rhodora 36:73. 1934. Type: U.S.A. South Carolina: Orangeburg, 18 Aug 
1905, AS. Hitchcock 450 (holotype: US 592752!). 

Panicum capillare subsp. hillmanii (Chase) Freckmann & Lelong, comb. & stat. 
nov Basionym: Panicum hillmanii [as "hinmani"]Chase,J. Wash. Acad. Sci. 14:345. 
1934. Type: U.S.A. Texas: Amarillo, plain, 11 Aug 1918, A.S. Hitchcock 16206 (holo- 
type: US 1037542!). 

Panicum dichotomiflorum subsp. bartowense (Scribn. & Merr.) Freckmann & 
Lelong, comb. & stat. nov Basionym: Panicum bartowense Scribn. & Merr, 
U.S.D.A. Dept. Agrost. Circ. 35:3. 1901. Panicum dichotomiflorum var. bartowense 
(Scribn. & Merr) Fernald. Rhodora 38:387. 1938. TYPE: U.S.A. FLORIDA. Polk Co.: in 
v^et reclaimed swamps at Bartow, 29 Sep 1898, R. Combs 1220 (holotype: US 80538!). 

Panicum dichotomiflorum subsp. puritanorum (Svenson) Freckmann & Lelong, 

Svenson, Rhodora 22:154, f. 1-5. 1920. TYPE: U.S.A. MAINE: dry sandy and gravelly 

beach, Half-way Pond, Barnstable, 13 Sep 1919, ML. Fernald 306 (holOTYPE: GH). 
Panicum hallii subsp. filipes (Scribn.) Freckmann & belong, comb. & stat. nov. 

Basionym: Panicum/i/ipes Scribn. in A. Ileller,Contr Herb. Franklin Marshal I Coll. 

1:13. 1895. Panicum ha/Hi var./ihpes (Scribn.) RR. Waller, SouthW. Naturalist 19:105. 

1974. Type: U.S.A. Texas: growing in rich shaded ground in the upper part of the 

arroyo at Corpus Christi, alt. 30 feet, 31 May 1894, A.A. Heller 1809 (holotype: US 

Panicum hirticaule subsp. sonorum (Vasey) Freckmann & belong, comb. & stat. 

nov. Basionym: Panicum sonorum Vasey in Beal, Grasses N. Amer 2:130. 1896. Type: 

MEXICO. Sonora: Lerdo. at the head of the Gulf of California, 1889, E. Palmer 947 

(holotype: us 2903025!). 

cum hirticaule var. miliaceum (Vasey) Beetle, Phytologia 47:383. 1981. 
Panicum hirticaule subsp. stramineum (Hitchc. & Chase) Freckmann & belong, 

comb. & stat. nov. Basionym: Panicum stramineum Hitchc. & Chase, Contr U.S. 

Natl. Herb. 15:67, f. 50. 1910. Panicum capillare var. stramineum (Hitchc. & Chase) 

Gould, Madrono 10:94. 1949. Panicum hirticaule var. stramineum (Hitchc. 61 

Chase) Beetle, Phytologia 47:383. 1981. Type: MEXICO. Sonora: Guaymas,Jun-Nov 

1887, E Palmer 206 (holotype: US 592753!). 
Panicum philadelphicum subsp. gattingeri (Nash) Freckmann & belong, comb. 

& stat. nov Basionym: Panicum capillare var. gattingeri Nash in Britt. & Brown, 

U.S.A. Tennessee: cedar glades near Nashville, Sep, A. Gattinger s.n. (lectotype: 
Iragment US! [designated by Hitchc. & Chase, Contr U.S. Natl. Herb.]). 

Panicum philadelphicum subsp. lithophilum (Swallen) Freckmann & belong, 
comb. & stat. nov Basionym: Panicum lithophilum Swalltn, Proc. Biol. Soc. Wash. 
54:43. 1941. Type: U.S.A. Georgia: on rocky slope, Stone Mountain, 23 Aug 1905, 

1940. Type: U.S.A. Florida: Sebring, m dry sand at tourist camp near lake, 3 Oct 
1925, P Weatherwa. 

i (Scribn. & C.R. Ball) Freckmann & belong, 
Panicum comhsii Scribn. & C.R. Ball, U.S.D.A. Div 
Agrost. Bull. 24:42, f . 16. 1901 [imprint date 1900]. Panicum longifolium var comhsii 
(Scnbn.&CR. Ball) Fernald, Rhodora 36:69(1934).Panicumngidulum var. combsii 
(Scribn. & C.R. Ball) Lelong, Brittonia 36:263. 1984. Type: U.S.A. FLORIDA. Wash- 
ington Co.: damp fertile flat woods [low pine barrens] at Chipley, 20 Aug 1898, R. 
Combs 583 (holotype: US 80589!). 
cum rigidulum subsp. elongatum (Pursh) Freckmann & belong, comb. & stat. 
nov BASIONYM: Panicum elongatum Pursh, Fl. Amer Sept. 1:69. 1813 [imprint date 
1814], non Salisb. (1796). Panicum agrostoides var elongatum (Pursh) Scribn., Tennes- 
see Agric. Exp. Sta. Bull. 7:42. 1894. Panicum rigidulum var elongatum (Pursh) Lelong, 

Bnttonia 36:263, 1984. TYPE: U.S.A. DELAWARE: F.T. Pursh s.n. (lectotype: K, photo US! 
[designated by Hitchc. & Chase, Contr. U.S. Natl. Herb. 15:104. 1910]). 
Pamcum slipUatum Nash m Scribn., U.S.D.A. Div, Agrost. Bull. 17 [rev. ed.l:56, f . 352. 1901. 
Panicum rigidulum subsp. pubescens (Vasey) Freckmann & Lelong, comb. & 
Stat. nov. Basionym: Pamcum anceps var. puhescens Vasey U.S.D.A. Div. Bot. Bull. 
8:37. 1889. Panicum longifolium var. puhescens (Vasey) Fernald, Rhodora 36:69. 1934. 
Panicum rigidulum var. puhescens (Vasey) Lelong, Brittonia 36:263. 1984. TYPE: 
U.S.A. Alabama: Mobile Co.: Mobile, C.T. Mohrs.n. (lectotype: US 80530! [desig- 
nated by Hitchc. & Chase, Contr U.S. Natl. Herb. 15:105. 1910]). 
We are very grateful to Mary E. Bar]<wortli for her continuous support and en- 
couragement of our efforts to produce new treatments of Panicum and 
Dichanthelium and for her assistance m preparing this manuscript. We are also 
very grateful to K.N. Ghandhi for his extensive and meticulous search of the 
literature for mfrageneric names and for his help in interpreting the applica- 
tion of the Code to some difficult nomenclatural problems. 

Bentham,G and J.D.Hooker. 18 

83. Genera plantarum.Vol. 3 



1 258. L Reeve & 

Co., London. 

Brown, W.V. and B.N. Smith, 1975. The genus Dichantheliur 

n (Gram 



Davidse, G. and F.O, Zuloaga. 

2000. Proposal to reject 

the nan 

le Par 

Ileum divergens 



Fernald, M.LI 936. Plants frorr 

1 the outer coastal plain of V 


•a 38:376-404. 

Fernald, M.L 1938. Contributi 

ons from the Gray Herbarii 

im of Ha 


University - No. 

CXXII.VIIl. New species, var 

ieties and transfers. RhodorE 

i 40:331- 


Fernald, M.L 1947. Additions 

to and subtractions from th 

le flora c 


nia. Rhodora 49: 

Gould, F.W.I 974. Nomenclatu 

ral changes in Poaceae, Brit 

tonia 26: 


Greuter, W.L, J. McNeill, FR.BARf 






0. Intern, 



cal nomenclature. Koeltz Scientific Books, Konigstein, ( 



, F.O. Zuloaga, and E.AKellog. 




ofthe grass subfamily Pan 

icoideae (Poaceae) shows rr 



of C4 photosyn- 

thesis. Amer.LBoL 88(1 1):1 


Gould, F.W. 1 974. Nomenclatu 

iral changes in the Poaceae. Brittoni, 

a 26:5? 


Hitchcock, A.S.1913.Mexican< 

grasses in the United States 


1 Herb, 

arium, Contr. U.S. 

Natl. Herb. 17:181-389. 
Hitchcock, A.S.I 91 5. Pan/cum 

LlnJ.H.Barnhart,ed.N Ame 




Hitchcock, A.S. 1 951 . Manual of grasses of the United States. Ed. 2, Rev. A. Chase. U.S.D.A. 

Misc. Publ. 200. 
HiTCHCOCK,A.S.and A.Chase. 1910.The North American species of Pan/cum. Contr.U.S.NatL 

Hitchcock, A.S. and A. Chase. 1 91 5. Tropical North American species of Panicum. Contr. U.S. 

Natl. Herb. 17:459-539. 
Hsu,C.C. 1965. The classification of Panicum (Gramineae) and its allies, with special refer- 

Lawrence, G.H. M.I 951. Taxonomy of vascular plants. Macmillan, New York. 

PiLGER, R. 1931. Bemerkungen zu Panicum und verwandten Gattungen. Notizblatt der 

Botanischen Gartens und Museums zu Berlin-Dahlen 1 1 (104):237-247. 
ZuLOAGA, P.O. 1 987. Systematics of New Word species of Panicum (Poaceae; Paniceae). In: 

T.R. Soderstrom, K.W. Hilu. C.S. Campbell, and M.E. Barkworth, eds. Grass systematics 

and evolution. Smithsonian Institution Press, Washington, D.C. Pp. 287-306 
ZuLOAGA, P.O., R.P Ellis, and 0. Morrone. 1 993. A revision of Panicum subg. Diclianthelium 

secX.Dichanthelium (Poaceae: Panicoideae: Paniceae) in Mesoamerica, the West Indies, 


rsiana B.L. Turner, sp. nov., is described and illustrated from n 

mg ledges. The sj 
md features of th 


e El Salto, Durango, Mexico. 
;a. 2720 m) y se dice que es 
: la especie esta emparentada 

Routine identification of Mexican Asteraceae has revealed the follow novelty: 
Senecio sandersiana B.L. Turner, sp. nov (Fig. l). Type: MEXICO. DURANGO: Mpio. El 

Salto, "rim of plateau W of La Ciudad, 3.1 km NW of Hwy 40 ... off Neveros rd. 

upper Arroyo Tascanto watershed," 2720 m, 19 Apr 1999, A.C. Sanders etal. 22680 


Perennial subscapose herhs to 60 cm high. Leaves glabrous or nearly so, 15-25 cm 
long, 3.0-8.5 cm wide, forming a persistent basal rosette; petioles of well-devel- 
oped leaves alate, mostly 10-15 cm long, 2-4 mm wide; blades broadly ovate to 
cordate, the margins decidedly serrulate. Heads 1-3 at the apices of elongate neariy 
naked scapes (bearing 2-4 much-reduced, bract-like leaves). Involucres calycu- 
late, the calyculae grading into the involucral bracts; inner bracts ca. 21 in a single 
series, ca. 8 mm long, 1.5 mm wide, sparsely glandular-pubescent along the mar- 
gins, otherwise glabrous. Receptacle plane to concave, epaleate, glabrous, ca. 4 
mm across. Ray florets 11-13, sterile; corollas yellow, glabrous; tubes ca. 6 mm long; 
hgules 4-6 nervate, 6-8 mm long, 2-3 mm wide. Diskjlorets 40-60, fertile; corol- 
las yellow, glabrous, ca. 7 mm long, the tube ca. as long as the cylindrical throat, 
the lobes 5, deltoid, ca. 0.5 mm long. Style branches hnear, truncate, minutely 
hispidulous apically. Achenes (immature) ca. 2.5 mm long, glabrous; pappus of 
numerous white bristles somewhat longer than the disk corollas. 

This taxon is closely related to the widespread species, 5. tolucannus of the 

\ - i ^ C ^ D 


Sec. Lugentes of Senecio. It might also be compared with 5. wootonii Greene of 
the southwestern U.S.A. and closely adjacent northwestern Mexico. Senecio 
sandersiana differs from both in having fewer heads to a capitulescence and 
markedly cordate leaf blades. The distribution of these several taxa in Mexico 
IS shown m Fig. 2. 

Etymology— The species is named for its primary collector, A.C. Sanders, 
Curator at UCR, an intrepid field worker When informed of its possible novel 

marked via ema 
r that 22680 was 

live object. This 

about....! have 5 more sheets. It was very common where I found it: I had t 
through beds of it on the shaded slopes to find other things." 

I am grateful to my wife, Gayle Turner, for the Latin diagnosis. Susan Wi 
provided the illustration. 


Pat Halliday. 2001. The Illustrated Rhododendron. (ISBN 0-88192-510-1, hbk.). 
Timber Press, Inc., 133 S.W. Second Avenue, Suite 450, Portland, OR 97204- 
3527, U.S.A. (Orders:, 800-327-5680, 503-227-2878, 
503-227-3070 fax). $69.95, 274 pp., 120 color illustrations, 4 line drawings, 
8 1/4" X 11 3/4". 


pretty pictui 


It IS certs 

anly a great deal more 


for more thai 

a 40; 

years, has 

■ culled from its archive 


;tion of the genus 




'ere prepared 


a herbar: 

ium material. lUustra- 



result is i 

1 comprehensive study 




vation of 

: rhododendrons. 

,e orij 

^mal iUustra 


; were ta 

ken from the pages of 

CY KoBER. 2000. With Paintbrush & Shovel. (ISBN 0-8139-1969-X, hbk.). The 
University Press of Virginia. Charlottesville and London. (Orders: Price not given, 280 pp., 8 photographs, 1 map, 
222 plates. 8 1/2" x 10". 

ersburg, Virginia, the author has woven threads of many colors and textures into a fascinating 
and sociological study There are 
ts and habitats were already dis- 


Zachary S.Rogers 
illustrated. Both sclerophyllous specie 


Se describe y se ilustra Weinmannia bradfordiana Z. Rogers y Weinmam 
dos especies esclerofilas son conocidas linicamente de las localidades d 
en la region de la Cordillera del Condor en la provincia de Morona Sai 

The family Cunoniaceae is composed of 26 genera and about 300 species 
(Bradford & Barnes 2001). The australly-distributed genus Weinmannia L ac- 
counts for about 150 species m five sections (Bradford 1998; Bradford 2002). All 
American species are in section Weinmannia, with the greatest species rich- 
ness found in northwestern South America (i.e. Venezuela, Colombia, Ecuador 
and Peru) (Bernardi 1961; Harhng 1999). 

Weinmannia bradfordiana and Weinmannia condorensis were collected 
by the late Alwyn H. Gentry (1945-1993) on his last major expedition, and were 
filed in a "family indeterminate" folder for a number of years, despite the good 
quality of the specimens in flower and fruit. Later, while sorting Gentry's col- 
lections, Ron Liesner (MO) came across the specimens but did not initially con- 
sider the specimens as belonging to the family Cunoniaceae because of their 
uncharacteristically short inflorescences, but he did notice a few small fruits 
that looked remarkably like the septicidal capsules of Weinmannia. Ron then 
showed the material to Jason Bradford, who was studying Neotropical 

annotated the specimens as new species of Weinmannia (Ron 

I Bradford, pers. comm.). 

as well as another recently described species of Weinmannia 

from the Condor (Rogers 2002), have adapted extremely sclerophyllous habits 
due to the very wet and windy weather conditions and the very thin, nutrient- 
poor sandstone substrate. Many other woody genera on the summits of the 
Cordillera del Condor have similar sclerophyllous adaptations, and many of 
these species will also turn out to be new and locally endemic to the range. 

from their types and are probably closely related, due to their similar leaf mor- 
phology and ecology and because they were collected from sites about 10 km 
apart, but significant differences exist between the two including: leaf size, leaf- 
let shape, number of flowers per inflorescence, and trichome density and dis- 
tribution on the stems, leaves, stipules and sepals. Based upon experience with 
other recognized species of Weinmannia and character variation within and 
among species, the differences observed from these specimens suggest they rep- 
resent distinct species. This analysis is based entirely on a subjective study of 
the type collections and available herbarium specimens, so more collections 
will be needed to elucidate the patterns of variation m these species. At this 
time, the type localities are unable to be revisited because the sites have been 
covered by deadly land mines since the border dispute between Ecuador and 
Peru m January 1995, which was about 18 months after the types were origi- 
nally collected. The mine field will not be cleared for at least several more years, 
but when the situation is resolved, further exploration will be needed and may 
expose intermediates that support a notion expressed by one reviewer (David 
Neill), who felt that both collections could represent a single species. 

These new species can be easily distinguished from other Ecuadorian spe- 
cies by their small trifoliolate (rarely unifoliolate) compound leaves on short 
petioles that can appear to be simple and sessile to the naked eye, and by their 
extremely short inflorescences (i.e. pseudoracemes) due to inconspicuous pe- 
duncles. The highly reduced inflorescences are very uncharacteristic of the 
genus because almost all Weinmannia have elongated inflorescence axes mea- 

Color images of the type collections can be found on the W3 TROPICOS 
database at: <>. 

Weinmannia bradfordiana Z. Rogers, sp. nov (Fig. 1). TvPl-: rxXLADOR. Moko\.\ 
Santiago: Canton Gualaquiza: Campamento Acliupalla, Cordillera del C ondor, 
15 km E of Gualaquiza, tepui-like bromeliad sward witli scattered, smal I trees, 03" 
27'S, 78" 22'W, 2090 m, 21 Jul 1993 (fl., fr.), Gentry 80312 IHOi o\ WE: QCN1:-092179; 
ISOTYPES: AAU, GB-186976, MO-5544000, NY, USj. 

Shrubs to 1 m; young stems covered with dense lanate or tomentose trichomes, 
the trichomes to 1.3 mm long, simple, unicellular, matted, stiff, white, nodes 
densely hirsute. Leaves to 5.3 mm long, opposite, compound, trifoliolate (rarely 
unifoliolate), densely congested; leaflets 2.0-4.9 x 1.4-3.3 mm, broadly obovate 
to nearly orbicular, broadly boat-shaped to nearly planar, upturned at apex, all 
leaflets similar m size and shape, coriaceous, thick, membranous and thin along 
the margins, shiny, drying dark brown on the upper surface and light brown 
on lower surface, midvein raised on upper and lower surface, thicker near the 
base, secondary veins 3-5 pairs per side, slightly raised to nearly inconspicuous 
on the upper surface, prominently raised on the lower surface, blade surfaces 
glabrous, usually lanate or tomentose along the upper one-third of the apical 
margins (usually more than 20 trichomes per apex), the trichomes are similar 
to those found on the young stems, 0.4-1.0 mm long, upper surface scrobicu- 
late, lower surface scrobiculate and rugose, base cuneate to slightly oblique, 
margin entire, apex obtuse to slightly apiculate; petioles 0.1-0.4 mm long, re- 
duced to inconspicuous or indiscernible protuberances, obscured by the dense 
pubescence at the nodes, dark red or black, densely hirsute or tomentose; peti- 
olules 0.1-0.4 mm long, swollen, dark red or black, glabrous, rugulose; stipules 
2.2-3.6 mm x 1.8-2.7 mm, mtcrpetiolar, paired, free, broadly ovate to suborbicu- 
lar, membranous to subcoriaceous, dark red or brown, outer surface densely 
lanate or tomentose, becoming more dense towards the apex and along the 
margin, tricfiomes to 1.1 mm long, similar to those found on the young stems, 
inner surface glabrous, base truncate, margin entire, apex rounded to obtuse, 
caducous. Pseudoracemes paired, unbranched, densely compacted, axillary on 
young growth at the terminus of the stem; peduncles to 0.9 mm long, reduced 
to inconspicuous protuberances, densely lanate or hirsute; fascicles (2-)4-8- 
flowered, densely compacted and congested; pedicels to 0.9 mm long in flower, 
to 1.9 mm long in fruit, sparsely lanate or hirsute; bracteoles not seen. Flowers, 
actinomorphic, bisexual, diplostemonous, recorded as white m color on the label; 
sepals 4, 1.0-1.2 x 0.5-0.9 mm m flower, to 1.4 mm long in fruit, ovate, subcoria- 
ceous, adaxial surface glabrous, abaxial surface densely lanate to sparsely to- 
mentose, denser near the apex and along the margin; petals 4, 1.0-1.2 x 0.8-1.0 
mm, broadly elliptical to broadly obovate, membranous, white, midvein con- 
spicuous, slightly raised and dark, glabrous, base truncate, margin entire and 
ciliated along the upper portion, apex rounded; stamens 8; filaments 0.8-2.0 
mm long, 0.4-0.6 mm wide at base, flattened, slender at apex, glabrous; 
anthers 0.3-0.4 x 0.4-0.5 mm, orbicular, introrse, dorsifixed, longitudinally 
dehiscent; nectary disk annular with 8 concrescent lobes, surrounding the ovary 
persistent in fruit; ovary 0.7-0.9 x 0.3-0.5 mm, superior, bicarpellate, syncar- 
pous, red or brown, glabrous; styles 2, 0.2-0.4 mm long in flower, 0.6-0.9 mm 
long in fruit, divergent near the apex of the carpels, persistent in fruit; stigma 
simple, capitate. Fruits 1.9-2.4 x 1.4-1.8 mm (length measurement not includ- 

ing the persistent styles), septicidal capsules, orbicular, occasionally widely 
ovate, dark red or brown, costate longitudinally glabrous; seeds not seen. 

Distri hution and Hahitat.—Weinmannia hradfordiana is known only from 
the type locality collected near one of the summits of the Cordillera del Condor 
at about 2100 m elevation, and located about 15 km SE of the town of Gualaquiza. 
The Condor mountain range is covered by "dwarf-forest" vegetation consisting 
of many species of shrubs and small trees that have evidently adapted 
sclerophyllous habits due to thm, nutrient-poor sandstone substrate, and ex- 
tremely wet and windy weather on the ridge. 

Etymology— The epithet was chosen in honor of Dr Jason C. Bradford, who 
has spent a number of years in the field collecting Cunoniaceae, and who has 
made valuable contributions to our knowledge of the family through his con- 
tinuing research efforts. 

lorensis Z. Rogers, sp. nov (Fig. 2), Type: ECUADOR. Morona 
Santiago: Canton Gualaquiza: Crest of Cordillera del Condor, ridge top 15 km ENE 
of Gualaquiza, high montane forest and bromeliad sward. 03°22'S, 7870'W, 2500 
m, 26 Jul 1993 (fl, fr), Gentry 80465 (holotype: QCNE-092303; isotypes: AAU, 

Trees to 3 m; young stems covered with dense lanate or hirsute trichomes, the 
trichomes to 1.1 mm long, simple, unicellular, matted, stiff, white, nodes densely 
hirsute. Leaves to 7.6 mm long, opposite, compound, trifoliolate (rarely unifoli- 
olate), congested; leaflets 3.6-6.8 x 1.9-4.5 mm, obovate to elliptical, broadly 
boat-shaped to nearly planar, upturned at apex, all leaflets similar in size and 
shape, coriaceous, thick, membranous and thin along the margins, drying dark 
brown, shiny midvein prominently raised on upper and lower surface, thicker 
near the base, darker than blade, secondary veins 2-5 pairs per side, raised to 
nearly inconspicuous on the upper surface, more prominently raised on the 
lower surface, blade surfaces glabrous, infrequently having a few sparse tri- 
chomes at the apical margins (usually 5 or fewer trichomes per apex when pu- 
bescent), the trichomes are similar to those found on the young stems, 0.1-0.4 
mm long, upper and lower surface scrobiculate and rugose, base cuneate to 
slightly oblique, margin entire, apex obtuse to slightly apiculate; petioles 0.7- 
1.2 mm long, often inconspicuous, obscured by the dense pubescence at the 
nodes, dark red or black, sparsely hirsute or tomentose; petiolules 0.3-0.6 mm 
long, swollen, dark red or black, glabrous, rugulose; stipules 2.2-3.5 mm x 2.0- 
3.0 mm, mterpetiolar, paired, free, ovate to suborbicular, membranous to sub- 

the inflorescence. All illusti 

)r from the type collec 

entire, apex rounded to obtuse, caducous P^cudo} 
densely compacted, axillary on young growth, de 
quent nodes at the term m us of the stem: peduncle 

. un branched, 
ng, reduced to 

lar to those found on the young stems; fascicles 1- or 2-flowered, com- 
md congested; pedicels to 1.2 mm in flower, sparsely hirsute; bracteoles 
[. Flowers actinomorphic, bisexual, diplostemonous, recorded as white 

in color on the label; sepals 4, 1.6-2.4 x 0.8-2.0 mm, broadly ovate, subcona- 
ceous, adaxial surface glabrous, abaxial surface lanate, denser near the apex and 
along the midvein; petals 4, 1.6-2.0 x 1.2-2.2 mm, broadly elliptical to broadly 
obovate, membranous, white, midvein conspicuous, slightly raised and dark, 
glabrous, base truncate, margin entire and ciliated, apex rounded; stamens 8; 
filaments 0.8-2.5 mm long, 0.3-0.5 mm wide at base, flattened, slender at apex, 
glabrous; anthers 0.3-0.4 x 0.2-0.4 mm, orbicular, introrse, dorsifixed, longitu- 
dinally dehiscent; nectary disks annular with 8 concrescent lobes, surround- 
ing the ovary, persistent in fruit; ovary to 0.9 mm long, to 0.6 mm wide, supe- 
rior, bicarpellate, syncarpous, red or brown, glabrous; styles 2, to 2.1 mm long, 
divergent near the apex of the carpels, persistent in fruit; stigma simple, capi- 
tate. Fruits (immature) 1.8-2.5 mm long (measurement not including the per- 
sistent styles), septicidal capsules, ovate, dark red or brown, glabrous; seeds not 

Distribution and Hahitat.-Weinmannia condorensis is known only from 
the type locality collected on a ridge-top of the Cordillera del Condor moun- 
tain range at about 2500 m elevation, and located about 15 km ENE of the town 
of Gualaquiza. The site, as in the case of W hradfordiana, is covered by "dwarf- 
forest" vegetation consisting of many species of shrubs and small trees with 
similar sclerophyllous adaptations. 

Etymology -The specific epithet refers to locality of the type collection. 

Aj/inities.— According to the key for the Cunoniaceae in the Flora of Ecua- 
dor (Harlmg 1999), W hradfordiana and W. condorensis would fit between the 
couplet W cochensis Hieron. and W mariquitae Szyszyl. because they have small 
compound leaves measuring less than 5 cm in length, but W. mariquitae is very 
different morphologically because it has much larger leaves with 4-8 leaflet 
pairs per leaf. On the other hand, W. hradfordiana and W. condorensis can be 
easily distinguished from the most morphologically similar species from Ec- 
uador, W. cochensis, because they have smaller, trifoliolate (rarely unifoliolate) 
leaves with smaller, entire-margined leaflets, and inconspicuous peduncles. No 
other currently described species of Weinmannia in Ecuador has such highly 
reduced compound leaves and peduncles. 

Weinmannia hradfordiana can be vegetatively distinguished from W 
condorensis by its smaller leaves, its smaller, more broadly obovate to subor- 
bicular leaflets, by the presence of 20 or more trichomes found along the apical 
margins of most leaflets, and by its densely pubescent stipules. The inflores- 
cences of W hradfordiana are more congested by more numerous flowers, and 
have slightly longer, more globular peduncles than W. condorensis. In contrast, 
most leaflets of W. condorensis are typically larger, narrowly obovate to elhpti- 
cal and glabrous. Weinmannia condorensis also has inflorescences that are re- 
duced to only 1 or 2 flowers, and stipules that are mostly glabrous except for a 
dense apical pubescence. 

Another novel and distinctive character separating the two species is that 
W. condorensis has a rare inflorescence architecture, where pairs of racemes 
develop at two successive nodes near the terminal end of the main stem. The 
inflorescence of section Weinmannia (i.e. all Neotropical species) is limited to 
a pair of racemes developing from axillary buds at the most distal node, and is 
a distinctive character of the section. The development of racemes at two sub- 
sequent nodes has not been found in other Neotropical members of the genus, 
but it has been observed in a small group of species from Madagascar and the 
Comores (Bradford, pers. comm.). Patterns in inflorescence architecture are rela- 
tively stable in Weinmannia, and in many cases, can be more helpful than plas- 
tic vegetative characters, which often prove to be similar among many closely 
related species (Bradford 1998). The interesting inflorescence architecture of 
W condorensis will definitely warrant further investigation once more collec- 
tions are made available. 


1 long, margins entire; petic 
Leaflets broadly obovate, margins usually ci 

2. Leaflets narrowly obovate to elliptical, r 
centrated along the apices and along tl 

The author thanks Jason Bradford (MO-DAV), Richard Barnes (HO) and David 
Neill (MO-QCNE) for their helpful review suggestions on the manuscript. The 
author would also like to thank Ron Liesner (MO) and Jon Ricketson (MO) for 
their comments, Roy Gereau (MO) for his preparation of the Latin diagnoses, 
Barney Lipscomb (BRIT) for waving the publication costs of this manuscript, 
and Uno Eliasson (GB) for expeditiously providing loan material. 


Bradford, J.C. 1 998. A cladistic analysis of species groups in Weinmannia (Cunoniaceae) 
based on morphology and inflorescence architecture. Ann. Missouri Bot. Gard. 85: 

id R.W. Barnes. 2001. Phylogeneti 
Bing chloroplast DNA sequences i 
Cunoniaceae.ln:G. Marling and L. 

Howard Garrett with Odena Brannam. 2001. Herbs for Texas. (ISBN 0-292-78713- 
3, hbk.). University of Texas Press, RO, Box 7819, Austin, TX 78713-7819, U.S.A. 
(Orders: 800-252-3206, 800-687-6046 fax). $60.00, 280pp. 8 photographs, 
1 map, 222 plates, 8 1/2" x U", 

1 msighf 
. photo - 

c\ LF^sSl^PAN 2001 Picturing Tropical Nature. (ISBN 0-8014-3881-0, hbk) 
Cornell Univ Press, 512 East State St , Ithaca, NY 14850, USA (Orders 
607-277-2211, 607-277-6292 fax) $35 00, 243 pp , numerous b&w photos, 
6" X 9 1/4" 


Jill D.AIford and Loran C Anderson 

Department of Biological Science 


Macranthera is monotypic; the single species M.Jlammea is endemic to the 
southeastern coastal plain of the United States and is commonly referred to as 
"flame flower" or "hummingbird flower." It is listed as endangered by the state 
of Florida (Coile 2000). The genus was formerly placed in the Scrophulariaceae, 
but recent studies have shown that family to be a heterogenous group (Olmstead 
et al. 2001); the hemiparasitic taxa are better placed in the Orobanchaceae. 

Here, an updated species description and range map are provided. More 
detail on the life history and pollination biology is documented by Alford (2000). 

The many names that have been proposed for this species indicate the diffi- 
culty that early botanists had in understanding the taxon and its generic place- 
ment (Table 1). William Bartram (1791) first described M.Jlammea as Gerardia 
flammea during his travels through the southeastern states during 1773-1777. 
In 1834, after examining an unlabeled specimen in the herbarium of the 
Academy of Natural Sciences, Philadelphia (PH), Nuttall published the genus 

Russelia flammea (Bartr.) Raf. 
Toxopus gymnanthes Raf. 


Conradia as a dedication to the late Solomon W. Conrad, a Professor of Botany 
at the University of Pennsylvania. Nuttall proposed the name Conradia 
fuchsioides, because "the corolla and long exerted stamens put one in mind of 
some gigantic Fuchsia" (Nuttall 1834). 

Bentham (1835) applied the name Macr anther a fuchsioides (Nutt.) Benth., 
because Conradia had been previously applied to a genus of Gesneriaceae by 
Martins. The name Macranthera means "long anther" Bentham and LeConte 
were both originally cited as authors for the genus (Hooker 1835), although 
Bentham (1835) felt that Torrey alone should receive credit for the generic name. 

Torrey (1837) described a second species, M. kcontei (sometimes spelled 
lecontii), from a specimen collected in Georgia in 1831 by LeConte. He believed 
it was distinct because the segments of the calyx were entire, linear-lanceolate, 
and approximately one-third the length of the corolla. In M.fuchsioides the seg- 
ments of the calyx are dentately lobed and often as long as the corolla tube [on 
LeConte's specimen (NY), Torrey applied the name Conradia kcontei Torr., along 
with a note stating the segments of the calyx are entire]. The specimen itself is 
damaged, but an attached illustration depicts the entire sepals. Torrey actually 
published the species (1837) not as C kcontei, but as M. kcontei, since Bentham 
(1835) had determined that Conradia was already in use. 

Pennell (1935) questioned the authenticity of the collection site of M. 
h'contei, which was purportedly along the Altamaha River in Liberty County, 
Georgia, because that site was far removed from other known occurrences of 
the species. However, the species has been observed along the Altamaha River 
as recently as 1997 (M. Hopkins, pers. comm.). 

Rafmesque (1837) proposed many new names for Macra nthera (see Table 1). 

Macranthera flammea Raf. was proposed on the basis of Bartram's original 

However, Rafinesque thought the genus name Macranthera was inappro- 
priate because he did not beUeve the anthers to be of considerable length, and 
he suggested the name Toxopus, meaning "incurved peduncles." He substituted 
Toxopus gymnanthes Raf. for M. hcontei Torr. and Toxopus calycinus Raf. for M. 
fuchsioides (Nutt.) Benth. He further proposed the genus Tomilix meaning "cut 
calyx" for M. fuchsioides, suggesting that this species be treated as either a sub- 
genus or new genus because of the condition of the calyx and the presence of 
what he felt to be a bifid style. He proposed the binomial Tomilix hracteata 
Raf. as an additional name for M. fuchsioides (Nutt.) Benth. The new genus 
Flamaria, as in Flamaria coccinea Raf., was also proposed 

Bertoloni (1853) described a new genus and species, Dasystoma tuhulosa, 
after examination of collections in GH, NY, and PH. Chapman (1860) treated M. 
lecontei as a variety of M. fuchsioides, and Kuntze (1891) resurrected the genus 
Conradia on the basis of the type specimen that Torrey had originally identi- 
fied as C. lecontei. 

Pennell (1913) combined the two species of Macranthera into one, writing: 
"this difference [m the calyx] is due wholly to age of the flower, as may be readily 
seen in the field; young flowers are as in lecontei, while with age the sepals con- 
tinue growing and become lobed while the corolla shrivels and shortens in situ, 
so bringing about \ht fuchsioides state." Pennell suggested that, because Bartram 
originally described the species, the accepted name must become Macranthera 
flammea (Bartr.) Pennell. He argued that Bartram named his species for its 
"flame-coloured" flowers which is a fitting description. Gerardia was rejected 
as a generic name because it is a synonym for Agalinis. 

Later, many were concerned about the validity of Bartram's binomials 
(Fernald 1944; Rickett 1944; Merri 1 1 1945). Rickett (1944) argued that Bartram's 
binomials are inappropriate because they appear in a publication that is not 
always consistent with the Linnean system of binomial nomenclature. Merrill 
(1945) defended Bartram's binomials, writing: "the proportion of binomials to 
polynomials, 358 to 2, is too great in favor of the former to justify the elimina- 
tion of Bartram's work." Wilbur (1971) argued that, if a Bartram binomial is ac- 
companied by an adequate description, the publication of the binomial should 
be considered valid. 

Additional controversy surrounds Bartram's description of his collection 
site. He probably confused the locality of his specimen of Macranthera with 
that of the mint Calamintha coccinea (Nutt.) Benth. While traveling along the 
Tensaw River near "Taensa" (near or in Baldwin Co, Alabama), Bartram (1791) 
recounts the following observations: "These stony gravelly heights produce a 
variety of herbaceous plants, but one in particular I shall mention on account 

of its singular beauty. I believe it is a species of Gerardia (GerardiaJlammeaX it 
grows erect, a single stem from a root, three or four feet in height, branching 
very regularly from about one half its length upwards, forming a cone or pyra- 
mid, profusely garnished with large tubular labiated scarlet or flame coloured 
flowers, which give the plant a very splendid appearance even at a great dis- 
tance." This description closely describes, for the most part, the form and ap- 
pearance of M.Jlammea, but the locale and scarlet flower color are more appro- 
priate for the mint Calamintha coccinea (Merrill 1945). Unlabeled specimens 
of both plants were found in Bartam's collection at the Natural History Mu- 
seum in London (Pennell 1935). Bartram's description is applicable in part to 
both plants (Table 2), but Pennell (1935) believed Bartram's description to be 
most applicable to M.Jlammea. 

Macrantheraflammea is generally an herbaceous biennial of the Orobanchaceae 
Rosette leaves on juvenile plants are oblanceolate to spatulate. Leaves are gla- 
brous with entire or pinnately lobed margins, depending on developmental 
stage. Rosettes are variable in leaf number and leaf size, having 3-46 leaves per 
rosette with lengths of 3-33 cm and widths of 0.8-3.6 cm (Fig. 1). 

Plants may be more appropriately classified as monocarpic perennials be- 
cause they may remain m the rosette phase for more than one year Rosettes 
that have attained a critical size are triggered to reproductive states by an in- 
crease in day length. Before a rosette bolts into a flowering adult, it begins to 
produce pinnately lobed leaves. 

Reproductive plants of M.Jlammea are mostly 15-30 dm tall, with stems 
that are obtusely four-angled and essentially nonbranchmg below the 
candelabralike spreading inflorescence. Cauline leaves are opposite and sessile 
or have short, winged petioles. Lower leaves are 8.5-16 cm long and 2-6 cm wide, 
lanceolate to ovate, and pinnately lobed. Leaves are progressively reduced up- 
ward along the stem; the uppermost are 4-7 cm long and 1-2.5 cm wide, slightly 
lobed, and toothed or entire. Blades are glabrous, or with minutely ciliate mar- 
gins. Rosettee and lower leaves are usually shed by the time of anthesis. 

branches (Fig. 2). Flowers are axillary to a reduced, usually entire, foliar bract 
(see illustration m Godfrey and Wooten 1981). Pedicels are 1-2 cm long, becom- 
ing ref lexed in fruit. The calyx is fused with five short lobes that are equal and 
linear and 7.5-15 mm long. The corolla is bright orange, fleshy, 2-2.5 cm long, 
short pubescent without, and glabrous or slightly pubescent at the base within. 
A whitish nectar chamber at the corolla base is 4 mm long. 

The corolla turns brown to black with age. The tube is cylindrical and much 
longer (20-23 mm) than the two-lipped lobes (3-4.5 mm), with an adaxial lip 

iy gravelly heights 

that IS two-lobed and erect and a three-lobed, spreading abaxial lip. The four 
contiguous stamens are equal in length and exerted. The filaments are orange 
and become about twice as long as the corolla tube, up to 46 mm. The ovary is 
ovoid and has a slender style that is to 46 mm and a linear-clavate stigma. Ma- 
ture bicarpellate capsules are 9.5-13 mm long, 7.5-10.5 mm wide, loculicidal, 
and slightly pubescent. The seeds are brownish black, 2.5-3 mm long, half as 
wide, irregular in shape (angular-lunate to triangular) with two or three fluted 
membranous wings. Specimens of M.flammea become black upon drying (due 
to presence of orobanchin or iridoid compounds as in related parasitic taxa). 

The species description is verified by examination of collections (Appen- 
dix 1) from major and regional herbaria: Angus K. Gholson (personal collection 
designated AKG; it will eventually become part of FLAS), University of Florida 
(FLAS), Florida State University (FSU), Gray Herbarium (GH), University of 
Southern Mississippi (no official abbreviation, designated HATT here for con- 
venience), Jacksonville State University (JSU), Louisiana State University (LSU), 
Missouri Botanical Garden (MO), New York Botanical Garden (NY), Old Do- 
minion University (OD), Academy of Natural Sciences of Philadelphia (PH), 
Smithsonian Institution (US), University of Georgia (GA), University of Missis- 
sippi (MISS), University of South Alabama (USAM), Valdosta State University 
(VSC), and Vanderbilt University (VDB). 

The species description is also supplemented by measurements taken from 
several plants at two field sites: Thomas Co., Georgia, at Greenwood or GWD 
{Alford 1077 at FSU) and Liberty Co., Florida, in the Apalachicola National For- 
est or ANF (Alford 1064 at FSU). 

Range and Habitat.— The range of M.flammea (Fig. 3) extends along the 
lower coastal plain from Georgia to southeast Louisiana. This species requires 
mesic habitats of bogs and thickets and is ecotonal, occurring in the sandy acid 
soils of pine f latwoods along the margins of seepage slopes and shrub-tree bogs 
or bays (Small 1933). It can occasionally be found in shallow water of cypress- 
gum ponds or depressions (Godfrey & Wooten 1981). It does not tolerate stand- 

(Determann et al. 1997). 

Macranthera flammea 

Woody plants associated with M.fla mmea include: Acer rubruniL., Am 
arbutifolia (L.) Ell., Ckthra alnijolia var. tomentosa (Lam.) Michx., Clifto 
monophylla (Lain.) Britt. ex Sarg., Cyrilla racemiflora L, Fraxinus carolinU 
Mill., Ilex coriacea (Pursh) Chapm., Ilex glabra (L.) Gray, Itea virginica 
Liriodendron tulipijera L., Lyonia lucida (Lam.) K. Koch, Magnolia virginu 
L., Myrica cerifera L., Nyssa syhatica var. biflora (Walt.) Sarg., Persea palus^ 

(Raf.) Sarg., Pinus elliottii En^ 
Rhododendron viscosumd.)! 
taxa include: Andropogon vir 
sulcatum (Fern.) H. Robins., 
gladiata Walt., Elephantopus 
integrijolium W 

(L.) Kuhn. 

lus palustris Mill, Pinus serotina Michx., 
urnum nudum L. Herbaceous associated 
L, Aristida stricta Michx., Arnoglossum 
naria gigantea (Walt.) Muhl, Coreopsis 
Gray, Erianthus giganteus (Walt.) Muhl, 
'ianthus angustifolius L., Hypericum 

Lam., Hypericum galioides Lam., Hypericum tetrapetalum Lam., 
2 (Raf.) Shinners,Juncus valtaus Coville, ludwigia hirtella Raf., Os- 
^amomeaL., Panicum scoparium Lam., and Pteridium aquilinum 

Common Name.— This species is commonly referred to as flame flower or 
hummingbird flower. In at least one instance, it was referred to as Spanish prin- 
cess (Pickens 1955). Flame flower is an appropriate name for M.flammea. It is 
representative of the specific epithet because of the flame-like appearance of 
the orange-yellow corolla (and the candelabra branching habit lends a flame 
shape to the inflorescence). Hummingbird flower is less desirable as is could be 
applied to any number of species that have red or orange tubular corollas. 


F. 3. Documented range of M.C 








iistb base pla 


ions on the 

morphology of repeating modular 




ic lca\cs 

tloweis etc 

because, akhough the numbei 



J size ot the \ 

s hole plain 


teatly with age and environmer 
tern size and form of modular m 



1Q77) DiKeK 



lepend on the morphological form 

Ol 1 


ing units Vj 

nation in l 


nay exist across the range of a s 


individual flo^ 
is affected by i 

the iniloiescer 

ndetei mmate as defined by Harper (1977). 
1 lowers and leaves, and the position of an 
ce letlects its age. By measuring changes 

■ detei mined how each individual flower 

■ architecture of an individual plant and 
in stating that the differences (in calyx 

length and lobmg) between the two species recognized at that tnne (M. 
fuchsioides and M. hcontei) were m fact attributable to age. 

We examined herbarium specimens (Appendix 1) to determine the extent 
of variability of morphological characters across the species' range. Characters 
considered were calyx length, corolla tube length, corolla lobe length, corolla 
width, pedicel length, bract length, fruit length, and fruit width. Because it was 
not always possible to know which part of the plant was represented by the 
herbarium specimen, we standardized measurements by choosing floral buds 
at the stage of development just prior to the separation of the corolla lobes. 
Lengths and width of mature fruits and, when available, the length and width 
of the lower and uppermost (just below the inflorescence) stem leaves were 

To examine the extent of variability within a population and also within a 
single plant, we randomly selected five plants from the established study site at 
GWD in Thomas Co., Georgia, that represented the range of environmental 
conditions for the site. Three flower buds (upper, median, and lower) were 
marked on each of three racemes of the inflorescence (terminal, upper axillary, 
and lower axillary), for a total of nine measured buds per plant (Fig. 4). Charac- 
ters previously mentioned were then measured on marked buds every four days 
for a period of one month. We measured fruit and foliage on 12 plants randomly 
selected from the population. 

Means and standard deviations were calculated for all measured charac- 
ters from herbarium specimens (Table 3). Means and standard deviations of flo- 
ral characters from the GWD sample population were calculated from charac- 
ters measured on 28 August 1997 in order to determine the amount of variability 
for characters within the plant architecture at a given time (Table 3). 

The standard deviations of measured characters (Table 3) for herbarium 
specimens reveal that although the floral tissue components (corolla tube) of 
the flower show little variability variability is higher m the chlorophyll con- 
taining components (calyx, pedicel, and bract). The GWD population exhib- 
ited more variability overall in corolla tube and calyx lengths than the her- 
barium specimens because smaller and larger buds were included. 

Plant 59 of the GWD sample population was selected to illustrate growth 
changes over time in corolla tube length (Fig. 4) and calyx length (Fig. 5) for 
each bud position of the three racemes examined. For each raceme, the corolla 
tube expanded rapidly (Fig. 4). On the terminal and lower axillary racemes, 
the tubes expanded to 22-24 mm before anthesis. Those of the upper axillary 
raceme reached only 20-22 mm. Corolla tube width was between 4 and 5 mm 
for all flowers examined. 

The calyces of the lower and median buds on the terminal raceme (Fig. 5, 
a) had nearly reached their limits of expansion at the initial measuring, but 

orolla lengths from different p 

plateau. Calyces oi buds on the upper and lower axillary racemes (Fig. 5, b-c) 
also demonstrated rapid expansion before leveling off. The calyces of the up- 
per buds were much shorter than those of the median and lower buds for all 

Figure 6 illustrates calyx growth for the upper buds of the three racemes 
for three of the plants comprising the sample population. It shows that the up- 
per axillary raceme had shorter calyces than did the lower axillary and termi- 
nal racemes for plants of the sample population. 

Pedicels exhibited the same growth pattern as calyces. Bracts generally did 
not expand further once anthesis had occurred for an individual bud. Bracts 

^^ 1 3. \'iean5 and standard 

deviations for s. 


\ features of Macrantheraflammea. 

Herbanun.spean.ens GWD plants 

X s N X s N 

did exhibit very rapid growth during bud expansion, and some of the bract 
lengths reached 50 mm for lower buds. 

To produce more valid assessment of the amount of variation in calyx and 
corolla tube lengths observed in the G WD population (Table 3), we selected buds 
at various positions within the inflorescence and did not standardize to stage 
of development. The mean lengths of calyx, corolla, pedicel, and bract were 
greater in the GWD sample population because lower buds were included. Fruits 
measured in the GWD population tended to be shorter, narrower, and less vari- 
able than those in the herbarium specimens. 

In general, variation was minimal in length and width of the corolla (Table 
3, Fig. 4). Recent pubhcations (Campbell et al. 1996; Smith et al. 1996; Temeles 
1996) suggest that corolla width is the floral character selectively influenced 
by hummingbirds, a major pollinator of M.flammea (Pickens 1927). 

The position of a flower within the framework of the inflorescence reflects 
its age (Harper 1977). Calyx length is determined by the position of a flower 
within the framework and is therefore determined by age of the flower. Obser- 
vations (Figs. 5, 6) indicate calyces reach a plateau in their growth and do not 
continue to grow after flowering begins. 

Pennell (1913) was correct in stating that the differences m the lengths of 
calyx lobes are due to the age of flowers. However, because lobes do not grow 
after anthesis, the position of the flower unit within the architecture of the in- 
florescence accounts for observed differences in calyx lengths. These growth 
characteristics are typical for plants with indeterminate inflorescences. Sepals, 
bracts, and pedicels are shorter for bud units at the tip of the raceme and longer 
for those near the bottom. Internodes at the tip of the raceme are truncated, 
those at the bottom elongated. Differences in the upper axillary branching po- 

sitions can be attributed to apical dominance. The terminal racemes are i 
the first to flower, followed closely by the lower axillary racemes. The ra 
flower upwards on the inflorescence so that the upper axillary racemes ; 
last to reach anthesis (Fig. 2). 

In general, as much variation appears to occur within a single plant or 
lation as throughout the entire range. Observed variation in the charac 
calyx, pedicel, and bract length can all be attributed to position effects ^ 
the architecture of the plant. Age determines the placement of individual 
units within an expanding inflorescence and therefore influences chc 

and the GWD popu 

ALABAMA. Baldwin Co.: 7 Sep 1912, F. W. Penndl 4553 (US, PH); 15 Oct 1940, M.G. Henry 2445 (GA); 
20 Aug 1947, M.G. Henry 4935a (PH); 23 Aug 1947, M.G. Henry 4942 (PH); 16 Sep 1964. S.M. McDaniel 
5314 (FSU); 11 Sep 1979, W.S.Judd 2288 (FLAS); 20 Aug 1981, R. Krai 67854 (VDB). Butler Co.: 11 Oct 

yield suffi 

vations of he 
cient evidenc 



ognition of subspecific 


smM. flan 



examined ar 


ated docum 

logical va 
of species 

iabihty of Macranthe 



'iedjield&JD Sn 

1951,D Demo 


1 D Den 

Jackson Co.: 11 bcp 1889 F5 Earle (NY), 16 Aug 1947 MG 
:i,cc 2S202 (N\ ) 20 Aug 1949, D Demaree 28300 (NY VDB), 
7 August 1952 R L Diena 509 (MO), 24 Aug 1953, R B Chan- 
ncll 3^3 (VDB), 11 Sep 1975, M Arguelles 1455 (VSC) Lamar Co.: 14 Aug 1927, F Cooh (US) Layton 
Co.: 15 Aug 1923, F Coo l2 (US) Marion Co.: 8 Aug 1955 J D Ra^(GA,GH,NY),5Sepl963 R Krai 19389 
(VDB) Pearl River Co.: 19 Sep 1982, SM McDam el 26507 {FSV) Perry Co.: 15 Aug 1947, MG Hen))' 
4904 (PH), 4 Oct 1995, S W Lconani 9J8J (FSU, 28 plants) Stone Co.: 24 Aug 1953, D Demmee 34081 
(GH, VDB), 8 Oct 1966, TM Pullen 661140 (UM) Winston Co.: 6 Sep 1932, FH Saygent (NY) County 
unknown: 30 Aug 1941, G.A. Girhart 87 (NY). 

We thank the Greentree Foundation for allowing access to the Big Woods at 

port in the field. We thank Jean C Puti 

for use of botanical illustrations. Nancy C. Coile, Sarah Matthews, A.B. Thistle, 

and an anonymous reviewer provided helpful suggestions on the manuscript. 

AiFORD, J.D. 2000. Taxonomy, reproductive biology, and demogi 

flammea (Batr.) Pennelf M. S.TIiesis, Florida State Univ.Jallahass 
BAHrRAM,W. 1791. Travels in North & South Carolina, Georgia, East ar 

and Johnson, Philadelphia, PA. 

NTHAM, G. 1 835. Synopsis of the Gerardiaeae, a tribe of 

the Scrophulariaceae.Compan- 

ion Bot. Mag. 1:198-212. 

iRTOLONi, A. 1 853. Memorie della Accademia della Scien 

ze dell' Institute di Bologna 4:75. 

AMPBEU, D.R., N.M. Waser, and M.V. Price. 1 996. Mechani 

sms of hummingbird-mediated 

selection for flower width in /pomops/soggregata. Ecology 77:1463-1472. 

HAFiN, E.G. 2000. Field guide to the rare plants of Florida 

.Florida Natural Areas Inventory, 


HAPMAN.A.W. 1860. Flora of the southern United States. 1 


oiLE, N.C. 2000. Notes on Florida's endangered and thr^ 

?atened plants. Contribution no. 

38, 3rd edition. Florida Dept. of Agriculture & Consun 

ler Services, Division of Plant In- 

dustry, Gainesville. 

ETERMANN, R., L.K. KiRKMAN, and H. NouRSF. 1 997. Plant conse 

rvation by propagation.Tipularia 


cities ,n the flora of eastern North 

America. Rhodora 46:1-21. 

OLiFREY, R.K.and J.W. Wooten. 1 981 .Aquatic and wetland plants of the southeastern United 

States: dicotyledons. University of Georgia Press, Ath^ 


ARPER,J.L. 1977. Population biology of plants. Academic 

Press, New York. 

5 Companion Bot.Mag. 



1 Plant 

arum 1:459. 


1945. In defense of 

the va 


m Bartram's bin 

omials. B 



!34. A description of 


ofthe rarer lit 

tie known plant 


United States,from the dried specimens in the herbarium of the Academy of Natural 
Sciences in Philadelphia.]. Acad. Nat. Sci.Philadelphia 7:88. 

Olmstead, R.G., C.W. dePamphieis, A.D. Wolfe, N.D. Young, W.J. Eusons, and PA. Reeves. 200 1 . Disin- 
tegration ofthe Scrophulariaceae. Amer. J. Bot. 88:348-361. 

Pennell, F.W. 1913. Studies in the Agalinanae, a subtribe of the Rhinanthaceae. Bull.Torrey 
Bot. Club 40:1 19-1 30. 

Pennell, F.W. 1 935. The Scrophulariaceae of eastern temperate North America. The Acad- 
emy of Natural Sciences of Philadelphia, Philadelphia. 

Pickens, A.L. 1 927. Unique method of pollination by the ruby-throat. Auk 47:346-352. 

Pickens, A.L 1955. The bird-flower as the apex of floral color display. Castanea 20:1-18. 

Rafinesque, C.S. 1 837. New flora and botany of North America 2: 58-72. Philadelphia. 

RicKETT, H.W. 1944. Legitimacy of names in Bartram's "Travels." Rhodora 46:389-391. 

Small, J.K. 1 933. Manual of the southeastern flora. 1 972 Reprint. Hafner, New York. 

Smith, C.E., J.T Stevens, E.J. Temeles, RW. Ewald, R.J. Hebert, and R.L Bonkovsky. Effect of floral 
orifice width and shape on hummingbird-flower interactions.Oecologia 1 06:482-292. 

Temeles, E.J. 1996. A new dimension to hummingbird-flower relationships. Oecologia 

ToRREY,J.1837.An account of several new genera and species of North American plants. 




Myrna R Landim^ Leslie R. Landrum 

Departamento de Biologia Department of Plant Biology 

Unlversidade Federc 

Arizona State University 

49 100-000, SdoQisto 

vac, SE, BRAZIL 




e Atlantic forest is one oft 



razil that has 

suffered the greatest destruc 



;till imperiled, especially i 




the states where the Atlantic 

: forest has 

m most greatly reduced, 1 

: fragments V 

'arying in size and degree of 


n. Despite their important 

:e, botanic 

:al studie 

;e fragments are still greatly 

needed. In 


s paper we report for the f 

irst time t 


■ence of four 

species of Campomanesiam 

Sergipe: C 



md C. VIC 

itoris. The la: 

St, formally thought to be kn 

,own from 


ibution in Sergipe, and IS four 

id m forest 


d restinga areas of differei 


time from 

> Northeast of Brazil. We hope that s 

the composi 


the local preservation of 1 


item and 

help future ; 

programs of forest restoratio 

n We em- 


nore studies ■ 

on plant taxonomy and the t 


«. specialists. 





s brasilei 

ros mais des 


lo, sendo a 


ciada no ^ 

>Jordeste . 

do BrasU. Un 

a de mata 


antica, Sergipe apresent: 

oje algur 

IS fragment( 

Ds florestais com tamanho 

e grau de 






■ saoamda 

de diversos municipios. Registra-se, ainda, i 
ao Nordeste. Espera-se que estudos sobre £ 
1 a preservagao desse ecossistema a nivel re- 
florestal. Salienta-se a necessidade de un: 

The objective of this note is to report the occurrence in Sergipe, Brazil of four 
species of Campomanesia Ruiz et Pavon (Myrtaceae), usually known locally as 
"guabirobas" or a variant of this name (e.g., guavirobas, gobiraba, gabiroba). None 

of these species, nor any other species of Campomanesia, was known for Sergipe 
when Landrum (f986) monographed the genus, and one species (C. guaviroha) 
has not been reported before in Brazil's Northeast region (Maranhao, Piaul, Ceara, 
Rio Grande do Norte, Paraiba, Pernambuco, Alagoas, Sergipe, Bahia). The fact 
that recent floristic exploration can reveal so much is an indication of the im- 
portance of continuing efforts in this field. 

Despite the great biological diversity of the Brazilian coastal forests, little 
of its original cover still persists. At the time of the European discovery such 
forest covered about 1,000,000 km- of Brazil, while today its area has been re- 
duced to an estimated 5% (Consorcio Mata Atlantica 1992), 6% (Prance et al. 
2000), or 8% (INPE and IBAMA 1990) of its original size. In spite of being pro- 
tected by law, destruction of coastal forest still continues. The remaining areas 
consist of fragments varying in size and degree of conservation and are "typi- 
cally small, isolated and highly disturbed" (Viana et al. 1997). Ranta et al. (1998) 
recently studied the degree of fragmentation in the Northeast Atlantic forest 
of Brazil, and emphasize the importance of fragment size and shape for the long- 
term survival of biodiversity. Smaller fragments and more irregularly shaped 
fragments have a higher proportion of their areas in edges, which are more vul- 
nerable to plant extinction. Because fragmentation in Sergipe has progressed so 

The historical pattern of almost complete elimi 

nation of the native forest 

covering m Brazil's Northeast is explained by the n 

elative gentle relief of the 

landscape and ease of access to all areas. Farther so 

uth, the mountainous ter- 

rain of the Serra do Mar makes access more difficult. 

, and thus greater portions 

of Atlantic rainforests there are relatively protected. Sergipe, the smallest of the 
Brazilian states, originally had about 41% its area covered by forests (Campos 
1912), but Its forest covering today is estimated to be less than 1%. For this rea- 
son, Sergipe has been excluded in some mappings of the remnants of Atlantic 
forest carried out at the national level. ITowever, floristic surveys in areas of 
Atlantic forest in the state (Landim et al. 1998) have shown it to be a forest with 
unique floristic and structural characteristics. Additional studies are urgently 
needed that contribute to knowledge of the forest's composition, structure and 
dynamics, with the aim being a knowledge base for reforestation initiatives. 

The family Myrtaceae is mainly a tropical and subtropical family 
(hley wood 1993), with about 3500 species and approximately 100 genera, with 
two main centers of development: tropical America and Australasia (Barroso et 
al. 1991). It is a family of great importance in the neotropical forests, being one 
of the dominant families in Atlantic coastal forest (Barroso & Peron 1994; Leitao- 
Filho 1993; Mori et al. 1983; Peixoto & Gentry 1990; Reitz et al. 1978). In a study 
of Atlantic forest in Sao Paulo, Mantovani (1993) found 38 tree species of 

istic family of arboreal species of Atlantic forest in the south coast of the State 
of Sao Paulo, with f loristic and structural importance." 

In general, species of Myrtaceae are quite common in the Atlantic forest 
areas of the Northeast, (Siqueira 1994), and this has proved to be true the frag- 
ments studied in Sergipe. In a phytosociological study in the Mata do Crasto, in 
the Municipality of Santa Luzia do Itanhy (Landim et al. 1998), this family ac- 
counted for 10.3% of the sampled trees, and was the sixth in basal area, and 
fifth in importance value index (I.V.I.). 

Voucher specimens of collections described below are deposited in the her- 
baria of the Department of Biology of the Federal University of Sergipe (ASE), 
the University of Brasilia (UB), and Arizona State University (ASU). 

Campomanesia can be distinguished from other genera of Myrtaceae by: 1) ovary 
with (3-)4-18 locules (Fig. ID); 2) ovules several per locule, biseriate, all or all but 
one aborting in each locule; 3) locule-wall in the mature fruit chartaceous to 
slightly woody strongly glandular, serving as a false seed coat (Fig. IC). In Sergipe, 
Campomanesia can be distinguished from most other genera of Myrtaceae when 
in flower by its 5-merous flowers and inflorescences of solitary flowers or dicha- 
sia. Psidium L. and Calycolpus O. Berg can be confused with Campomanesia in 
flowering specimens but generally have coriaceous leaves, whereas 
Campomanesia usually has submembranous to membranous leaves. 

Campomanesia is quite distinctive in fruit. There seem to be several seeds 
in each fruit arranged in a ring, each with a glandular covering (Fig. ID), fiow- 
ever, each of these "seeds" is a locule, and usually only some of them have a seed 
inside. In others the ovules have all aborted. Other genera of Myrtaceae either 
have few seeds (e.g., 1 or 2 in Myrcia, Eugenia, or Calyptranthes) or the seeds are 
numerous and hard (e.g., Psidium, Calycolpus). In no other American genus of 
Myrtaceae do the seeds have a glandular covering. 

The bark of the trunk of Campomanesia is usually rough with numerous 
narrow, papery plates, unlike most other genera that have smooth to scaly bark. 
The lateral veins of the leaves are generally prominent and broadly arch near the 
margin and the smaller veins often show an intricate reticulate pattern (Fig. 2). 


Campomanesia viatoris 

Campomanesia aromatica (Aubl.) Grisebach 

Shrub or tree 2-20 m high; leaves immature at anthesis, eUiptic, ovate, lanceolate 

or oblanceolate, 4-12(-15) cm long, 2-4(-7.5) cm wide, 1.7-4 times as long as 
wide; peduncles 0.3-2.6 cm long, uniflorous; bracteoles 1-4 mm long; calyx- 
lobes hemiorbicular to oblong-truncate, 2-4.3 mm long; hypanthium obconic 
to campanulate, 1.5-2.5 mm long; stamens 80-90, 4-7 mm long; anthers 0.5-0.8 
mm long; ovary 4-6-locular; ovules 4-7 per locule; fruits ca. 1 cm long, black. 

1 locally as "gabiroba" or 
atinga region was called 
"cadeia brava." 

In his revision of this genus, Landrum (1986) cited only a few collections 
of C aromatica in the Northeast, namely in the states of Maranhao, Ceara, Rio 
Grande do Norte, Paraiba and Bahia. The species also occurs in Bolivia (Landrum 
1986), an interesting disjunction of over 2000 km. In the present study, it has 

f coastal forest (Mata Atlantica) as well as caatmga, 
dry areas. A greater sampling effort is needed to bette 

I dichotoma (O. Berg) J.R. Mattos 
Shrub or tree up to 10 m high; leaves mature at anthesis, elliptic, elliptic-oh 
long, ovate or suborbicular, 3-9.5 cm long, 1.5-6 cm wide, 1-2.3 times as long a 
wide; peduncles 2-6 cm long, bearing 3-15 flowered dichasia; bracteoles ca. 
mm long; calyx nearly closed in the bud, the calyx-lobes 1-2 mm long, the ca 
lyx tearing between the lobes at anthesis; hypanthmm (including closed ca 
lyx) 5-8 mm long; stamens 200-350, 3-10 mm long; anthers 0.8-1 mm lonj 
ovary 6-10-locular; ovules 7-18 per locule; fruits ca. 1 cm long. 

Specimens examined: Mun. Santa Luzia do Uanhy: Mata Atlantica, Mata do Crasto, 19 Apr 1995, A 

The common name "bacalhau" is used for this species as well as the more com- 
mon generic designation of "gobiraba." 

Campomanesia dichotoma is known from the coastal region of the North- 
east of Brazil and the state of Rio de Janeiro (Landrum 1986). So far in Sergipe, 
examples have been found in only one fragment of Atlantic coastal forest. 
Campomanesia guaviroba (A.R de CandoUe) Kiaerskou 
Tree up to ca. 12 m high; leaves mature at anthesis, mainly elliptic, less often 
ovate, lanceolate, suborbicular, or ovate, 4-13 cm long, 1.7-8 cm wide, 1.4-3 times 
as long as wide; peduncle 0.3-2.5 cm long, unif lorous; bracteoles ca. 4 mm long; 
calyx-lobes broadly triangular or rounded, 1.5-3 mm long; hypanthmm obconic 
to campanulate, 4-6 mm long; stamens 250-500, 3-9 mm long; anthers 0.5-1 
mm long; ovary 7-i44ocular; ovules 13-20 per locule; fruit 1-3.5 cm long, yel- 
low or orangish. (Fig. 1 B), 

Until recently Campomanesia guaviroba was known only from Brazil's Atlan- 
tic forest from Espirii 
Paraguay (Landrum 

the north (ca. 1200 km) and are the only ones known to us from the Northeast 
of Brazil. Recently a collection has extended the known range to Bolivia [Santa 
Cruz, Velasco Province, Parque Nacional Noel Kempff Mercado, 5 km S del 
campamento Las Gamas (14" 4814"S, 60° 23'59"W), 850 m, A Rodriguez &J. 
SuruhV 558 (ASU)l h thus has a disjunct distribution similar to that of C 

In one area of Sao Paulo (Mantovani 1993) this species was represented by 
only two individuals, having an I.V.I of 0.58. Apparently rare, it was not found 
in the forest gap areas in the same study, although other Myrtaceae were, e.g., 

Eugenia, Marlieria, Calyptranthes, Myrcia and Gomidesia. This may mean that 
it is less tolerant of disturbance (anthropogenic or not) and thus is at a higher 
risk of going extinct through habitat disturbance and fragmentation. 
Campomanesia viatoris Landrum 

Shrub or tree to ca. 12 m high; leaves mature at anthesis, elliptic to ovate, 3-13 
cm long, 2-6.5 cm wide, 1.5-2 times as long as wide; peduncle 0.5-2 cm long, 
unif lorous or bearing a 3-f lowered dichasium; bracteoles ca. 3 mm long; calyx- 
lobes truncate-auriculate, 1-3 mm long; hypanthium 8-10 mm long, attenuate 

thers ca. 1-2 mm long; ovary 7-8-locular; ovules 11-16 per locule; fruit globose 
except for an attenuate base, up to ca. 2.5 cm in diameter (Fig. 1 A). 

Sao Cristovao: restinga, 21 Mar 1985, G. Viana 1112 (ASE). Mun. Pirambu: Reserva Biologica Santa 
Isabel, dunas no km 11 de praia na diregao de Ponta dos Mangues, 28 Jan 1992, C Farney et al. 2944 
(ASU). Mun. Santa Luzia do Itanhy: cerca de 2.5 km do Distrito de Crasto, na estrada para Sta. Luzia 
do Itanhy, 27 Nov 1993, A.M. A. Amorim et al 1500 (ASU); Mata Atlantica, 7 Apr 1993, M. Landim 282 
(ASE, ASU, UB), 3 May 1995, M. Landim 355 (ASE, ASU, UB), 5 Jul 1995, M. Landim 456 (ASE, ASU, 
UB); Mata Atlantica, Mata do Crasto. 7 Apr 1995, M. Landim 285 (ASE, ASU, UB), 16 Dec 1996, M. 
Landim 1108 (ASE, ASU), Mun. Sao Cristovao: restinga, 9 Apr 1996, M. Landim 904 (ASE, ASU). Mun. 
Areia Branca: Mata, Serra de Itabaiana, 28 Mar 1997, A. Vicente 7 (ASE). 

The common names for this species are "guabiraba," "gabiroba," "gabirobinha," 
"gobirabinha," and "gobiraba." 

Until recently only two collections for this species were known, the type 
and one other, both from Alagoas (Landrum 1986). The type was collected by 
Gardner over 150 years ago and the additional specimen has no date. Since the 
species was thought to be extinct or near extinction, Landrum (1986) urged 
botanists m Alagoas to search for this rare species. The collections cited above 
from Sergipe indicate that Campomanesia viatoris is at least locally common 
m the state and additional specimens have also been found in Bahia. le.g., Mun. 
Apora, 31 km S of Olmdina along highway BR-116, 310 m, 1 Apr 1976, G. Davidse 

The anthers of Campomanesia viatoris anthers are unusually elongate and 
similar to those of C. laurifolia. This unusual shared anther morphology may 
indicate an interesting pollination mechanism deserving more careful study. 

Phytogeographic analysis of species of Myrtaceae is greatly hampered by the 
difficulty in identifying material, due to the fact that American species of 
Myrtaceae often are very similar in the majority of their characters (McVaugh 
1968), and because of the cryptic nature of the characters used to identify the 
genera (Landrum and Kawasaki 1997). Some floristic and/or phytosociologi- 
cal studies list great numbers of species or individuals of Myrtaceae, but au- 
thors often are unable to identify them to species or genus (e.g., Guedes 1992). In 

a study of the flora of Atlantic rainforest, Myrtaceae are cited as the family with 
the greatest number of unknown specimens (Siqueira 1994). 

Ecological studies of the Northeast of Brazil will require many more spe- 
cialists to identify the great number of collections unknown to species, genus, 
or even family (Siqueira, 1994). Three conditions need to be improved: 1) there 
needs to be more collecting in general as this paper demonstrates; 2) 
monographers must study the flora of the Northeast and include specimens 
from the regional herbaria; and 3) there should be a greater investment in train- 
ing more specialists. 

The absence of reports of Campomanesia in Sergipe in Landrum's 1986 re- 
vision of the genus reflects the low sampling effort in the state until recently. It 
is still difficult to describe the distribution of the species in the state because 
they are known from so few collections. The present study indicates the exist- 
ence of a flora that is still relatively unknown in the fragments of Atlantic 
rainforest in Sergipe and indicates the importance of the exploration and con- 
servation of these areas. 

The sympatric occurrence of these four Campomanesia species m one frag- 
ment, the Mata do Crasto, in Santa Luzia do Itanhy, a coastal region in the south 
of the Sergipe, provides opportunities for more studies. These might involve the 
identification of possibfe differences between these four species in flowering 
and fruiting phenology, pollinators and fruit predators, and reproductive iso- 
lating mechanisms that may have led to speciation. 

Finally, it is worth noting that even in regions with forest coverings insuf- 
ficient to be registered m some national surveys, such as Sergipe, the forest rem- 
nants that do exist are the last representatives of an entire ecosystem, includ- 
ing plants, animals, fungi, and micro-organisms. These fragments, mostly of 
medium to small size, are the best possible estimation of the original ecosys- 
tem that will ever be available. Studies on the composition of these fragments 
can contribute to the preservation of that ecosystem at the regional level and 
will be useful for future reforestation programs. In order to avoid the drastic 
loss of large portions of biodiversity, the establishment of reserves is necessary, 
at least for some of these fragments, and conservation management that in- 
cludes corridors between fragments as well as the involvement of neighboring 
human communities in this process is desirable 


V offered many helpful suggestions for which 


Jniversidade Federal de Vigosa,Vi(;osa. 

Barroso, G.M. and V. Peron. 1 994. Myrtaceae. In: M. P. M. Lima and R. R. Guedes-Bruni, orgs. 

Reserva ecologica de Macae de Cima, Nova Friburgo, RJ. Aspectos flori'sticos das 

especies vasculares., Vol. 1 . Pp. 261-302. 
Campos, G. 1912. Mappa florestal. Rio de Janeiro, Servigo Geologico e Mineralogico do 

Brasil.Consorcio Mata Atlantica/UNICAMR 1 992. Reserva da Biosfera da Mata Atlantica: 

Piano de Agao. Referencias basicas, 1 . Campinas. 
GuEDEs,M.L.S. 1 992.Estudo floristico e fitossociologico de um trecho da Reserva ecologica 

da Mata de Dois Irmaos Recife - Pernambuco.Tese de Mestrado, Universidade Federal 

Rural de Pernambuco, Recife. 
HEYwooD,V.H.(ed.) 1983. Flowering plants of the world. Oxford University Press, New York. 
Inpe (Instituto Nacional de Pesquisas Espaciais) and Ibama (Instituto Brasileiro do Meio Ambiente e 

DOS Recursos Naturais Renovaveis) 1990. Atlas dos remanescentes florestais do dominio /index.asp. 

nal de Botanica.Sociedade Botanica do Brasil, Salvador. Resumes. 
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und Lumo (Myrtaceae). FI.Neotrop.,Monogr. 45. The New York Botanical Garden, New 

.ANDRUM, L.R. and M.L. Kawasaki. 1 997.The genera of Myrtaceae in Brazil: an illustrated syn- 
optic treatment and identification keys. Brittonia 49:508-536. 
eitao-Filho, H.F (org.). 1 993. Ecologia da Mata Atlantica de Cubatao (SP). Editoras UNE5P/ 

/IcVaugh, R. 1968.The genera of American Myrtaceae-an interim report.Taxon 17:354- 

/Iantovani, W. 1 993. Estrutura e dinamica da floresta atlantica na Jureia, Iguape-SRTese de 

Livre Docencia. Universidade de Sao Paulo, Instituto de Biociencias, Sao Paulo. 
/1ori,S. A., B.M. Boom, A.M. CARVALH0,and T.S.Santos. 1983. Ecological importance of Myrtaceae 

in an eastern Brazilian wet forest. Biotropica 15:68-70. 
'EixoTo,A.L.and A.Gentry. 1990. Diversidade ecomposigaofloristica da mata detabuleiro 

na Reserva Florestal de Linhares (Espirito Santo, Brasil). Revista Brasil, Bot. 1 3:1 9-25. 
rance, G.T., H. Beentje, J. Dransfield, and R. Johns. 2000. The tropical flora remains 

undercollected. Ann. Missouri Bot. Card. 87: 67-71 . 
ANTA, P., T. Blom, J. Niemela, E. Joensuu, and M. Siitonen. 1998. The fragmented Atlantic rain 

forest of Brazil: size, shape and distribution of forest fragments. Biodiversity and Con- 

ti!z, R., R.M. Klein, and A. Reis. 1 978. Projeto madeira de Santa Catarina. Sellowia 30(28- 

lANA, V.M, A.A.J. Tabanez, and J.L.F. Batisia, 1 997. Dynamics and restoration of forest frag- 
ments in the Brazilian Atlantic moist forests. In: W.F.Laurance and R.O. Bierregard,eds. 
Tropical Forest Remnants — Ecology, Management, and Conservation of Fragmented 
Communities. University of Chicago Press,Chicago, IL. Pp.351-365. 


Terrell (2001) presented a taxonomic study of the genus Stcnaria (= Hedyotis 
nigricans and closely related cohorts, sensu Turner 1995). In this he recognized 
H. nigricans var. gypsophila as occurring in Trans-Pecos, Texas, citing among 
his "selected representative specimens" four sheets (cf. Fig. 1), two from 
Culberson Co., a north-south ridge of limestone outcrops extending for ca. 50 
miles from near Van Horn, Texas to near the New Mexico border, and one from 
the same mountain range in closely adjacent Hudspeth Co.; the fourth collec- 
tion (Webster 4501, MICH) was said to be from the "Madera Canyon" m igneous 
soils of Jeff Davis County 

Terrell distinguished H. nigricans var. gypsophila (Fig. 3) from the rest of 
his concept of the wide-ranging, highly variable H. nigricans by the following 

west Texas var. gypsophila 

longer than wide; corollas 2-8 mm long [var. nigricans and closely related c 

Careful scrutiny of the above key will show that the only meaningful cl 
ter used by Terrell to define his "var. gypsophila" is leaf shape. Indeed, Terrell 
himself (2001), states that "the most conspicuous character tdistinguishing be- 
tween these elements] is the elliptical leaves on rather small plants...Iand] 
not find any other characters significantly different from var. nigricans..." 

Regardless, I would like to place on record here that I take many if not 
most, of the specimens cited by Terrell (2001) as occurring outside of the re- 
gional distributions of H. n. var. gypsophila (Turner 1995) to be misidentifications 
of H. n. var. nigricans, these but leaf and/or habital forms occurring among oth- 
erwise typical populations of var. nigricans. 

This can be readily attested to by reference to the single sheet (Webster 4501) 
of H. nigricans var. gypsophila cited in the above introduction. A duplicate of 
this collection at SRSC (Fig. 2) is clearly linear-leaved and could not be identi- 
fied as var. gypsophila by Terrell's key to varieties. Additionally, its habit and 
floral features do not conform to those of var. gypsophila as described by Turner 
(1995). Indeed, 1 have reexamined 200 or more specimens of H. n. var. nigricans 

nigricans liar, nigricans. 

from Trans-Pecos, Texas (LL, SRSC, TEX) and find not a single plant referable to 
H. n. var gypsophila, including most of the specimens so cited by Terrell (2001). 

He further ventures that "I [Terrell] consider the taxon as misnamed, as I 
recorded only one collection out of 49 Mexican collections from 19 herbaria 
that mentioned a gypsum substrate. It may be noted that the type specimen 
came from a gypsum habitat." 

However, had Terrell examined the 49 sheets of this taxon on file at LL.TEX 
he would have found 12 sheets with labels noting their occurrence on gypsum 
substrates, and most of the rest from areas known to possess such soils. The 
truth is that most early collectors did not know the difference between gypsum 
substrates (CaS04) and limestone substrates (CaC04), or else they ignored such 
distinctions. The highly endemic flora occurring on gypsum substrates need 

zr (1977) and Turne 
t should be noted t 
ler 1993) that "Thi; 

snomenon being well c 
ind Powell (1979). 
t Terrell seemingly igno 
axon [var, gjypsophiZfl] is 

collections at LL, TEX, mostly obtained from gypseous soils in the state of Nuevo 
Leon." Tliis is not to say that occasional plants of var. gypsophila in north cen- 
tral Mexico might not approach var nigricans in this or that character, but such 
"intermediates" do not occur in Texas. Indeed, var nigricans can be found in 
relatively close proximity to var gypsophi la m Nuevo Leon without clear inter- 
mediates, the former occurring on calcareous soils (e.g., Hinton et al. 2^351, 
2^575, 25940, etc., all TEX), the latter on gypseous soils (e.g., Hinton etal 24373, 
27183, etc., TEX). In fact, my initial taxonomic sense was to recognize vzr.gypso- 
phila as specifically distinct, but such treatment would deny the occasional 
intermediates, these mostly occurring in regions of near geographical contact. 

It should be emphasized that var gypsophila is distinguished from var 
nigricans by more than habit, leaf shape and calyx vestiture. The upper leaf 
surfaces of the former are more nearly papillose; those of the latter having sur- 
face-sculpturing more like that of the epidermis of a peanut. In short, there are 
a syndrome of characters which provide for the recognition of both taxa, al- 
though the occasional plant from this or that population might lack one or an- 
other of the characters concerned. 

Such identification problems are touched upon by numerous workers, in- 
cluding my own (Turner et al. 1988). Leaf shape and habit vary considerably 
among populations of var nigricans in Texas. Thus, heavily browsed plants will 
appear much branched from the base, and newly produced basal shoots will 
possess somewhat broader leaves. No doubt such plants occasioned the attri- 
bution of var gypsophila to Texas, and probably those Mexican collections cited 
by Terrell as occurring outside the distribution of var gypsophila sensu the 

Finally, I can't help but note that I curre 
nigricans var nigricans m Trans-Pecos, Tex 

populations from throughout this region and nearly all consist of p 
tifiable as var. nigricans as defined by both Terrell and myself, exci 
one might let the occasional over-browsed plant cloud one's identifi 

Delprete, whose comments greatly improved the present contributi 

PowEn,A.M.and B.LTurner. 1977. Aspects of the plant biology of the gypsum outcrops ( 
the Chihuahuan Desert . Natf Park Service Trans. & Proc, Ser. 3:3 1 5-325. 

Terrell, E.E. 2001 .Taxonomy of Stenaria (Rubiaceae: Hedyotideae), a new genus includin 
Hedyotis nigricans. Sida 1 9:591 -61 4. 

Turner, B.L. 1995. Taxonomic overview of Hedyotis nigricans (Rubiaceae) and closely allie 
taxa. Phytologia 79:12-21 





Wendy B.Zomlefer Gerald L.Smith 

Plant Biology Department Biology Department 

University of Georgia High Point University 

2502 Plan t Sciences Box 359 1, University Station 

Athens, GA 30602-7271, U.S.A. High Point, NC 27262, U.S.A. 

The Melanthiaceae sensu the Angiosperm Phylogeny Group (APG 1998) com- 
prise five tribes (11-16 genera; ca. 154-201+ spp.) of predominately woodland 
and/or alpine perennial herbs occurring mainly in the temperate to Arctic zones 
of the Northern Hemisphere (with one species of Schoenocaulon extending into 
South America; Zomlefer et al. 2001). Of concern here are two genera of the 
tribe Melanthieae, Stenanthium s.l. and Zigadenus s.l. As traditionally and 
broadly circumscribed, both genera form poorly defined complexes (summary 
in Zomlefer 1997). Stenanthium s.l. has sometimes been divided into two gen- 
era: a monotypic Stenanthium (5. gramineum) and Stenanthella (2-4 spp.); 
Zigadenus has a more complex taxonomic history involving several proposed 
segregates, although contemporary botanists have typically accepted only the 
monotypic segregate Amianthium with the remaining ca. 19 species maintained 

The circumscription and relationships of genera within the tribe 
Melanthieae were recently evaluated using parsimony analyses of ITS (nuclear 
ribosomal) and trnL-F (plastid) DNA sequence data (Zomlefer et al. 2001). Based 
on the cladograms generated in this study, Zigadenus s.l. is polyphyletic and 

Stenanthium s.l. is biphyletic, with Stenanthium s.s. and Stenanthella embed- 
ded within two different "Zzgadenus" clades. In overview, the species of 
Stenanthium and Zigadenus, as traditionally circumscribed, form five strongly 
supported clades that correlate with distinctive geographical distribution, chro- 
mosome number, and certain morphological characters. The Stenanthium chde, 
comprising Stenanthium gramineumiStenanthium s.s.), Zigadenus densus,and 
Z leimanthoides, is defined by the morphological synapomorphies of a slen- 
der (cylindrical) bulb and one obscure (or lacking) gland per tepal. However, 
the chromosome number for this group, 2n = 20, has been documented only for 
Stenanthium gramineum (Miller 1930; Sato 1942; as S. rohustum, see Fernald 
1946). Since chromosome number is a significant feature for genera of tribe 
Melanthieae, 20 was predicted as the likely mitotic count for the other two spe- 
cies now transferred to the recircumscribed Stenanthium, Stenanthium densum 
and S. leimanthoides (Zomlefer et al. 2001; Zomlefer &Judd 2002). 

Several plants of Stenanthium densum and S. leimanthoides were collected dur- 
ing April to June 2001 (see Table 1) and transplanted to pots maintained at the 
Plant Biology Dept. Greenhouse Facility at the University of Georgia and the 
Biology Dept. Greenhouse at High Point University Dividing root tip cells were 
prepared for examination according to general protocols outlined by Flory and 
Smith (1980), Jones and Luchsmger (1986), and Smith (1984). Once the plants 
were well-established, actively growing root tips were harvested at noon and 
treated with 0.2 % colchicine for 4 hours, rinsed in distilled water, and then 
fixed in Carnoy's solution (3 ethanol: 1 acetic acid) overnight or longer Follow- 
ing this fixation, the roots were rinsed in distilled water, hydrolyzed in 1.0 N 
HCl at 43*^ C for 25 minutes, macerated with a dissecting spatula on a glass 
microscope slide, and stained with 1% aceto-orcein. After application of a cover 
slip, the slide was gently heated with an alcohol lamp, placed between blotters, 
and subjected to additional pressure. Slides were mounted in euparal for future 
reference. Well-spread metaphase chromosomes were traced under a Leica 
DMLB Research Microscope with a camera lucida attachment. Herbarium speci- 
men vouchers (Table 1) are deposited at GA. 


The mitotic chromosome number of 2n = 20 for Stena nthium, as circumscribed 
by Zomlefer and Judd (2002), is verified for all species (Table 1, Figs. 1-3). Chro- 
mosome number is a useful taxonomic character for genera within tribe 
Melanthieae (see Fig. 4), especially the synapomorphic 2n numbers of 20 for 
Stenanthium and 22 for Toxicoscordion. A probable base chromosome number 
of X = 8 is often cited for the tribe Melanthieae (Sen 1975; Tamura 1995; Lowry et al. 

i (McDearman 1984), some ai 

Locality & collection 

distinguishing characters, 

(Desr.) ZomleferS 

Creek, 4 Af 
Zomlefer 7. 

4.3 mi N of Butler, 

792 (GA) 

6.1 mi S of Butler, 

1987; Zomlefer 1997), and multiples c 
lent m the group (Fig. 4): Ani 
Sdxotnocaulon, and sister groups 
Mdanthium). Unfortunately, di; 

number (or perhaps of 4) are preva- 
(which includes Stenanthella), 

aumniumj. uniortunateiy, uue lo the small chromosome size of members of 
e tribe (ca. 2.0-4.0 [im in length), the few karyotype studies (e.g., Lee 1985) are 
not detailed enough to infer a possible mechanism of chromosomal evolution, 
although the chromosome numbers (Fig. 4) indicate the prevalence of polyp- 
loidy and/or aneuploid variation. The functional outgroup for Melanthieae (see 
Zomlefer et al. 2001), Z.glahernmus (Zigadenus s.s.), has an unpublished count 
of 2n = 52, tentatively reported by Preece (1956). Probable base numbers for the 
other immediate outgroup taxa of the tribe Melanthieae (the remaining four 
tribes of the Melanthiaceae) are five and 15, respectively for sister taxa Parideae 



\ >*•< 


Zigadenus s.s. all contain species of the polypliyletic Zigadem 

(Trilliaceae) and Xerophylleae, and 17 and possibly six, respectively, for sister 
taxa Heloniadeae and Chionographideae (Lowry et al. 1987; Zomlefer 1996, 1997). 
Confirmation of the chromosome number for Z. glaherrimus (a possible 
polyploid) and the reassessment of the base number for the tribe merit further 
investigation (Zomlefer m prep.), especially in relation to the probable base 
numbers of the other tribes. Available data, however, already demonstrate that 
chromosome number strengthens support for the monophyly of Toxicoscordion 
and Stenanthium, as recircumscribed. Chromosome counts for the superficially 
similar plants in the Melanthieae validate chromosome number as a signifi- 

cant character, useful for elucidating and predicting groupings and relation- 
ships. Variation in chromosome number has also proven phylogenetically sig- 
nificant in other monocot clades, such as Polygonatum/Polygonateae (Tamura 
1993; Tamura et al. 1997a, b). 


Stephanie Bergamo, Richard Carter, Angus Gholsonjr, David E. Giannasi, Mary 
Jo Godt, and Elissa Totin enthusiastically assisted WBZ m the field; S. Bergamo 
also shared her time and expertise with the laboratory work. We are grateful to 
Andrew W. Tull (Plant Biology Dept. Greenhouse Facility, University of Geor- 
gia) for maintaining plants collected in the field; Walter S. Judd and Douglas E. 
Soltis for helpful comments on the manuscript; and William Carromero for the 
Spanish translation of the abstract. Funds provided by the University of Geor- 
gia Department of Plant Biology financed the collecting trips associated with 
this project. 


p[APG]. 1998. An ordinal classificc 

3tion for the families of flower- 


cations and reidentifications of N 

orth American Plants. Rhodora 

1980. The chromosomes of Hab 

'fanthus martinezil, H. robustus, 

3nt Life 36:54-62, 

nomic study of the Korean Veratr 

urn species. Korean J. Pl.Taxon. 

LORY, W.S. and G.L. Smith. 

and their F, hybhd.Pk 

ovvRY,RP.,RGoLDBLATT,and H.ToBE. 1987. Notes on the floral biology, cytology an^ 

ology of Campynemant/ie(Liliales:Campynemataceae). Ann. Missouri Bot.Garc 

Jones, S.B. and AE.LucHsiNGER. 198 

6. Preparing sguashes of plant root tips. Ir 

r Plant system- 

atics, 2nd ed.,McGraw Hill, Ne 

?w York. Pp. 181-182. 

McDearman, W.W.I 984. Systemat 

:ics of Zigadenus densus and Z leimanthoi 

des and phylo- 

genetic implications of breec 

ling systems in the Veratreae. [Abstract]. / 


MiLiFR, E.W. 1930.A preliminary 

note on the cytology of the Melanthioic 

leae section of 

the Liliaceae.Proc. Univ. Durh 

am Philos.Soc. 8:267-271. 

Preece, S.J. 1 956. A cytotaxonomic study of the genus Zigadenus. Ph.D. dis: 

sertation. State 

College of Washington, Pullm 



Hales. Feddes Repert. 86:255-305. 

Sato, D. 1942. Karyotype alterati( 

on and phylogeny in Liliaceae and allied 


Bot 12:57-161. 

Small, J.K.I 903. Flora of the sou- 

theastern United States.Tublished by th- 

e author," New 

York, New York. Pp. 250, 252. 

Smiih, G.L. 1 984. Revision of Piptocarpha. Ph.D. dissertation. University of Georgia, Athens. 

SoRRiE,B.A.,and A.S.Weakley.2001. Coastal plain vascular plant endemic5:phytogeographic 
patterns. Castanea 66:50-82. 

Tamura, M.N. 1 993. Biosystematic studies on the genus Polygonotum (Liliaceae) III. Mor- 
phology of stanninal filaments and karyology of eleven Eurasian species. Bot. Jahrb. 

Tamura, M.N. 1995. A karyological review of the orders Asparagales and Liliales 
(Monocotyledonae). Feddes Repert, 1 06:83-1 1 1 . 

Tamura, M.N., M. Ogisu, and J.-M. Xu. 1 997a. Heteropolygonatum, a new genus of the tribe 
Polygonateae (Convallariaceae) from west China. Kew Bull. 52:949-956. 

Tamura, M.N„ A.E. Schwar/bach, S. Kruse, and R. Reski. 1997b. Biosystematic studies on the 
genus Polygonatum (Convallariaceae) IV. Molecular phylogenetic analysis based on 
restriction site mapping of the chloroplast gene trnK. Feddes Repert. 1 08:1 59-1 68. 

ZoMLEFER, W.B. 1996.TheTrilliaceae in the southeastern United States. Harvard Pap. Bot. 

ZoMLEFER, W.B. 1 997.The genera of Melanthiaceae in the southeastern United States. Har- 

ZoMLEFER, W.B. and W.S. Judd. 2002. Resurrection of segregates of the polyphyletic genus 
Z/gac/enus S.I. (Liliales: Melanthiaceae) and resulting new combinations. Novon 12:299- 

ZOMLEFER, W.B.,N.H.WiLUAMs,W.M.WHiTTEN,and W.S. JuDD.2001. Generic circumscription and 




Waterloo, Ontario, Canada N2L 3Q / 
)some numbers are reported for 85 collections of Cerastium (Caryopfiyllace: 
ionh America belonging to 22 taxa. First published reports include those fo 
1 Kearney & Peebles (In = 34) and C texanum Britton (2n = 36). First reports 
.merican origin include C hialynickii Tolm. (2n = f08), C. hrachypodum (En; 
»n (2n = 34), C dichotomum L. (2n = 38), C diifusum Pers. (2n = 72), C/ischeric 
), C.giomeratum ThuiU. (In = 72) and C semidecandrum L (2n = 36). 

los numeros cromosomaticos de colecciones de Cerastium (Caryophyllai 
J continental pertenecientes a 22 taxa. Entre ellos se encuentran los primeros : 
/ar. objectum Kearney & Peebles (In = 34) y C texanum Britton (2n = 36). Prim 

hialynickii Tolm. (2n = 108), C hrachypodum (Eng. e 
L (2n = 38), C. di//usum Pers. (2n = 72), C.fischerianw 
= 72) y C semidecandrum L. (2n ^ 36). 

The purpose of this paper is to put on record previously unpubhshed data on 
chromosome numbers m the species of Cerastium, both native and introduced, 
occurring in contmental North America. 

Mitotic chromosomes were studied in root-tip preparations. Plants were raised 
in the greenhouse, either from seed or from material transplanted from the wild. 
Root-tips were taken from actively growing plants, prefixed for about one hour 
in a saturated aqueous solution of paradichlorobenzene and fixed in 1:3 glacial 
acetic acid and ethanol (absolute). They were hydrolyzed in N hydrochloric acid 
at 60°C for 10—15 minutes and squashed in aceto-carmine. In a few cases flower 
buds were used for counts of meiotic chromosomes. The buds were fixed in 
Carnoy's Fluid (acetic acid Lchloroform lethanol 3). Developing anthers were 
teased out and squashed in aceto-carmine or aceto-orcein. The cultivated plants 
were grown to maturity and voucher specimens taken for comparison with 
those collected m the wild. Voucher specimens are retained in the author's re- 

Ont.: Hudson Bay - Cape 

.: Kananaskis to Coleman re 

Alaska:Taylor Hwy. - Chicken, / 
Alaska: Alaska Pen.- Ugashik, 
Talbot 103-25 

Calif-Alpine Co.- Round Top Mt., 

Colo.: Rocky Mt. Nat. Pk.,Ay/^7334 
Colo.: Mt. Evans, NA7345 

Nfld.:Cape St. George, /V/\5409 

Que.:Gaspe Pen,- Perci 
Wyo.: Park Co. -Snow L 
Wyo.:Beartooth Mts.-E 

ium (Bast.) Guepin 

eCo.- nrTazewell, 

■Cowan -0 


10 E of Swan 

•, NA4549 

Lanark Co. - 


Imonte, NA2} 

Bruce Pen. - 


ermory, NA37 

: Mendocino Co. - 


Tenn.iGatlinburg, iV/\4443 

Cpumilum Cuals 

Ont.: Bruce Pen.- 

N.Y.: New York City -State 


Ohio: Erie Co.- Catawba 1^ 

;., Sandusky, 



Worfon s.n. 

Ont.: Essex Co.- Point Pelee, /V/^4476 



Pa.: Chester Co. -Lees Brie 

ige, NA3770 

Calif.: Marin Co. -Pt. Reyes 


•t Bragg, 

Stebbins 7007 

Calif.: Sonoma Co.- Bodec 

ja Beach, 


)n Beach, 

Chambers s.n. 

Oreg.:Clatsop Co.- Onion 

search collections and partial sets of duplicates have been 
MICH, DAO and VDB. Collection numbers prefixed with ^ 
J.K. Morton and Joan M. Venn collections (Table 1). 

Many of these chromosome counts confirm those from previous workers. How- 
ever, the following appear to be the first published counts for their respective 
taxa: Cerastium nutans var ohjectum - 2n = 34 and C texanum - 2n = 36. That 
for C terrae-novae - In = 108, is the first definitive count for that species though 
Bocher (1977) reported a chromosome number of "lower than 72 (perhaps 68)." 

The following chromosome counts appear to be the first published counts 
for North American material of these alien species: C dichotomum - In = 38, C 
dijjusum - In = 72, C.glomeratum -In = 11 and C stmidzcandrum - In = 36. 

Amongst native species that also occur in other regions, that for C. 
bialynickii - In = 108, is new. Cerastium bialynickii has not previously been 
reported from North America, having been originally described from Siberia, 
where it is reported to have a chromosome number of 2n = 72. It is a very com- 
pact and hispid relative of C. heeringianum with obtuse sepals. 

Also our count for C.fischerianum is the first from this continent. The pres- 
ence of two different chromosome numbers in adjacent populations of this spe- 
cies is unusual. The differences appear to be genuine, for the preparations were 
carefully checked. The explanation may be that the population at sea level, with 
In = 66, was growing in a weedy situation on mine tailings. As C.fischerianum 
frequently intergrades with C heeringianum it is possible that these plants were 
of hybrid origin. 

The count for C hrachypodum (In = 34) is the first for North America, but 
Beaman et al. (1962) reported the same chromosome number in material from 
Central America. 

I am indebted to the following persons for their assistance m supplying seeds 
used m this research: L.C. Bliss, K.L. Chambers, WJ. Cody R. Old, M.J. Oldham, 
DJ. Pmkava, S.S. Talbot, E.K. Winterhalder and the late G.L Stebbms. 

Beaman, J.H., D.C.D. De Jong, and W.P. Stoutamire. 1 962, Chromosome studies in the alpine 

and subalpine floras of Mexico and Guatemala. Amer. J. Bot. 4941 -50. 
Bocher, TW. 1977. Cerastium alplnum and C arcticum, a mature polyploid complex. Bot. 

Notiser 130:305-309. 


Sally WAbOWSKl 2002 Gardening with Praine Plants How to Create Beautiful Native 
Landscapes. Photography by Andy Wasowski USBN 0-8166-3087-Q pbk ) 
outh Suite 290 Mmne- 
cdu 612-627-1932, 612-,cs81/2"xir 

■isityof Mil 
apohs,MN 55401-2^20 (Oidetb w \\ w i 
627-1980 lax) ^.2995 304 np 24Lcolo 

*Alina Freire Fierro Diana Fernandez 

Catalina Quintana 

sidad Central del Ed 
Escuela de Biologia 

srbario QCA 

12-584, Quito, ECUADOR 

s. De las 124 especies utiles. 27 (21,8%) son alimenticias, especialtr 

y espirituales; solamente 4 (3,2%) son ornamen 
i de herramientas, escobas, encofrados, forraje p 

;nera and 553 species regist< 
It QCNE, QCA and the litera 


El Ecuador mcluye 553 especies (Renner et al. 1999) de las 4500 existentes en el 
mundo (Mabberley 1997). Registros de usos de Melastomataceae, junto con otras 
familias de plantas se ban concentrado principal men te a areas o a grupos etnicos 
determinados del pais. Asi, Ceron (1996) registra los usos de plantas del Parque 

Nacional Machalilla (prov, de Manabi), Kvist y Holm-Nielsen (1987), basados 
en sus estudios en la comunidad indigena Cayapa (prov. de Esmeraldas) y en la 
literatura, describen rituales curativos y usos de plantas por comunidades 
indigenas de las tierras bajas. Para la region Andina, Kothari (1993) lista las 
plantas usadas por pobladores de La Esperanza, comunidad mdigena Zuleta 
(prov. de Imbabura), Ceron (1996) reporta los usos de plantas de la Reserva 
Geobotamca Pululahua (prov de Pichincha), RIos (1993) lista las plantas de una 
comunidad mestiza del noroccidente de Pichincha, EUeman (1991) publica los 
usos maderables realizados por la comunidad Saraguro (prov de Loja), y Cueva 
(1997) y Van den Eynden (1997) listan las plantas alimenticias de la provincia 
de Loja. Muchos otros trabajos etnobotanicos han sido realizados en 
comunidades indigenas de la Amazonia ecuatoriana, entre ellos podemos citar 
a Alarcon (1988), Ceron (1996), Ceron y Montalvo (1998), Lescure et al. (1987) y 
Vickers y Plowman (1984). A una mayor escala geografica, Schultes y Raffauf 
(1990) presentan usos etnobotanicos de Melastomataceae y demas familias de 
angiospermas presentes en la Region Amazonica Nororiental. 

La ausencia de publicaciones sobre los usos de la familia a nivel del Ecua- 
dor nos motive a la recopilacion de informacion sobre nombres vernaculares, 
usos actuales y potenciales de especies de Melastomataceae a partir de las 
etiquetas de especimenes depositados en los dos herbarios con mayor numero 
de colecciones del pais (QCNE y QCA). 

La informacion sobre usos de plantas se la obtuvo a partir de los especimenes con 
informacion etnobotanica depositados en el Herbario Nacional del Ecuador 
(QCNE) y herbario de la Pontificia Umversidad Catolica del Ecuador (QCA), 
ambos en Quito. De cada especimen se recopilo el nombre de la especie, el colector 

determinacion de los especimenes botanicos depositados en am bos herbarios fue 
realizada principalmente por Elvira Cotton (actualmente en el Herbario AAU, 
Aarhus, Dinamarca), especialista en Melastomataceae ecuatorianas quien residio 
en Ecuador por varios anos. Informacion publicada en la literatura, y aquella 
producto de nuestra experiencia de campo tambien fueron incluidos en el presente 
trabajo. Datos sobre el numero total de especies por genero fueron tomados de 
Renner (1993) y Mabberley (1997), e informacion sobre el numero de especies por 
genero y su distribucion en el Ecuador se obtuvo de Wurdack (1980), Renner et al, 
(1999) y Freire-Fierro (en prensa). El material examinado esta listado en el Anexo, 
en el cual se incluyen a los autores de los nombres cientificos. 
Aciotis D. Don 

Este genero herbaceo incluye 13 e 
especies que se distribuyen principa 

diUera Occidental y en las faldas onentales de la Cordillera Oriental. Aciotis 
rubricaulis, una especie distnbuida a ambos lados de la cordillera es utilizada 
para curar picaduras de culebras. Similar uso se registra para Aciotis ornata, 

noroccidente. Ceron (1994) registra el uso de Aciotis leoyara (probablemente 
refiriendose a A. levyana Cong., un smonimo de A. rubricaulis [Freire-Fierro 
(en prensa)]) como antiofidica. 
Arthrostemma Pavon ex D. Don 

Este genero arbustivo presenta siete especies, y solamente Arthrostemma ciliatum 
esta registrada en el Ecuador, en donde se distribuye en la region costera, andina y 
amazonica. Esta especie es utilizada para curar la diarrea en ninos y para curar 
dolores de garganta. Los tallos suculentos son un tanto acidicos y son generalmente 
utilizados para calmar la sed. Esta especie tambien podria ser utilizada como orna- 
mental debido a su habito trepador y f lores de un vistoso color rosado. 
Axinaea Ruiz y Pavon 

Este genero arboreo o arbustivo consta de 30 especies, 10 de las cuales ocurren 
en los Andes del Ecuador De estas, Axinaea affinis, A. macrophylla, A. quitensis, 
y A. sclerophylla son utilizadas principalmente para la construccion y como 
combustible en calidad de lefia y carbon. Un potencial uso del genero seria como 
ornamental ya que las flores son bastante vistosas. 

Dos especies de Axinaea son consideradas maderables, A. macrophylla por 
Ceron (1994) y A. sclerophylla (como A. schhrophylla) por Elleman (1991). 
Bellucia Neck, ex Raf . 

Este genero arboreo consta de siete especies, y solamente Bellucia pentamera 
esta registrada en el Ecuador, en donde se distribuye en la Costa, Andes y 
Amazonia. La madera de esta especie es utilizada para la construccion. Los 
frutos, que nacen directamente de los troncos, pueden alcanzar hasta 10 cm de 
diametro, son comestibles y de sabor ligeramente dulce. 

El fruto maduro de Bellucia pentamera (como B. weherhaueri Cogn.) es 
comestible para los Kichwas (como Quichuas) de la Amazonia Ecuatoriana 
(Alarcon 1988, Lescure et al. 1987) y por colonos mestizos de la comunidad 
Ahuano (provincia del Napo) (Rios y Caballero 1997). El nombre Bellucia 
axinthera (probablemente refiriendose a B. axinanthera Triana, un sinonimo 
de B. pentamera) es listado como alimenticio por Ceron (1994). 
Blakea P Browne 

Este genero arbustivo incluye 100 especies, 31 de las cuales ocurren en el Ecuador, 
en donde se distribuyen en la Costa, Region Andina y Amazonia. Blakea hracteata, 
B. glandulosa, B. involvens, B. punctulata, B. rosea, y B. suhconnata son utilizadas 
para tratar heridas, picaduras de culebras, y prevenir ampollas. Tambien son 
utilizadas para curar el "enduendado" [hechizo] y para la construccion de casas. 

Especies de este genero tambien tienen gran potencial ornamental, especialmente 
para reforestar parques y avenidas debido a que sus f lores son bastante Uamativas, 
puesto que existe un contraste entre los petalos, generalmente rosados, con los 
estambres y estaminoides que son de un amarillo dorado. 

El especnnen Viewers y Plowman 255 (QCA) listado como ''Blakea aff. 
ciliata Mgf. or B. rosea" por Vickers y Plowman (1984), y como B. rosea por 
Lescure et al. (1987), es conocido por los Siona-Secoyas como "Yayurua" y es 
utilizado para tratar quemaduras. B. sawadae es utilizada por los Cofanes como 
medicinal (Ceron 1994). B. portentosa Wurdack es conocida por los Huaoranis 
como "Namuntaqui" y tiene frutos que son consumidos por aves pequefias 
(Ceron y Montalvo 1998). 
Brachyotum (A. DC.) Triana 

Brachyotum es un genero arbustivo de 58 especies, 21 de las cuales ocurren en el 
Ecuador, en donde se distribuyen unicamente en los Andes. Cinco especies, 
Brachyotum alpinum, B. azuayense, B. Icdifolium, B. rostratum y B. trichocalyx 
son utilizadas para curar el catarro en los polios, tefiir, construir, asi como para 
leiia y escobas. Este genero puede ser tambien utilizado como cercas vivas. 
Algunas especies de Brachyotum tienen potencial como ornamental ya que sus 
flores presentan generalmente calices de un color rojo vivo y corolas morado 
oscuras (e.g. Brachyotum andreanum Cogn.) o amarillas (e.g. B. ledijolium); sin 
embargo, las plantas tendrian que ser sembradas especialmente en lugares frios 
ya que el genero es tipicamente de paramo. 

Brachyotum ledijolium es utilizada medicmalmente por los Kichwas 
andinos (Ceron 1994) y como combustible y para escobas en la zona del 
Pululahua (Ceron 1996). 
Clidemia D. Don 

Este genero arbustivo incluye 120 species, 38 de las cuales ocurren en el Ecua- 
dor, en donde se distribuyen tanto en la Costa, Region Andina como Amazonia. 
Las siguientes 14 especies son utilizadas: Clidemia acostae, C. allardii, C. 
C. hirta, C. ostrina, C. purpurea, C. scptuplinervia, C. sprucei y C variifolia. Estas 
especies son utilizadas para tratar heridas, ulceras, diarrea con sangre, asi como 
para cicatrizar, calmar el mareo, y bajar la fiebre. Ademas, una de estas especies 
es utilizada por los shamanes para "asustar a los espiritus", especialmente los 
"malos" espiritus lentidades sobrenaturales que pueden hacer dafioj. Debido a 
que las bayas son agri-dulces, estas son utilizadas para calmar la sed. Muchas 
especies de CI idemia presentan form icarios en la base de las hojas o en los tallos 

sus nidos. Los formicarios tienen un sabor acido debido a esta situacion y son 
consumidos como alimento por algunos grupos etnicos de la Amazonia. 

Clidemia dentata, conocida como "Paitzi", es utilizada por los Kichwas de 

la Amazonia ecuatoriana para bajar la fiebre (como Clidemia sp, Alarcon 1988, 
Lescure et al. 1987). Los frutos de C. dimorphica y C. sprucei son comestibles 
para los indigenas Huaoranis (Lescure et al. 1987), y es tambien conocida como 
medicinal por los Cofanes (Ceron 1994). Los indigenas Kichwas y mestizos de 
la comunidad Ahuano (provincia del Napo), tambien consumen los frutos de 
una especie de Clidemia no determinada (Rios y Caballero 1997). Clidemia 
heterophylla es listada como medicmal por Ceron (1994). 
Conostegia D. Don 

Conostegia es un genero arbustivo de 43 especies, 16 de las cuales ocurren en 
Ecuador, en donde se distribuyen principalmente en la Costa, aunque tambien 
se han registrado especies para la Region Andina y la Amazonia. Conostegia 
attenuata, C centronioides y C. dentata son utilizadas para curar enfermedades 
[no especificadas en las etiquetasj. 
Diplarpea Triana 

Este genero herbaceo o arbustivo tiene solamente una especie, la cual esta 
presente en el Ecuador. Diplarpea paleacea ocurre en los flancos de la Cordi- 
llera norte andina y es utilizada para tratar dolores de estomago. 
Graffenrieda DC 

Este genero arbustivo o arboreo consta de 44 species, 11 de las cuales existen en 
Ecuador, y estan distribuidas principalmente en la region andina. Dos especies 
del genero, G. af f . intermedia y Graff erieda sp. son utilizadas en la construccion 
y como lefia por los Huaoranis de Quehueiri-Ono (provincia del Napo) (Ceron 
y Montalvo 1998). 
Henriettella Naudin 

Este genero arbustivo o arboreo incluye 51 species, siete de las cuales se 
encuentran en Ecuador, tanto en las tierras bajas como en los flancos occidental 
y oriental de los Andes. De acuerdo a Rios y Caballero (1997), el fruto de una 
especie no identif icada es conocida como comestible por los pobladores Kichwas 
y mestizos de la comunidad de Ahuano (provincia del Napo). 
Leandra Raddi 

Este genero arbustivo incluye 175 especies, 19 de las cuales ocurren en el Ecua- 
dor, en donde se distribuyen principalmente en la Amazonia, aunque tambien 
hay registros de la Region Andina y de la Costa. Leandra caquetana, L. 

caquetensis, L. granatensis, L. secui 

Ida y L suhseriata, son utilizadas como cc 

mestibles por sus frutos, y para cura 

r ulceraciones de la piel. Los pequenos frutc 

carnosos son comidos por las aves. 

Loreya DC. 

Loreya es un genero arbustivo que o 

omprende 14 especies. De las cuatro especie 

ecuatorianas, una se distribuye en : 

la Costa y las tres restantes en la Amazon!; 

De acuerdo a Lescure et al. (1987), los 

; frutos de Loreya spruceana (bajo L. eollatat 

Wurdack) son comes 
puercos salvajes (~bajc 

tibles. Estos frutos 

tambien son comic 

Maieta Aubl. 

Este genero arbustivo : 

incluye unii 

;tas, Mai 



dolores estomacales, y de cabeza, mareos, y para cicatnzar heridas. Esta especie 
tambien tiene frutos carnosos que son comidos por las aves. 
Meriania Sw. 

Este genero arbustivo y arboreo consta de 74 especies, 27 de las cuales ocurren 
en el Ecuador Una sola especie esta registrada para la Costa y las restantes se 
distribuyen en los Andes. Los troncos de Meriania tomentosa son utilizados para 
la elaboracion de carbon (Ceron 1996). Este genero tambien tendria gran 
potencial para forestacion de parques y avenidas por las vistosas f lores. Aunque 
este ultimo uso no se da en el Ecuador, en Colombia especies del genero son 
utilizadas como arboles ornamentales de avenidas (D. Neill, com. pers. 2001). 

La madera del "Quihle o Quishle bianco," una especie indeterminada de 
Meriania, es utilizada en la construccion por los Saraguros (Elleman 1991). 
Miconia Ruiz y Pav 

Miconia es el genero mas grande de la familia, incluye aproximadamente 1000 
especies, 240 de las cuales estan registradas para el Ecuador La mayoria de 
especies son andinas, aunque tambien ocurren en la Costa y en la Amazonia. 
La alta diversidad de especies es acompafiada por la diversidad de usos de 
especies de Miconia en el Ecuador. En los herbarios revisados se tienen 
registradas como medicinales a las siguientes especies: Miconia affinis, M. 
harhinervis,M- hracteolataM- huhalina,M-crocea,M.erioclada,M-goniostigma, 
M. gracilis, M. lugonis, M. nervosa, M. pahacea, M. papulosa, M. procumbens, M. 
theaezans, M. trinervia, M. triplinervis, M. wnulosa, y M. zuhenetana, las que 
son utilizadas como vermifugo, para estimular las dilataciones durante el parto, 
curar dolores de garganta y del cuello, tratar la tuberculosis, dolores de muelas, 
infecciones bucales, micosis, sarnas y para curar diarrea de nifios recien nacidos, 
e inclusive pueden ser utilizadas como antidoto de picaduras de la hormiga 
"Conga" (Paraponera spp.) (ver Ceron 1994, 1996, Ceron y Montalvo 1998; 
Schultes y Raffauf 1990). Miconia buhalina (como Miconia sp.) es utilizada por 
los Kichwas de la amazonia para tratar pequenos abcesos (Alarcon 1988; Lescure 
et al. 1987). 

La madera de Miconia aeruginosa, M. astroplacama, M. aureoides, M. 
M. decurrens, M.flori bunda, M. lutescens, M. matthaei, M. napoana, M. obscura, 
M.ochracea,M. pahacea, M. papulosa, M-pustulata,M.salicifolia,M.splendens, 
M. subspicata, M. theaezans, y M. tinijolia, es utilizada principalmente para 
construccion, para fabricacibn de herramientas, produccion de carbon y como 

combustible (Ver Ceron 1994, 1996; Ceron y Montalvo 1998; Cueva 1997; 
EUeman 1991; Van den Eynden 1997; Lescure et al. 1987; Schultes y Raffauf 1990; 
Vickers y Plowman 1984), De acuerdo a Ceron y Montalvo (1998), otra especie 
utilizada en la construccion y lena es Miconia ternijolia Triana, sin embargo, 
este nombre no esta registrado en el Catalogo de Plantas Vasculares del Ecua- 
dor (Renner et al. 1999) ni en la base de datos TROPICOS (Missouri Botanical 
Garden 2002). 

Si bien los frutos bacaceos de Miconia alcanzan hasta 1 cm de diametro, 
algunos son dulces y comestibles, tal es el caso por ejemplo de Miconia 
asclepiadea, M. aspergillaris,M- crocea, M. latijolia, M. ledifolia, M. lutescem, M. 
cf. sphndens y M. theaezans (Ver Ceron 1994, 1996; Ceron y Montalvo 1998; 
Cueva 1997; Van den Eynden 1997). 

Los frutos de Miconia huhalina, M. calvescens, M. cazaletii, M. centrodesma, 
M. cercophora, M. decurrens, M. nervosa y M. oraria son comidos por aves (Ver 
Cer6n y Montalvo 1998). 

Algunas especies tendrian tambien potencial ornamental, por ejemplo, 
Miconia crocea tiene paniculas de cerca de 20 cm de longitud con abundantes 
flores con petalos blancos y estambres amarillos. Otra especie con potencial 
ornamental podria ser Miconia salicifolia, la cual se distribuye en la Region 
Andina y presenta arbustos con apariencia de candelabros debido a la densa y 
erecta disposicion de los tallos y a la posicion adpresa de las hojas, las cuales 

Los indigenas Kichwas de la Amazonia ecuatoriana utilizan una especie 
no determinada de Miconia conocida como "Chiquitapayansu" para tratar 
granos infectados (Alarcon 1988). La madera de M. astroplacama es utilizada 
por los Siona-Secoyas para pilares en construccion de casas (Lescure et al. 1987; 
Vickers y Plowman 1984). Los frutos de otra especie no identificada de Miconia, 
y conocida por los Kichwas como "Payanzo", son comestibles (Lescure et al. 1987). 
Los frutos de M. hdifolia, M. lutescens y dos especies no determinadas son co- 
mestibles en la provincia de Loja (Cueva 1997; Van den Eynden 1997). Ceron 
(1994) registra el uso de Miconia crocea en la medicina y M. palacea, M. pustulata 
y M. theaezans en la construccibn. EUeman (1991) lista como maderables para 
los Saraguros a M. caelata, M. corymhijormis, M. ohscura, M. cf. tinifolia y dos 
especies no identificadas. 

1 (DC.) Naudm 

Este genero arbustivo mcluye 45 especies, y solamente tres estan registradas 
para el Ecuador, en donde se distribuyen en la region andina. Monochaetum 
hartwegianum es utilizado como ornamental y M. lineatum para lena. Esta 
ultima especie es conocida como maderable por los Saraguros (EUeman (1991). 
Monolena Triana 
Monolena es un genero herbaceo de 10 especies, y unicamente M. primulaejlora 

esta registrada tanto en la Costa y Region Andma como en la Amazonia 
ecuatoriana. Esta especie es utilizada para tratar la gripe, dolores de estomago, 
parasites intestinales, conjuntivitis, siendo ademas usada como antiofidica. 
Debido al sabor acido de los peciolos suculentos, e.sta cspecie tambien es utilizada 
para calmar la sed. Monolcna primulacjlora podria ser explotada como planta 
de interior debido a que las hojas acaulescentes tienen peciolos rojizos y 
suculentos, laminas variegadas y flores bastantc llamativas. 

Kvist y Holm^Nielsen (1987) mencionan que esta especie es usada por los 
Coaiqueres como vermifuga y que es fuertemente hemetica, y Ceron (1994) la 
lista como laxante. Schultes y Raffauf (1990) registran el uso de la misma especie 
para tratamientos de mordeduras de serpientes en la provincia de Pastaza. 
Mouriri Aubl. 

Este genero arboreo consta de 81 especies, 10 de las cuales se han registrado para 
el Ecuador, en donde ocurren principalmente en la Amazonia. De las tres 
especies registradas como utiles, M. oligantha es utilizada como anticonceptivo 
y esterilizante; y los frutos de Mouriri grandiflora y M. nigra son comestibles. 
Su uso como comestible esta registrado tambien por Ceron (1994). 
Ossaea A.P. DC. 
Ossaea es un genero arb 
la Costa y Amazonia e 

especies iitiles: Ossaea 
0. rohusta. Los herbai 
utilizadas para bajar k 
como comestibles. 

Una especie identi 
por los Kichwas de la 
utilizada para tratar gri 

hracteata, 0. laxivenula, 0. micrantha, 0. quadrisuka, y 
:ios estudiados registran que estas especies han sido 

ficada como Ossaea sp. Imuestra Alarcon 34] y conocida 
Amazonia ecuatoriana como "Hagapania/Payanzo" es 
mos infectados y sarna (Alarcon 1988; Lescure et al. 1987). 
imen fue mas tarde identificado como Miconia affinis. 

Tessmanianthus Mark^ 
Este genero arboreo ne 
registradas para la An 
herbario de este genei 

^otropical incluye siete especies, de las cuales dos estan 
lazonia ecuatoriana. Aunque no se estudio material de 
■0, Ceron y Montalvo (1998) mdican que el fuste de I 

heterostemon Markgr. es utilizado por los Huaoranis de Quehueri-Ono para 
construccion y para lena. 
Tibouchina Aubl 

Este genero arbustivo incluye 240 especies. Solamente 14 especies est£ 
registradas para el Ecuador, las cuales se distribuyen principalmente en L 
Andes, aunque tambien existen especies en la Costa y Amazonia. Siete especit 
Tibouchina gleasoniana, I.grossa, T. laxa, T. lepidota, T. longifolia, T. mollis,y 
oroensis son utilizadas para tratar cataratas, extraer sarcoma de los ojos, trat; 
dolor de muelas, para forraje de los animales, lena o uso ornamental. 


Tihouchina lepidota es muy comun en la region andina nor-oriental, y 
podria ser utilizada no solo como combustible, sino tambien como ornamental. 
Las inflorescencias estan compuestas de grandes flores con colores blancos, 
rosados y fucsias y con estambres amarillos vistosos. Con la excepcion de 
Ti houchina oroensis, las otras especies de Ti bouchina ornamentales observadas 
en el Ecuador son importadas del Brasil (D. Neill, com. pers. 2001). 

El uso como combustible de Tihouchina mollis esta registrado en Ceron 
Tococa Aubl. 

Tococa comprende cerca de 50 especies de plantas arbustivas usualmente 
mirmecofilas. Ecuador posee 12 especies distribuidas principalmente en las 
tierras bajas de la Amazonia, existiendo menos en la Region Andina y la Costa. 
Tococa guianensis es utilizada para tratar los granos infectados de la piel. Los 
especimenes examinados tambien registran que los frutos de T. spadiciflora y 
de T. symphandra son comidos por las aves. Aunque las etiquetas de los 
especimenes herborizados no lo registren, es posible que las pequenas y vistosas 
bayas de otras especies de Tococa scan tambien comidas por aves. 

Tococa guianensis (-erroneamente escrita ''Cococa") es utilizada por los 
Kich was de la Amazonia ecuatoriana para curar granos infectados y es conocida 
como "Payanshu" (Alarcon 1988), o como "Jatunpayanshu" (Lescure et al. 1987). 
Los frutos maduros de esta misma especie, bajo el sinonimo Tococa chuivensis 
Wurdack van ecuadorensis Wurdack, son comidos por personas y por aves 
(Ceron y Montalvo 1998). 
Triolena Naudin 

Triolena es un genero herbaceo de 22 especies, 11 de las cuales se encuentran en 
Ecuador, especialmente en las tierras bajas tanto de la Costa como de la 
Amazonia. Triolena hirsuta, T ohliqua, T. pluvialis, T. pedemontana, y T. 
pustulata son utilizadas en Ecuador como medicinales. Estas especies son 

subsiguientes hinchazones, y para tratar tuberculosis y tos. 

Los Cayapas y Coaiqueres utilizan Triolena para tratar mordeduras de 
culebras, y los Siona-Secoyas usan T. pluvialis para tratar dolores de muelas 
(Kvist y Holm-Nielsen 1987; Vickers y Plowman 1984). Similar uso es dado para 
esta especie por los Kichwas de la Amazonia ecuatoriana (Alarcon 1988; Lescure 
et al. 1987) y por los Cofanes (Schultes y Raffauf 1990). Ceron (1994) tambien 
registra el uso medicinal de esta especie. 



El presente trabajo se baso primordialmen 
de las etiquetas de dos herbarios ecuato 
generalizada de que Melastomataceae n 

rianos. Contrariai 
es una familia ( 

mente a la opinion 

nportante (e.g. Schultes y Raffauf 1990), al menos una de cada cinco especies 

je ocurre en Ecuador tiene alguna utilidad. 

Mas de la mitad de los generos registrados para el Ecuador son utiles, sin 

nbargo, a pesar de que algunos generos estan presentes en el pais y de ser 
o utiles (e.g. Adelohotrys DC. y Centronia D. Don 
) registran uso alguno en Ecuador. 

medicinalmente, en segundo lugar esta el uso para lena, carbon, alimentos de 
animales, tmciones, etc. [Otros]; una de cada cuatro especies utiles de la familia 
es usada en la construccion, menos del 22% de especies es utilizado en la 
alimentacion, muy pocas especies son utilizadas en rituales magico religiosos y 
solamente cuatro de las 124 especies es utilizada como ornamental. Segun 
Schultes y Raffauf (1990), sin embargo, la unica importancia economica que la 
familia tiene es por las especies ornamentales. Probablemente esta discrepancia 
se deba a que si bien en Ecuador Melastomataceae no es muy utilizada a nivel 
ornamental, este uso es comun en otros paises. 

Debido a la diversidad de especies, Miconia es el genero que mayor numero 
de especies utiles presenta, principalmente en la construccion, medicina y usos 
varios. Clidemia es el segundo genero mas importante (14 especies utiles) y la 
mayoria de sus especies es utilizada en la medicina, aunque C crenulata es 
tambien utilizada en ritos magico religiosos. En tercer lugar estan Blakea y 

Tibouchma, gener 

OS con 7 esp 

ecies utiles cada uno. Los otros generos tienen de 

una a cmco espec 

;ies utiles, i 

\ nivel de especies, Miconia theaezans, Blakea 

suhconnata, Micon 

'ia hracteol 

ata, Miconia hubalina, Miconia crocea, y Ossaea 

micrantha registv'c 

iron el mayc 

)r numero de usos, tanto en la alimentacion, como 

m medicina y con 


A nivel de uso 


ia y Monokna son los mas utilizados para calmar 

la sed debido a sus 


ulentos y de sabor acido; y Bellucia, Henriettella, 


son predom 

Linantemente usados como alimenticios debido a 

que sus frutos sor 

1 relativam^ 

mte grandes y bastante dulces. Las especies de 

[, Bellucia, Grafjenrieda, y Tessmanianthus son las mas utilizadas en la 
especialmente debido al habito arboreo de las mismas. Mas del 
60% de las especies de Aciotis, Arthwstema, Blakea, Clidemia, Conostegia, 
Diplarpea, Maieta, Monohna, Ossaea, y Triohna, son utilizadas 
medicmalmente. A nivel magico religioso, Aciotis es el mas utilizado y menos 
del 50% de las especies de Blakea, Clidemia, Conostegia, Miconia, y Ossaea es 
tambien usado para este fin; ninguno de los otros generos registrados para Ec- 
uador es utilizado para el mismo proposito. De los 24 generos usados en Ecua- 
dor, unicamente Monochaetum (M. hartwegianum) y Tihouchina (T.gkasoniana 
y T. grossa) son en la actualidad utilizados como ornamentales. Mas del 55% de 
las especies de los generos arboreos y arbustivos Axinaea, Brachyotum, 
Grafjenrieda, Loreya, Meriania, y Tihouchina es utilizado primordialmente 
para carbon, lefia o por la fauna local como alimento. 

De las 124 especies utiles, 13 de ellas (cerca del 10%) fueron citadas en la 
literatura pero no estaban representadas en los dos herbarios (Ceron 1994, 1996, 
Ceron y Montalvo 1998; Elleman 1991; Rios y Caballero 1997), y de las HI especies 
restantes, solamente 36 (32%) fueron citadas en estudios etnobotanicos 
anteriores (Alarcon 1988; Ceron 1994, 1996; Ceron y Montalvo 1998; Cueva 1997; 
Elleman 1991; Freire-Fierro en prensa; Kvist y Holm-Nielsen 1987; Lescure et al. 
1987; Rios y Caballero 1997; Schultes y Raffauf 1990; Van den Eynden 1997; 
Vickers y Plowman 1984). Esta compilacion da una pauta sobre la necesidad de 
publicar informaci6n que de otra manera permanece restringida unicamente a 
los investigadores locales, quienes podrian darla a conocer a la comunidad 
mternacional por medio de publicaciones similares en otras familias de plantas. 

La informacion sistematica sobre usos de Melastomaceae podria servir de 
base por ejemplo, para la busqueda de principios activos en las especies listadas 
como medicinales; o para la seleccion de especies para la construccion y ornato 
que mejor podrian crecer en determinada region geografica; o para estudios 
ecologicos de las especies utiles endemicas, especialmente de aquellas 
amenazadas de extincion (Cotton 2000); para futuros planes de manejo 
sustentable y conservacion. Adicionalmente, estudios etnobotanicos y de campo, 
enfatizados en la familia, podrian verificar los usos de Melastomataceae 
registrados aqui y en la literatura. 

Material de Melastomataceae examinado en los herbarios QCA y QCNE. Usos: 
A = Alimenticio; C = Construcccion; M = Medicinal; MR = Magico-Religioso, O = 
ornamental; R = Otros. Distribucion Qorgensen y Leon 1999, Renner et al. 1999, 
Freire-Fierro en prensa): AZU = Azuay BOL = Bolivar, CAN = Canar, CAR = 
Carchi, CHI = Chimborazo, COT = Cotopaxi, ORO - El Oro, ESM = Esmeraldas, 
GUA = Guayas, 1MB = Imbabura, LOJ = Loja, RIO = Los Rios, MAN = Manabi, 
MOR = Morona Santiago, NAP = Napo, PAS = Pastaza, PIC = Pichincha, SUC = 


Payanzo (K), 

Sacha guayaba (K), (QCA,QCNE), 


3n/20(QCA), Alarcon 

■■6/54 (QCA), Ceron ( 


r 40049 (QCA) 

'od 4161 3 (QCA) 



1 66933 (QCA), 


) 75440 (QCAj 


ai 6883 (QCA, 


iQnez 75 (QCA, 


majerol 27 {QC 


1 66862 (QCA) 


1 66575 (QCA) 


) 70/2 (QCA) 


/ 060 (QCA), 


I 85 (QCA) 


, 84 (QCA) 


106/ (QCNE), 


3077 (QCA) 

Rios 434 (QCA) 


Kvist 40307 (QCA) 


KvistyAsanza 40896 


Kvist 405 W {QCA), 

Rios 357 (QCA) 


Asdall 8228 {QCA), 


), Kohn 11 62 {QCA, 




13805 (QCA, 



14090 {QCA, QCm, 

Palaaos 10254 {QCA, 

QCm, Ponce 226 

{QCA).Van Asdall 

82-28 {QCA, QCm 

centronioides COT, ESM, RIO, 


Kvist, 40528 QCA 


Rios 496 (QCA) 


Titemenca (H) 


Chanchak chichi (A), 

Balslev 301 6 iQCA). 

Egimo (H), Payas (K) , 


Putus panga (K), 

Lewis etal.l 3899 iQCA, 



(QCA,QCNE),/?/05 477 




oi fr/feen 59/04 (QCNE), 

Mordnetol.l 5 {QCA. 


Ayamohue (H), 

Alarcon 35a (QCA), 


3, AulestiayNenquerei 




Tiuyu iQCA,QCm,Hurtado 



Montalvo (1 998), 

Cerdn 7850 (QCA,QCNE) 






Alvarez etal.Tl 98 (QCA, Cerdn {' 

QCm. Ayala 2 iQCA), 1996) 

Cerdn etal. 932 (QCA, 


10436 {QCA. QCm. 
Cerdn etol. 10718 (QCA, 
QCNE), Cerdn era/. 10841 
(QCA, QCNE), Cerdn era/. 
/0997 (QCA, QCNE), 

Irvine /27 (QCNE), 

lombrevernaculo Especi'menes 


Beck / 777 (QCA) 


d 41014 {QCM 
d 48099 {QCA), 
1 02 {QCA) 

Elleman 66671 {QCm Ellemc 

4iconia nutans Donn. Sm. 



Cuamacds & Gudino 
245 (QCA,QCNE) 
Ceronetal. 10867 
(QCA, QCN E),Cerdn 
etal. // 069 (QCA, 



Barfod 41495 {QCA, 



/2236 (QCA, QCNE), 




fino (S) 

r/pazJi (QCA, QCNE) 


f 40489 (QCA,QCNE) 

7(QCNE) Ellem 

CAR,COT,ESM, x x Apanguira, Aurutullo Aulestia &Aulestia 1321 

LOJ,MOR,NAP, (K),Auru-tulla,Corazon (QCA,QCNEMu/est/o 

PAS.PICJUN, pangajeenshiimujcu, ef a/. / 456 (QCA.QCNE), 

ZAM Lyofasufsapa'cco, Barfod 41582 (QCA), 

Lyunk (U), Nimon Ceron 186 (QCA), 

onabo, Quenca Kohn 1 156 (QCA, QCNE), 

vgra (DC.) 
-)ligantha P 


Nombre vernaculo 

19 {QC\QCm. Nei 
7728 (QCA), 0llgaari 
57359 (QCA), Palacic 
10291 (QCA) 


41013 {QCA, 



Kvist 40364 (QCA) 


dez 111 (QCA), 


162 (QCA) 


9, 390 (QCA) 

CAN, CAR, sue 

/ (QCA, QCNE) 
lore 628 (QCA) 



Hallilumba,Payanshu Alarcon 27 (QCA), 
{K),Jatunpayanshu (K) Blanc % ?07 (QCA) 

Barfod 48032 (QCA; 

Sucumbios, TUN = Tungurahui\, ZAM = Zamora Chmchipe. A = Achuar, F = 
Cofan, H = Huaorani, I = Siona-Secoya, K = Kichwa, O = Colorado, Q = Coaiquer, 
S = Saraguro, U = Shuar, Y = Cayapa. * = Especie citada bajo un sinonimo, mal 
escrita o no identificada. 

Agradecemos a todas aquellas comunidades que desinteresadamente 
compartieron sus conocimientos con los colectores; a todos los colectores 
listados en el apendice quienes registraron los usos de las plantas en sus etiquetas, 
especialmente a A. Barfod, C.E. Ceron, L. Ellemann, J. Jaramillo, LP. Kvist y M. 
Rios; al personal de los herbarios QCNE y QCA, quienes nos permitieron revisar 
las colecciones de Melastomataceae, especialmente a Mercedes Asanza, David 
Neill, Katya Romoleroux y Renato Valencia. David Neill y Rosa Ortiz-Gentry 
(Missouri Botanical Garden), Kevin Janni y un revisor anonimo, contribuyeron 
mucho para el mejoramiento del articulo gracias a sus comentarios en versiones 
anteriores. La primera autora agradece al Plutarco Naranjo de la Universidad 
Andina Simon Bolivar, por la oportunidad de presentar este trabajo en las II 
Jornadas Ecuatorianas de Etnomedicina realizadas en 1999, a FUNDACYT y al 
Missouri Botanical Garden por todo el apoyo logistico, y a Joseph C. Welling por 
toda su ayuda durante la recopilacion de la informacion. 


CfrOn,C.E. 1994. Manual de botanica ecuatoriana:Sistematica y metodos deestudio. Uni- 
versidad Central del Ecuador, Quito. 
ChRON, C.E. 1996. Plantas utiles de la Reserva Geobotanica del Pululahua, Provincia de 

Ceron, C.E. y C.G. Montalvo. 1 998. Etnobotanica de los Huaorani de Quehueiri-Ono, Napo, 
Ecuador. Abya-Yala, Quito. 

eds.LibroRojode las plantas endemicas del Ecuador 2000. Publicaciones del Herbario 
QCA, Pontificia Universidad Catolica del Ecuador, Quito. Pp. 230-250. 

CurvA, E. 1 997. Arboles y arbustos productores de frutos comestibles en la provincia de 
Loja. En: Ri'os, M. y H.B. Pedersen, eds. Uso y manejo de recursos vegetales. Memorias 
del II Simposio Ecuatoriano de Etnobotanica y Botanica Econ6mica.Ed.Abya-Yala,Quito. 

Elleman, L 1 991 . El uso de la madera del bosque montano por los Saraguro. En: Rios, (VI. y H. 
Borgtoft Pedersen (eds.), Las Plantas y el hombre. Herbario QCA y Abya-Yala, Quito. 

Freire-Fierro,A. En prensa. Monograph of Aciotis (Melastomataceae). Monog.5yst.Bot. 62. 

J0RGENSEN, PM. y S. LeOn-Yanez (eds.). 1999. Catalogue of the vascular plants of Ecuador, 

KoTHARi, B. 1993.Nucanchic Panpa Janpicuna: plantas medicinales del campo. Ed.Aby 

Yala, Quito. 
KvisT, LP.y LB. Holm-Nielsen. 1 987. Ethnobotanical aspects of lowland Ecuador. Opera B( 


inventario critico de los datos disponibles en Quito. ORSTOM-P.U.CE.-I.N.C.R.A 

Programa Nacional de Regionalizacion Agraria PRONAREG, Quito. 
Mabberlly, D.J. 1 997.The plant book: a portable dictionary of the vascular plants. 2nd e 

Cambridge University Press. 
Renner,S.5. 1 993.Phylogeny and classification of the Melastomataceae and Memecylacec 

Nord. J. Bot. 13:5 19-540. 

Gard. 75:561 -585. 
Rios, M. 1 993. Plantas utiles en el noroccidente de la provincia de Pichincha. Hombre 

Ambiente 26:1-185. 
Rfos, M. y J. Caballero. 1997. Las plantas en la alimentacion de la comunidad Ahuar 

Amazonia ecuatoriana. En: Rios, M. y H.B. Pedersen, eds. Uso y manejo de recurs 

vegetales. Memorias del II Simposio Ecuatoriano de Etnobotanica y Botani 

Economica. Ed. Abya-Yala, Quito. 
ScHULTES, R.E.y R.R Raffaue. 1 990.The healing forest: medicinal and toxic plants of the Nort 

west Amazonia. Dioscorides Press, Portland, QR. 
Van den Eynden, V. 1 997. Plantas comestibles en la provincia de Loja. En: Rios, M. y H 

Pedersen, eds. Uso y manejo de recursos vegetales. Memorias del II Simpos 

Ecuatoriano de Etnobotanica y Botanica Economica. Ed. Abya-Yala, Quito. 
ViCKERS,W.T.yT. Plowman. 1984. Useful plants of the Siona and Secoya Indians of easte 

Ecuador. Fieldiana, Bot. 1 5:1-63. 
WuRDACK, J. 1980. Melastomataceae. En: Marling, G. y B. Sparre, eds. Flora of Ecuador 1 









PNG National Forest Service 

P.O.Box 5055 



mops ledermannii (Thymelaeacea 
taxa are currently being confused 



involved in the eaglewood trad 
the true eaglewood IS readily ( 



noli kohmGyrinopsledcrmann 



hymelaeaceae) long 
3ve long Papua New 



In recent years, a lucrative trade has emerged in the Hunstein subdistrict of 
East Sepik Province, involving a natural product commonly known as 'eaglewood' 
(Fig. 1). The newly established market is based on an arborescent species which 
accumulates aromatic oils in older wood. Demand for the fragrant wood can be 
likened to that for sandalwood, Santalum spp. (Santalaceae). However unlike 
sandalwood, the eaglewood essence appears to be a contingent response to in- 
jury or trauma, rather than a normal outcome of maturation. In its most desir- 
able form, eaglewood becomes blackened by concentrated deposits of aromatic 
compounds, attracting prices of K800-l,000 per kg for the highest grades. Fol- 
lowing separation of the oils, the fragrant extracts are currently used in the 
production of expensive incense and perfumes, for eventual purchase by Asian 

Due to its recent appearance in commerce and unusually high value, 
eaglewood has become the focus of intense interest. Although the product is 
known primarily from lowland and low montane forest, there has been con- 
siderable uncertainty over the identity and number of species involved in the 
current trade. In an attempt to resolve the taxonomic issues, the eaglewood 
source areas near Ambunti township were visited by the authors in August 2001, 

as part of a general floristic study of the upper Sepik. The following account 
presents the results of these investigations.^ 

During the 1989 National Geographic Society expedition to Mt. Hunstem (cf . 
Bakker 1994; Sohmer et al. 1991), over 1,200 botanical specimens were obtained 
from areas where eaglewood is now harvested. Among the survey vouchers was 
Takeuchi 4848 (A!, LAE!), collected from a tree described by village guides as 
having a fragrant black wood which could burn even when wet. This specimen 
was identified as Gyrinops kdermannii Domke on the original distribution 
(also confirmed in 1995, cf . annotation on the A duplicate), a species considered 
endemic to the upper Sepik and previously known with certainty only from 
the type gathering (Ledermann 7401; Pentecost Mt along the May River). The 
identification of the 1989 collection was facilitated by comparisons against two 
additional specimens obtained within the last 50 years, but the other reference 
materials have since disappeared from the Lae National Herbarium. 

The Ledermann locality lies ca. 100 km WNW of the April River site des- 
ignated as 'Kamelsriicken' (Samsai Mt) by early German explorers. Samsai vil- 
lagers report that eaglewood is very common on the west-facing slopes of 
Kamelsriicken, m the direction of Pentecost Mt. Because the original specimen 
was taken from the May River, the local testimony implies that the distribu- 
tional center for the species lies further to the west of existing production areas. 
The Hunstein populations are possibly only secondary occurrences. 

Although Ledermann's type was lost in the 1943 destruction of Berlin Her- 
barium, the National Geographic survey voucher was unambiguously keyed 
to the given species using Ding Hou (1960). After eaglewood became a resource 
deserving of scrutiny, suspicion was thus directed to Gyrinops as the likely 
source for the new product.^ However, in the absence of herbarium material 
specifically identified as eaglewood by local producers, it was difficult to be 
sure of the presumed correspondence. Especially as eaglewood was recently 
discovered in Jayapura (West Papua or Papua Barat, the former Irian Jaya), where 
it is there identified as Aquilariafilana (Oken) Merr (Thymelaeaceae), consid- 
erable confusion existed over the plant's identity 

A fertile collection taken during our Ambunti reconnaissance (Towati & 
Golman 159) was pointed out by village suppliers as an example of the tree har- 
vested for the eaglewood trade. The Ambunti voucher includes both flowers 
and fruits, and its completeness leaves no room for doubt that Gyrinops 
ledermanmi is the Hunstein eaglewood. In the Ambunti specimen, the pen- 

men also alternating with a timbnate pctaloid appendage This clearl) climi 
nates Aquilaria from consideration because m that genus the number ot sta- 
mens is twice that of the calyx lobes (Dmg Hou 1960) Aqmim la filay la is 
otherwise very similai to Gyrinops kdcrmannn paiticularly m then shared 
characteristic of loculicidal capsules that develop by ruptuimg thiough one 
side of the corolla tube. It is easy to get the two species contused unless the flow- 
ers are carefully examined. Reports of eaglewood being present in West Papua, 
and its identification as Aquilaria jilaria, need to be reevaluated. Aquilaria has 
thus far been recorded in New Guinea only from the Vogelkop region, so its 
alleged presence at other West Papuan localities such as Jayapura is not unrea- 
sonable. Due to the similarities between the respective taxa, it is entirely plau- 
sible that Aquilaria represents another source for eaglewood. However photo- 
graphs of the Jayapura provenance taken by M. Golman are more suggestive of 
Gyrinops ledermannii than of Aquilaria jilaria (Figs. 2, 3). 

Uncertainty over the identification of eaglewood is also evident among lo- 
cal villagers searching for the plant. Thymelaeaceae is a prominent family in 
the Hunstein flora, with very close similarities among its representatives. 
Within the existing source areas, Phaleria coccinea (Gaud.) F.v.M. 
(Thymelaeaceae) is the species most often mistaken for eaglewood. In superfi- 
cial aspect its inflorescence is comparable to Gyrinops^ and the leaves are also 
of similar appearance, particularly with respect to venation. As with other 

members of the family PI' 

laleria coccinea has an e 

;xtremely strong inner bark 

which is often used m the i 

construction of white bi 

lums (i.e., local netbags used 

as carry-alls) noted for the 

;ir soft texture. 

Due to the black colore 


/OS ptjpuflna Val. (Ebenaceae) 

is also mistaken by village 

rs for eaglewood, even th. 

ough the differences mmor- 

phological aspect are rath 

er obvious. For exampk 

^ unHke eaglewood the leaf 

venation inD.papuana is 

bifacially prommulous 

, and the fruits are mdehis- 

cent and large. The flowe 

rs are also unisexual an 

d dioecfous, rather than bi- 

sexual. Table 1 summarize 

s some of the salient dis 

tinctions between Gyrmops 

ledermannii and the plan 

ts with which it is presei 

ntly confused. 

In addition to the pr 

oblems caused by Sim i 

lanties with other taxa, G. 

Zedermannu IS highly var 

lable and can exhibit co 

nsiderable difforence in leaf 

size between individuals. 

The collection from the 

1989 survey has lanceolate 

blades averaging 5.0 x 2.5 c 

:m but the more recent n 

laterial has elUptic leaves ca. 

19.5 X 6.5 cm. Presumably 

these distinctions are at 

least partly under environ- 

mental control, especially 

m view of the wide ran 

ge of substrates comprising 

eaglewood habitat (cf . Tak 

euchi&r Golman 2002:6: 

3-64; this issue). The nonde- 

'reproductive characters fot Gynnopsledermannii arid the taxaw\lhwU 
I.Tokples names are piovided with the village locality where the na 
!:H&C = R.Hoogland and L Craven, CSIRO survey fronn 1 966;T = W.Takeui 

Scientific name/Tokples r 




Although the economic value of eaglewood affords the promise of substantial 
returns for stakeholders, high market prices clearly create incentives for 
overexploitation and eradication of natural populations. In order to ensure long- 
term viability of existing stocks, appropriate measures in resource management 
will eventually require enactment. However the present dearth of information 
on Gyrinops ledermannii will undoubtedly constrain its sustainable develop- 
ment. Even 90 years after its initial discovery the plant remains poorly under- 
stood by botanical and forestry science. The demography, distribution, and ecol- 
ogy of G. ledermannii are unknown. This situation will require correction before 
eaglewood can be properly developed as an economic asset. There is also an 
obvious need for natural products research into the chemical principles which 
are responsible for eaglewood's desirable qualities. Depending on the nature of 
the resource, the species could conceivably serve as a valuable subject for silvi- 
cultural and natural products development. 

clan of Bugabugi for their goodwill and hospitality during our visits to the April 
River Reviewers John Pipoly III (Fairchild Tropical Garden) and Kevin Janni 
(University of Hawaii) provided helpful comments on the manuscript. 


Bakker, E. 1 994. Return to Hunstein forest. Natl. Geogr. Mag. Feb., 40-63. 

Ding Hou. 1 960.Thymelaeaceae. Flora Maiesiana ser. 1, 6(1 ):1 -48. 

SoHMER, S.H., R. KiAPRANis, A. Allison, and W. Takeuchi. 1991 (unpublished). Report on the 
Hunstein River expedition-1989. Report to the National Geographic Society, Wash- 
ington D.C., U.S.A. 

TAKEucH!,W.and M.GoLMAN.2002.The present status of Ledermann's April River localities in 
Papua New Guinea. Sida 20:55-70. 

ZiCH, F.and J.Compton. 2001 .The final frontier. Towards sustainable management of Papua 
New Guinea's agarwood resource. Traffic Oceania and WWF South Pacific Programme: 


VIasi, 2001. The Sunflower Family in the Upper Midwest: a 
iuide to the Asteraceae in Illinois, Indiana, Iowa, Michigan, 
i Wisconsin. (ISBN 1-883362-11-3, hbk ) Indiana Academy 
lanapolis m collaboration with Chicago Botanic Garden, 
^ A S^OOO 41^1 pp numctouscoioi photos, maps, 7" x 10" 

lentb on Photography' 





US of Konza Piairie Biological Static 

sas are listed along with their 
5 species representing 336 gen 

ual grasses Th. 
provide a flora 
sat Plains regio 

sobrepasa cualquier otra area de tamano compai ihic tn tsl 

vancy initially purchased 371 ha of the site in 1971, followed by the remaining 
procurement in 1977. The preserve is managed for ecological research by the 
Division of Biology at Kansas State University. To study how fire and grazing 
affect the ecosystem, Konza Prairie is parceled into 52 watershed units that are 
burned at different frequencies and in different seasons, and are either ungrazed 
or grazed by large herbivores. Mowed fireguards or gravel roads separate the 
watersheds. Approximately 200 ha of agricultural land and abandoned brome 
fields are scattered throughout the site. The headquarters area consists of old 
homesteads with renovated buildings, a corral, and cultivated fields. 

Topographically the prairie landscape comprises upland plateaus, lime- 
stone hillsides, and fertile lowlands. The upland soils are relatively shallow, silty 
clay loams overlying limestone and shale layers (Udic Argiustolls and Lithic 
HaplustoUs), whereas the lowland soils are deeper coUuvial and alluvial de- 
posits (Pachic Argiustolls). Two major creeks meander through the lowlands, 
but water flow is intermittent and dependent upon precipitation patterns. Per- 
manent pools in the lower creek channels, localized springs, seeps, and several 

small stock ponds constitute the wetland habitat. Gallery forests (bands of de- 
ciduous trees bordering the creeks and ravines) cover about 8.5% of the site. 

Prairie vegetation is dominated by warm-season perennial grasses, with 
numerous interstitial forb species. Although the grassland flora consists pri- 
marily of tallgrass prairie plants, species that are characteristic of midgrass 
prairies thrive on the uplands and hillsides, and many shortgrass prairie spe- 
cies occur on the shallow, droughty sites. 

For nearly 100 years, the site was a cattle ranch with both year-long and 
season-long grazing operations. After acquisition by The Nature Conservancy, 
Konza Prairie was not grazed by large herbivores until 1987, when a bison (Bos 
bison) herd was established in a 992-ha internal enclosure. Seasonal grazing by 
domestic cattle (Bos taurus) resumed in 1992 in 6 adjacent pastures (510 ha), 
although not all are grazed every year. The remainder of Konza Prairie is 
ungrazed by ungulates, except for a prolific white-tailed deer (Odocoileus 
VI rgmiflnus) population. 

Konza Prairie has a temperate continental climate, characterized by hot 
summers, cold winters, and moderately strong surface winds. Average annual 
precipitation is 835 mm, with 75% of this occurring in the April through Sep- 
tember growing season. The average frost-free season lasts 180 days. Mean an- 
nual temperature is 13° C, with average minimum and maximum temperatures 
ranging from -3° C in January to 27° C m July. Elevations on Konza Prairie range 

After a decade of collecting, 436 species of vascular plants (plus 5 varieties) 
were initially reported to occur on Konza Prairie (Freeman & Hulbert 1985), 
with another 27 species subsequently added (Freeman & Gibson 1987). Six years 
later, Kazmaier (unpublished pamphlet) listed 524 vascular plant species for 
Konza Prairie. Because of Konza Prairie's role as a benchmark tallgrass prairie, 
it is essential that the flora inventory and pertinent ecological synopsis of the 
species be current and readily accessible. With more than 70 new species dis- 
covered since Kazmaier's unpublished checklist, questionable validity of some 
previously listed species, and extensive nomenclatural revisions, an updated 
inventory of the Konza Prairie vegetation is necessary. Format for this flora fol- 
lows standards proposed by Palmer et al. (1995). Voucher specimens are stored 
in the Kansas State University Herbarium (KSC). 

The vascular plants known to occur spontaneously on Konza Prairie are ar- 
ranged alphabetically by family, genus, and species within four groups: 
Ptendophyta (ferns and fern allies), Pinophyta (gymnosperms), Magnoliopsida 

(dicots), and Liliopsida (monocots). Each entry includes scientific name, com- 
mon name, relative abundance, primary habitat, flowering period, and life span 
(Appendix I). In addition, non-native species and the photosynthetic pathway 
of grass species are listed. 

Nomenclature follows Kartesz (1999), although authority abbreviations 
conform to Brummitt and Powell (1992). A synonym is enclosed in brackets for 
those species that have been annotated recently or if the scientific name changed 
from that listed by Freeman and Hulbert (1985). Infraspecific taxa (subspecies 
and varieties) are not delineated, although naturally occurring hybrids are 

The common name for each species was selected from regional sources 
(Anderson 1961; Bare 1979; Great Plains Flora Association 1986) to reflect cus- 
tomary usage for this area. Most species are identified with a single colloquial 
name, but alternatives are presented in a few instances. Structure and format of 
common names generally follow guidelines recommended by Kartesz and 
Thieret (1991). 

The listed habitat indicates where each species is most likely found on 
Konza Prairie; however, it does not imply that the plant is restricted to that site. 
Many species occur at different densities in heterogeneous locations. Designated 
habitat categories for the Konza Prairie flora are: 

Prairie — Open grassland that is subdivided by topographic position: uplands 
(clayey, cherty, or shallow soils on top of the hills), lowlands (deep bot- 
tom land soils), and slopes (rocky hillsides). Additionally, some species 
are listed as occurring predominantly in grazed or infrequently burned 

Woodland— G 

forest and sit 

es that an 

^shaded by 


Wetland— We 

t sites 

that include 

ponds, en 

eeks, seeps, 

stock tanks, and mesic 



Disturbed— S 

ites wl 

lere the soil 

has been 


include trails, wallows, 


roadsides, homeste; 

ads, lawn5 

5, and culti\ 

^ated fields. 

Richness amc 

.ng the 

different pla 

nt habitai 

.s was estimated by assigning each 

species to one 


It. Because m; 


2S occur m ; 

i variety of habitats, as- 

signment to .i 

i speci 

lie category I 

na)' be dc 

•batable. Ht 

)u-ever, the criterion for 

assigning spec 

is based on wl 




cally associat( 

:d wit 

h on Konza P 


'cics that arc found predominantly 

in disturbed s 

ites \\'( 

:rc not consic 

Icred prai 

ne plants e 

ven if they appear spo- 

radically mg 

razed c 

prainc. 1 i 

kcwise, wo. 

odland species that mi- 

grate into unt 



M-c not grc 

lupedin th( 

^ prairie habitat. 

Relative abundi 

\luation t 

>l how com 

mon and widespread a 

species is for the particular habitc 

It, and IS 1 


tensive field experience 

of the author.: 


er, abundance 



1 some perennial species 

may fluctuate 


'm response t 


ation patter 

■ns and grazing or burn- 

s. Desc 


£ five abundance classes are: 

Abundant — Species is dominant or co-dominant in the habitat. 

Common — Species is found easily m the particular habitat. 

Occasional — Species occurs sporadically but is not difficult to find. 

Infrequent — Species is relatively scarce and not easily found. 

Rare — Species is seldom encountered or found only in specific locations. 
The flowering period is when peak blooming normally occurs on Konza Prairie 
and often indicates when the species is most likely found. For vascular crypto- 
gams, the dates represent spore formation. Flowering periods are based on field 
observations, herbarium specimens, and phenology records for this area. 

Each species is classified by life span as either annual, biennial, or peren- 
nial. Non-herbaceous perennial plants were additionally subdivided into tree, 
shrub, and woody vine categories. Although some biennials and short-lived 
perennials may persist for only one year, designations were based on the ex- 
pected longevity of the species for this area. 

Grasses (Poaceae) are designated as either C3 or C4 species (Waller & Lewis 
1979). The C3 grasses are cool-season plants that initially produce a 3-carbon com- 
pound during photosynthesis and characteristically flower in May and June. The 
C4 grasses are warm-season plants that initially produce a 4-carbon compound 
m the photosynthetic pathway and characteristically flower in July and August. 

Species that were not present in this region before European settlement are 
considered introduced (Brooks 1986). Cultivated crops from the agricultural 
fields (e.g., wheat, oats, sorghum, corn, and soybean), and other non-natural- 
ized annual species are not included in this checklist. However, exotic horticul- 
tural and ornamental species that were planted decades ago near the headquar- 
ters area and have subsequently persisted, are listed in Appendix II. 

The known vascular flora on Konza Prairie consists of 576 species, represent- 
mg 336 genera and 96 families (Table 1; Appendix I). Families with the most 
species are Poaceae (84), Asteraceae (79), Fabaceae (49), and Cyperaceae (33). 
Forty families are represented by a single species. The most species-rich genera 
are CarcxiU species), Polygonum (12 species), Symphyotrichum (9 species), and 
Asclepias (9 species). 

Twenty-three species from the last inventory (Kazmaier, unpublished pam- 
phlet) are deleted from the present list because the specimens were misidentified 
(n = 8), had been reduced to synonymy (n = 4), or because there are no vouchers 
and no bona fide evidence of their presence on Konza Prairie. Most of the un- 
substantiated species appeared in preliminary inventories (see Freeman & 
fiulbert 1985), and their unverified presence was subsequently perpetuated. 
Taxonomic revisions of Kazmaier's unpublished checklist also transferred 55 
species into different genera and changed the specific epithet of 19 species. 

When species are categorized to one predominant habitat, 40.6% of Konza 
Prairie plants are characteristically associated with prairie grassland, followed 
by disturbed (22.4%), woodland (22.2%), and wetland (14.8%) habitats (Table 
2). In the prairie habitat, 16.2% of the species are grasses and 76.9% are herba- 
ceous forbs. Only seven woody species are characteristically associated with 
the prairie habitat; but without frequent fire, many woodland species invade 
and proliferate. Species richness is lowest in wetland sites, but these areas are 
the smallest and most fragmented habitats on Konza Prairie. 

When partitioned into life span categories, 316 species (54.9%) are herba- 
ceous perennials, 56 species (9.7%) are woody perennials, 180 species (31.3%) 
are annuals, and 24 species (4.2%) are biennials. Within the prairie habitat, 76.1% 
of the species are perennials, 20.9% are annuals, and 3.0% are biennials. In com- 
parison, 23.3% of the species in disturbed sites are perennials, 69.0% are annu- 
als, and 7.8% are biennials. 

Non-native species (n = 96) account for 16.7% of the total flora. Families 
with the most introduced species are Poaceae (21), Brassicaceae (13), Fabaceae 
(9), and Asteraceae (8). Annuals and biennials comprise 59.4% of all introduced 
species, compared to 30.8% of the native flora (Table 3). The highest density of 
exotics is in the headquarters area where many species were planted intention- 
ally or are cosmopolitan weeds. Most introduced species are either restricted to 
disturbed soils or are weak competitors in native prairie. However, some inva- 
sive species are capable of displacing native vegetation. Because the aggressive- 
ness of four exotic species (Bothriochloa bladhii, Bothriochloa ischaemum, 
Carduus nutans, and Lespedeza cuneata) potentially threatens the integrity of 
tallgrass prairie, they have been specifically targeted for eradication. 

The C4 photosynthetic pathway occurs in 59.5% for all grass species and in 
65.1% of the native grasses. Of the 24 annual grasses that occur on Konza Prai- 
rie, 17 species (70.8%) use the C4 photosynthetic pathway Most dicotyledons 
use the C3 photosynthetic pathway, but 14 species representing five genera 
(Amaranthus, Bassia, Chamaesyce, Portulaca, and Trihulus) have C4 photosyn- 

Table 3.Ther 


— — -- 

-— -'-— °- 

Native Species 

Introduced Species 

Life form 

Number % 

Number % 

thesis (Downton 1975). All of the Konza Prairie forb species that utilize the C4 
photosynthetic pathway are annuals. 

Although field collections of Konza Prairie flora have been ongoing for more 
than 25 years, the compilation of vascular plants vv'ill continue to be dynamic. 
New immigrants, ephemeral annuals that fluctuate with precipitation patterns, 
and superficially similar species will undoubtedly be found in subsequent years. 
Species also will disappear Some species have not been observed on Konza Prai- 
rie for more than 10 years and may be extirpated. Other species occur in a single 
locale or consist of only a few plants. Few areas of tallgrass prairie have been 
surveyed as intensively as Konza Prairie, and the diversified habitats created by 
management and research activities provide a flora richness that is likely 
greater than any other area of comparable size in the Great Plains region. 

Annotated Checklist ol Spt 
PTERIDOPHYTA - Vascular Cryptogams 4, 

EQUISETACEAE- Horsetail family i 

^^^"!Vl""I "/" /"T I""'"^ r^Ti°es^MT -^Jun ANACARDIACEAE - Cashew family 

quent on uplands and lowlands Apr-May 
OPHIOGLOSSACEAE - Adder's tongue family shrub 

rattlesnake fern /?/iusg/a6raL smooth sumac common on shal 

Ophioglossum engelmannii PrantI adders Toxicodendron mdicans (L) Kuntze poison ivy 
tongue rare in woody lowlands Apr-Jun common in woodlands May-Jun woody 

PTERIDACEAE - Maidenhair fern family ANNONACEAE - Custard-apple family 

[Notholaena dealbata] false cloak fern rare lands and along creeks; Apr-May; tree, 

ft // Tu^^MI I'Ma^Txtuhn TrJ^oith cTff brake- ^^^^^^^^^ ' Parsley family 

DIVISION PINOPHYTA - Gymnosperms Chaerophyllum tainturieri Hook,; erect chervil; 

it woodland edges and in d 


iomatium foeniculaceum (Nutt.) Coult. & Rose; 
Class Magnoliopsida - Dicots uplands and slopes; Apr-May; perennial. 

ACANTHACEAE- Acanthus family Osmorhiza longistylis (Torr.) DC; long-style 

I 1 Link;hedge-parsle 
il introduced annual. 

APOCYNACEAE - Dogbane family 

-Milkweed family Arnoglossum plantagineum ^ Raf.; [Cacalia 

,^ y : I ' casional on uplands and lowlands; Jun-Jul; 

coreopsis beggar-ticks; infrequent in wet 

Bidensbipinnata L; Spanish needles; rare in wet 

Bidens cernua L; nodding beggar-ticks; infre- 

e/densfrondosoL; devil's beggar-ticks;occasional 
in wet sites; Sep-Oct; annual. 

euporono/We5];false-boneset; common on 

Valter; spider milkweed; com- imliuuulcu uieniiidi. 

? (Michx.) Pers.; climbing-milk- nion m unburned lowlands; Aug- 

ASTERACEAE - Sunflower family Jun-Jul; perennial. 

Achillea millefolium L.;western yarrow;common Qy^,-^^ w/gore(Savi) Ten.; bull thistle; rare in dis- 

on all prairie sites; May-Jun; perennial, Curbed sites; Jun-Sep; introduced biennial. 

woodlands; Aug-Oct; perennial. Sep;annual. 

Ambrosia psilostachya DC; western ragweed; annual 

perennial. prairie dogweed; occasional along roads; 

Ambrosia triflda L.; giant ragweed; occasional in Jul-Oct; annual. 

disturbed sites and at woodland edges;jul- Echinacea angustlfolia DC; black-sampson 

Oct;annual. echinacea- common on slopes and rocky 

Amphiachyris dracunculoldes (DC) Nutt.; nrairiP- inn'nPrPnni;,! 

Erechtites hierac 

gayfeather; common on shallc 
Nothocalais cuspidata (Pursh) Greene 

'lianthus maximilianii Schrad.; Ma> 
Aug-Oct; perennial. 

woodland edges; Aug-0 

•jm (L.) Small; [Solidago rigida]; 

Nutt.) Weber & Love; [Senecio 

right prairie-coneflower; 

Aug-Sep; perennial. 

er; rare 

Hieracium longipilum Torr 



Hymenopoppus scabiosaeu 

s L'Her. 

Solidago canadt 

Solidago gigant 

nail prairie sites;Aug-Oct; perennial. 

Solldagopetiolaris Aiton;downy golder 
quent on rocky prairie; Sep-Oct; p 

Solidagospeciosa Nutt.;showy-wand g 
infrequent on uplands; Sep-Oct; p 

BETULACEAE- Birch family 

ynoglossum officinale L.; Iiount 

on uplands; Sep-Oct; perennial, Myosotis vema Nutt; Virginia forget-me-not; in- 

Symphyotriclium oolentangiense (Riddell) frequent on uplands;May;annual. 

Nesom [Aster oolentangiensis]; azine aster; Onosmod/c/m mo//e Michx.;western marbleseed; 

Draba cuneifolia Nutt. ex Torn & A. Gray; wedge- [qIj^jj^ siphilitica L , blue 

sites; Apr-May; annual. TrioTanis lepwcarpa {Nw 

Draba reptans (Lam.) Fernald; white whitlow- nus'-looking-glass; i 

pidium oblongum Small; t 
weed; occasional in d'n 

Triodanisperfoliata (L.) NieuwI.; clasping Ver 
''''^^ looking-glass; occasional on uplands 

jnd CANNABACEAE- Hemp family 

niai. Cannabis sativa L; marijuana, hemp; occasr 
ion in ditches, at woodland edges, and aro 

ual. CAPPARACEAE- Caper family 

ndalon roads' CAPRIFOLIACEAE Honeysuckle 

'■[Thlaspi perfo- -'"'-"" ' ' ' '' """ 

fior/ppo po/usfr/s(L.) Besser;bog yellowcress;rare 

/?onpp(7se55////7ora (Nutt.) Hitchc.;stalkless yellow- "os eu per o i u ., 9 ' 

Ror;ppos;nuota(Nutt.)Hitchc.;spreading yellow- ^ 

cress; infrequent along creeks; May-Jun; CARYOPHYLLACEAE - Pink family 

CACTACEAE - Cactus family Cerastium brachypodum (Engelm. ex 

CELASTRACEAE - Staff-tree family 9'"^^' '^"f^^^^^'^ '^i aisiuroea lowiana. 

Celastrus scandens L ■ American bittersweet- in- ^^P' P^'^^nnial. 

frequent in woodlands and on creek banks; CORNACEAE - Dogwood family 

Euonymus atropurpurea Jacq,; eastern wahoo; vines and on wooded slopes; May 

CHENOPODIACEAE - Goosefoot family 

CRASSULACEAE - Stonecrop family 

Penthorum sedoides L.; ditch stonecrop; infre 

CUCURBITACEAE - Cucumber family 

Cucurbita foetidissima Kunth; buffalo gourd; ir 


)dland edges and 



sites; Apr-May; annual. 

wort; infrequent on uplands and tov 

CUSCUTACEAE - Dodder family 

Cuscuta glonierata Choisy; cluster doddei 

Standley's __ parasitic on various Asteraceae sped 
" iscuta indecora Choisy; showy dodder 

EUPHORBIACEAE - Spurge family 

Chamaesyce glyptosperma (Engelm.) Small; [Eu- Tragia b 
phorbia glyptosperma]; ridge-seed mat- occe 

Chamaesyce maculata (L.) Small; [Euphorbia on shallow uplands and rocky sites;Jun-Aug; 

maculata]; spotted mat-spurge; occasional perennial. 

on grazed uplands and in disturbed sites; p^BACEAE - Bean family 

Chamaesyce missurica (Raf.) Shinners; [Euphorbia '"°^J^ ^^ p^airie^sites- Ju^n-Juh shru'b ' ^°^'^°" 

Chamaesyce r}utans [lag.) Small; [Euphorbia . , , , 

nutans]; eyebane; occasional in unburned Amnhirarnaea hracteata (I ) Fernald- hoa-nea- 

uplands and lowlands; Jul-Oct; annual. nut;occasionai on shaded creek banks; Aug- 

prostrata]; prostrate mat-spurge; occasional ^...fgl^'^^/s L.; Canadian milk-vetch; 

iroduced'amual '' ^'^^^'^^'-^^P' rare on shaded slopes; Jul; perennial. 

Chamaesyce serpens (Kunth) Small; [Euphorbia vetch- occasional on unlands- Aor-Mav 

serpens]; serpent mat-spurge; infrequent on nerennial 

grazed uplands and in disturbed sites; Jul- /^,,,agalus htiflorus Hook.; lotus milk-vetch; in- 

ChZaeTcesfictos ora En elm Small- Eu her ^''^'"'^ °' '^'"°'' 'P''""^' '""^ ''°P''' 

6,o^sr,cro5pora];slim-seed rTiat^spurge;infre- AstrTgX'mZZens,s Nutt.; Missouri milk- 

"* '" IS ur e 51 es u c, annua. vetch; infrequent on shallow uplands and 

iTnthTt^^^Z^ntseed Astragalul'l^a7te!^^^^^^^ 

isional on shallow uplands and slopes; e^pJ'LTsf'r^/s (l^^Bn ex Alton; blue'wTd^^^^^^^^^^ 

■^^P'^^^^i- ^, ^ common on uplands; May-Jun; perennial. 

la coroliata L; flowering spurge; infre- g^^^,.^,^ ^ 6,co/or Greenm.S Larisey;[e.aasrra//5 

It on uplands and lowlands; Jul Sep; ^ g bracteata]; bicolor wild-indigo; rare on 

'^'^'^ ■ ^ uplands; May; perennial. 

lacyaopora urray.pain e spurge, eapf/s/abracteafoMuhI.ex Elliott;plains wild-in- 

.dsites;Jul-Sep;annual. ^^ ,^^,3^^ 

3 hexagona Nutt. ex Spreng.; six-angle roronjiir, varir 

l-Sep;annuaL prairie adj. 

"^^.^ Jun-Sep;ir 


Dolealeporina (Alton) Bullock;fox-tail prairie-cio- Medicago saliva L.; alfalfa; r; 

clowlands;Aug-Sep;annual. brome fields and ditches; Jun-Sep; intro- 

is officinalis (L) Lam,;yellow s' 

mosa nuttallii (DC) Turner; [Scliranl<io nu 

diomelum argopiiyilum (Pursh) Gr 
[Psoralea argophylia]; silver-leaf scu 

n lowlands;Aug-Sep;perennial. Pediomelum esculentum (Pursii) Rydb.; [Psorale 
inosum (Muhl.ex Willd.) Wood; escu/enro];breadrootscurfpea;occasionalo 

clover;occasional at woodland edges; Aug- Senna marilandica (L.) Ur\k;[Cassia nnarilandica]; 

Sep; perennial. wild senna; infrequent at woodland edges 

Desmodium sessilifolium (Torr.) Torr. & A. Gray; and on lowlands; Jul-Aug; perennial. 

sessile tick-clover; infrequent on lowlands Sfrop/?osfy/es/ie/vu/a (L) Elliott;trailingwildbean; 

and slopes; Aug-Sep; perennial. rare on shaded creek banks;Aug-Oct;annual. 

Qleditsia triacanthos L.;honey-locust;occasional Stroptiostyles leiosperma (Torr. & A. Gray) Piper; 
in woodlands and unburned lowlands; (Vlay; smooth-seed wildbean; infrequent on low- 
tree, lands; Aug-Sep; annual. 

G/ycyrr/i/za/ep/doro Pursh; American licorice; in- Trifoliunn pratense L.; red clover; infrequent 

frequent on uplands and !owlands;Jun-Jul; around headquarters;May-Sep;introduced 

perennial. perennial. 

Gymnoc/adL/sc//o/cus(L.) Koch; Kentucky coffee- Trifolium repens L.; white clover; infrequent 


Nepefacafar/a L, catnip, occc 

isional in shade 

GENTIANACEAE - Gentian family 


troduced pere 

Gentiana puberulenta Pringle, downy gentian 

- Prunella vulgaris I, selfheal. 

infrequent in V 

rate on lowlands Sep-Oct, perennial 

lands and along creeks. 


GERANIACEAE- Geranium family 

m , [Salvia piu 


Pitcher's sage, blue sac 


Salvia reflexa Hornem,lar 

ice leaf sage; 

GROSSULARIACEAE- Currant family 


>s and along , 

)uri gooseberry 

1 t iht, ' iii.i. Apr-May,shrub 


ocky sites- Mav 

HIPPOCASTANACEAE - Buckeye family 


Ohio buckeye, infrequent Teucnum canadense 
1 along creeks, Apr-May, common on lowl 

HYDROPHYLLACEAE - Waterleaf family 

LYTHRACEAE - Loosestrife family 

WK, M m^M--i '^ " ' vvww^.o, ,^., r^uy w.., MALVACEAE - Mallow family 

oh/sp/da Pursh;roughfalse-penny-roya!; sionaln dsr bed In lad 

itus (L.) Britton et al.; [Trichostema 

(Torr, & A. Gray) A. Gray; 

OROBANCHACEAE - Broomrape family 

MENISPERMACEAE - Moonseed family 

MOLLUGINACEAE - Carpetweed family OXALIDACEAE - Wood-sorrel family 

MORACEAE- Mulberry family ^ Jun; perennial 

CH fi ■; h -H Oxo//s srr/cro L; yellow wood-sorrel; occ. 
hedge-apple; infrequent in unburned sites ^ 

NYCTAGINACEAE - Four-o'clock family 

; narrow-leaf PHYTOLACCACEAE - Pokeweed famil; 

inds;Jun-Sep; Phytolacca americana L.; pokeweed; occa 

ONAGRACEAE - Evening-primrose family 

PRIMULACEAE - Primrose family 

POLEMONIACEAE - Polemonium family ^^^^ sites, Apr-May; annual. 

Phlox divaricata L • blue phlox- occasional in ^y5'mac/]/aa//ofaL -fringed loosestrife;rareir 

POLYGALACEAE - Milkwort family RANUNCULACEAE - Buttercup family 

Polygonum hydropiper L.; swamp water-pepper; 

sites; Apr-May; annual. 

rare in wet sites; Jul-Oct; annual. 

Polygonum hydropiperoides Michx,; mild water- 

in wet sites; Apr-May; biennial. 

pepper; rare in wet sites; Jul-Oct; perennial. 

r/ia//cfrt;mdasycorpumFisch.&Ave-Lall.; purple 

Polygonum lapathifolium L.;pate smartweed;in- 

meadow-rue; rare in woodlands; May-Jun; 

frequent in wet sites; Jul-Oct; annual. 


Polygonum pensylvanicum L; Pennsylvania smart- 

RHAMNACEAE - Buckthorn family 

Polygonum persicana L; lady's-thumb smart- 

Ceanof/i[is/ief6aceusRaf; New Jersey tea, inland 


ceanolhiis, common in unburned uplands 

duced annual. 

and slopes; Apr-May; shrub. 

Polygonum punctatum Elliott; dotted smartweed; 

ROSACEAE- Rose family 

infrequent in wet sites; Jun-Oct; perennial. 

Agrlmonia pubescens Wallr.; downy agrimony; 

Polygonum ramosissimum Michx.; bushy knot- 

rare at woodland edges;Jul-Aug;perennial. 

weed; infrequent in disturbed sites and 

Craregusmo/foScheele; downy hawthorn, sum- 

along roads; Aug-Oct; annual. 

mer haw; rare on wooded slopes and along 

quent at woodland edges and in disturbed 

Geum conodenseJacq.;whiteavens;common in 

sites; Aug-Sep; perennial. 

woodlands and unburned lowlands;Jun-Jul; 

Polygonum tenue Michx.; slim knotweed; infre- 


quent on uplands;Jul-Oct;annual. 

Potentillaarguta Pursh;tall cinquefoil; infrequent 

Polygonum virginianum L.; Virginia smartweed; 

on lowlands; Jun-Jul; perennial. 

PORTULACACEAE - Purslane family 

-ned lowlands; May- Leucospora multifida (Michx.) Nutt.; paleseed; 

ih-bush blackberry; A4/mu/usg/ofcirafusKunth;yellow monkey-flower; 

and unburned low- infrequent in wet sites;May-Aug; perennial. 

Penstemon cobaea Nutt.; cobaea beardtongue; 

occasional in woodlands;Jul-Sep;perennial. 

RUTACEAE - Citrus family 


arshall; black will 


SOLANACEAE - Nightshade family 

Datura stramonium L.;jimsonweed; rare in di: 

turbed sites; Jul-Oct; introduced annual. 
Physalis hederifolia A. Gray; heart-leaf grounc 

Apr-Jun; perennial. Physa/ZspubescensL; downy ground-cherry; n 

woodlands and along creek5;Jun-Jul;an 

SCROPHULARIACEAE - Figwort family p^ sails pumila Nutt prairie qround-cherr ( 

Agalinisaspera (DougI.ex Benth.) Britton;rougli ^ Jof^ on all nr^airie sites- Jun-Sen-nereni 

Solanum ptychanthum Dum\;b\ackn\g[MsUade; Verbena srr/cfaVent.;woollyverbena;occasionai 

rare in wet sites and ditches;Jun-Oct;annual. on uplands and along roads;Jun-Sep;perennial. 

So/anumrosrratu/nDunal;buffalo-bur nightshade; Verbena urticifolia L.; white verbena, nettle-leaf 

occasional in disturbed sites;Jun-Sep;annual. verbena;occasional in woodlands and along 

ULMACEAE- Elm family 

woodlands; Apr-May; tn 

in wet sites; Aug Oct; annual, ZYGOPHYLLACEAE - Caltrop family 

VERBENACEAE - Verbena family troduced annual 

Glandularia bipinnatifida (Nutt.) Nutt.; [Verbena Class LILIOPSIDA - Monocots 

bipinnatifida]:D3kot3 verbena;occasional on ^GAVACEAE - Agave family 

lanceolata]; northern 

COMMELINACEAE - Spiderwort family 

at woodland edges; Jun-Oct; perennial. Larexv 

spiderwort; occasional in mesic lowlands; '^^P^^^ 

Tradescantia ohiensis Raf.;Ohio spiderwort; rare Sep; annual. 

in mesic lowhnds-Mav-Jun'Derennial Cyperus bipartitusJorc.lCyperus rivularislbroo 
flat-sedge; infrequent in wet sites; Jul-Sef 
CYPERACEAE- Sedge family annual 

Bolboschoenus fluviatilis (Torr.) Sojak; [Scirpus Cyperus esculentus L.; yellow nut-sedge; rare ii 

CarexalbicansWMd.exSpteng-lCarexartitecta]; Cyperus lupulinus{Spreng)Marcks;p\a\nsM-sedgi 
white-tinge sedge; infrequent in woodlands; common on uplands; Jun-Jul; perennial. 

May-Jun; perennial. Cyperus x mesochorus Geise; [C. lupulinus x C 

Carexbrachyglossa Mack.;[Carexannectens]:ye\- flat-sedge; rare in wet sites;Jul-Aug;annual. 

low-fruit sedge;occasional in wet sites;May- Cyperus strigosus L.; straw flat-sedge; infrequent 

June; perennial. in wet sites; Aug-Sep; perennial. 

Carex brevior (Dewey) Mack, ex Lunell; straw Eleocharis compressa Sull.; flat-stem spike-rush; 

sedge; common on uplands and lowlands; common in wet sites; Apr-Jun; perennial. 

May-Jun; perennial. Eleocharis erythropoda Steud.; red-foot spike-rush; 
Carex davisii Schwein. & Torn; Davis' sedge; rare common in wet sites; May-Jun; perennial. 

Carex gravida Bailey; heavy sedge; c 

Carex/iysreric/noMuhl.exWilld.;bottle-brush sedge; 'R'DACEAE Ins family 

common in wet sites; May-Jun; perennial. Sisynnciiium campestre Bickr 

Corex mead// Dewey; Mead's sedge;abundanton JUNCACEAE - Rush family 

jdulus Muhl.;drooping bulrush;com- 

LEMNACEAE - Duckweed family u^.,nu.,nuu u 

Lemna minor L .; lesser duckweed; rare in shak bladhii]; d 

ow water, u- ep,perennia. Jul-Oct;in 

LILIACEAE- Lily family Bothnochloais 

infrequent on uplands; Aug-Sep; perennial. Bothriochloalaguroides (DC.) HenerAAndropogon 

Asparagus officinalis L.;asparagus;rareat wood- sflccf]aro/c/es];5ilverbluestem;rare in ditches 

Erythronium albidum Nutt.;white fawn-lily; infre- Bouteloua curtipendula (Michx.) Torn; side-oats 

quent in woodlands;Apr; perennial. grama; abundant on uplands and slopes; 

Ornithogalum umbellatum L.;star-of-Bethlehem; Jul-Sep; perennial C 

rare around headquarters; Apr-May; intro- Bouteloua gracilis (Willd. ex Kunth) Lag. ex 

duced perennial. Griffiths;bluegrama;occasional on uplands 

Po/ygonarum6//'/o/-um (Walter) Elliott; Solomon's- 3^^ slopes; Jul-Aug; perennial C 

„,„^^i-,,^^(^gi^Piial Bouteloua hirsuta Lag.; hairy grama; infrequent 


NAJADACEAE - Naiad family 

Najasguadalupensis (Spreng.) Magnus;cor 
naiad;occasional in ponds;Aug-Sep;a 
ORCHIDACEAE - Orchid family 

POACEAE - Grass family 


Agrostis stolonifera L; redtop; infrequent along infre 

creeks; Jun-Jul; introduced perennial C3. Jul- 

Alopecurus carolinianus Walter; Carolina foxtail; Chloris ' 

s Muhl. ex Willd.; 

(-tail barley; infrequet 





woolly dich 


; occasional ( 

rie sites; Mc 




prairie site< 

n oligosar 


perennial C 


It.) Pilg.; [b 

grass; infre- 

uplands an 

d slopes; Jul-Sep;pere 







;ncato (Beauv.)Fernald, 

grass; occa 

sional in d 

itches and ( 

ep; annual 


jsine indica 

(L) Gaertr 

1 /goose gr; 





and along roads; Jul-Oct;annu; 

roads; Jul-5ep;annual C,. creeks; Aug-Oct; perennial Q. 

Festuca subverticiliata (Pers.) Alexeev; [Festuca Muhlenbergia mexicana (L.) Trin.; leafy muhly; 

obtusa]; nodding fescue; infrequent in infrequent in woodlands and along creeks; 

woodlands; May-Jun; perennial C3. Sep-Oct; perennial C^. 

Glycerio striata (Lam.) Hitchc.;fowl manna grass; Muhlenbergia racemosa (Michx.) Britton et al.; 

woodland edges and in mesic lowlands, 

5o/gha.trumnuram(L.) Nash; Indian grass;abun- 

Aug-Oct; perennial C,. 

dant on all prairie sites;Aug-Oct; perennial C,. 

yVluh/enfeerg/a sc/ireber/Gmel,; nimblewill; occa- 

sional in woodland5;Aug-Oct; perennial C4. 

near cultivated fields; Jul-Oct; introduced 

Panicum capillare L ■conimon witch grass;occa- 

perennial Q, 

sional in disturbed sites and along roads;Jui- 

Panicum dichotomiflorum Michx.; fall panicum; 

ditches; Jul-Sep; perennial C,. 

occasional in disturbed sites and along 

Sphenopholis obtusata (Michx.) Scribn.; prairie 

roads; Aug-Oct; annual C,. 

wedgescale;occasional on uplands and low- 

lowlands and in mesic uplands; Jul-Sep; pe- 

Sporobolus compositus (Poir.) Merr.; [Sporobolus 

rennial C,. 

osper];tall dropseed;abundant on all prairie 

sites and in abandoned brome fields; Aug- 

all prairie sites; Jul-Oct; perennial Q. 

Oct; perennial C,. 

Pascopyrum smithii (Rydb.) Love; [Agropymn 

Sporobolus cryptandrus (Torr.) A. Gray; sand 


osus Poir.; long-leaf pondweed; 

rennial Q. Potamogetonpusillus L.;baby pondweed; rare in 

taria pumila (Poir.) Roem. & Schult.; [Setaria ponds- Jun-Auq- perennial 

d\l'^!^fhe7!!a^sMln^^^^^^ SMILACACEAE - Catbrier family 

TYPHACEAE - Cat-tail family 


Numerous individuals have contributed to compiling the Konza Prairie flora, 
but the most prominent predecessors were Craig Freeman, Richard Kazmaier, Lloyd 
Hulbert, and Linda Marple. Ted Barkley and Iralee Barnard were instrumental 
in persuading me to collect formally and update the Konza Prairie flora. Earlier 
drafts of this manuscript were improved by comments from Carolyn Ferguson, 
Mark Mayfield, and Iralee Barnard. I also appreciate the meticulous review and 
insightful comments from Robert B. Kaul and Craig C. Freeman. The National 
Science Foundation Long-Term Ecological Research Program, Konza Prairie 
Biological Station, Kansas State University Herbarium, and the Kansas Agri- 
cultural Experiment Station provided support for this project. This report is 
contribution no. 02-122-J of the Kansas Agricultural Experiment Station. 

Anderson, K.L. 1 961 . Common na 

Stat. Tech. BulL No. 1 1 7, Manhc 

Bare, J. E. 1 979. Wildflowers and v^ 

DowNTON,WJ.S. 1 975.The occurrence of C. 

Freeman, C.C. and L.C. Hulbert. 1 985. An an 

Research Natural Area, Kansas. Trans, h 

Freeman, C.C. and 

D.J.G1BS0N. 1987. Additions to 

the vascular flora of Konza 

Prairie Resear 

Natural Area, 

Kansas.Trans. Kansas Acad. Sci. 90:81-84. 

Great Plains Flor 

A Association. 1 986. Flora of t! 

he Great Plains. University 

Press of Kans 

Lawrence, KS. 

Kartesz, J.T.I 999. 

A synonymized checklist anc 

J atlas with biological attnt 


cular flora oft 

he United States, Canada, an- 

i:J.T. Kartesz ai 


1. Synthesis of the North Am 

ericanflora,Version 1.0. North Carolina E 

tanical Garde 

n. Chapel Hill. 

Kartesz, J.T and. 

I.W.Thieret. 1991. Common nc 

^mes for vascular plants: G 



3ns. Sida 14:421-434. 

Palmer, M.W., G.L 

.. Wade, and R Neal. 1 995. Standards for the writing of f 



American gra 

.K.Lewis. 1979.0ccurrence of C 
sses. J. Range Manage. 32:1 2- 



Marco Antonio Bataiha Fernando Roberto Martins 

Department of Botany Department of Botany 

Federal University of Sao Carlos Institute of Biology State University of Campinas 

Box 676, Sao Carlos, SP, 13565-905, BRAZIL PO Box 61 09, Campinas, SP, 1 3083-970, BRAZIL 

rrado, a savanna-like ecosystem, is the second largest vegetation type in Brazil, originally cov- 

l about 133,000 ha, is one ol the most important reserves within the cerrado. From November 
3 October 1999, we carried out a floristic survey of all the cerrado physiognomies of the ENP 
and 601 species, belonging to 303 genera and 80 families. Among the collected species, 12 were 
and seven were new to science. The herbaceous to woody species ratio was 3.03:1. The richest 
ts were Asteraceae (88 species), Fabaceae (87), Poaceae (51), Myrtaceae (39), and Lamiaceae 

:ance of the ENP to cerrado conservation, since from 8 to 20% of the species recorded for this 
tion type occur within the reserve. We emphasize the need for more floristic surveys in which 

ae (24), compreendiendo 48% del t 


The Cerrado Domain, the second largest Brazilian phytogeographic province, 
once covered about 2 million km-, or 23% of the Brazilian territory, with its 
core area in central Brazil (Ratter et al. 1997). As its name implies, cerrado veg- 
etation prevails m the Cerrado Domain. The cerrado vegetation is not uniform 
m physiognomy (Coutinho 1990), ranging from grassland to tall woodland, but 
most of its physiognomies within the range defined as tropical savanna 

(Sarmiento 1983). In the Cerrado Domain, interspersed with the prevaihng 
cerrado vegetation, there are other vegetation types, such as seasonal forest, ri- 
parian forest, rocky campo, and wet campo. 

Although frequently neglected in the past, the cerrado vegetation stands 
out in Its high floristic richness (Ratter et al. 1997). After comparing a large 
number of floristic and phytosociological surveys carried out in cerrado sites 
from all over Brazil, Castro et al. (1999) estimated that 3,000 to 7,000 vascular 
plant species occur in this vegetation type. In addition to its high floristic rich- 
ness, the cerrado presents a high degree of endemism. Lenthall et al. (1999), for 
instance, listed 234 woody species occurring in 10 cerrado sites and verified 
that 80% of them were restricted to this vegetation type. Owing to its high rich- 
ness, high degree of endemism, and present conservation status, Fonseca et al. 
(2000) included the cerrado among the biodiversity hotspots for highest prior- 
ity conservation m the world. 

According to Rizzini (1963), the cerrado flora consists of an herbaceous 
and a woody component, which compete because both are heliophilous, i.e., 
there is no shade-adapted ground layer Following Coutinho's (1990) concept of 
cerrado, the importance of the herbaceous component decreases from open to 
closed physiognomies, whereas the importance of the woody component in- 
creases. In most forms of cerrado, the herbaceous component is much richer in 
species, as demonstrated by Mantovani and Martins (1993), who found a ratio 
between herbaceous and woody species ranging from 2:1 to 3:1 in the compari- 
son of some southern cerrado sites. However, despite its richness, Castro et al. 
(1999) pointed out the almost complete absence of existing surveys on the her- 
baceous component of the cerrado. These authors also highUghted the uneven 
distribution of surveys, which tend to be concentrated on few well studied ar- 
eas (Castro et al. 1999). 

The Emas National Park (ENP) is the largest and one of the most impor- 
tant reserves among those in the Cerrado Domain (Conservation International 
1999). Although some papers about its fauna and wildfires exist (e.g., Ramos- 
Neto & Pivello 2000; Rodrigues & Monteiro 2000), the ENP's vegetation remains 
poorly studied. The present work was designed as an intense and systematic 
floristic survey to provide a better knowledge of the ENP's flora in particular 
and of the cerrado flora in general, especially concentrating on the still poorly 
known herbaceous component. Furthermore, this survey is intended to pro- 
vide a basis for other studies to be carried out in the ENP and for phytogeo- 
graphical studies on the cerrado flora. 

To achieve these aims, we have tried to answer the following questions: What 
is the floristic composition of the cerrado physiognomies in ENP? Which fami- 
lies are the richest ones in its flora? Does the herbaceous to woody species ratio 
lie within the range described by Mantovani and Martins (1993), that is, be- 

'een 2:1 and 3:1? Are there woody species that should be included in the check- 
t elaborated by Castro et al. (1999)? 

Created m 1961, the ENP is located m the Brazilian Central Plateau, southwest^ 
ern Goias State (17°49'-1878'S, 5239'-5310'W), in the cerrado core region, and 
comprises 132,941 ha. Regional climate is humid tropical with wet summer and 
dry winter, classified as Aw following Koppen's (1948) system. Annual rainfall 
varies from 1,200 to 2,000 mm, concentrated from October to March, and mean 
annual temperature hes around 24,6°C (Ramos-Neto & Pivello 2000). Three 
quarters of the ENP consist of flat tableland, 820-888 m high, and the remain- 
ing area consists of hilly terrain, 720-820 m high (Ramos-Neto & Pivello 2000). 
Recently, ENP was included by UNESCO (2001) in the World Natural Heritage 
List as one of the sites containing flora, fauna, and key habitats that character- 

The cerrado in ENP exhibits almost all physiognomies found m this veg- 
etation type. Following Coutmho's (1990) classification and Sarmiento's (1984) 
translation, the cerrado in ENP ranges from campo limpo (a grassland) to cerrado 
sensu stricto (a woodland). In the reserve, open cerrado physiognomies— cam]90 
limpo, campo sujo (a shrub savanna), and campo cerrado (a savanna woodland)— 
prevail, covering 68.1% of the total area, especially on the flat tableland (Ra- 
mos-Neto & Pivello 2000). The more closed cerrado sensu stricto covers 25.1% 
of the reserve, mainly on the hilly terrain. Other vegetation types, such as wet 
campo (4.9% of the total area) and riparian and seasonal semideciduous forests 
(1.2%-)), also exist within the park. 

We surveyed all cerrado physiognomies occurring within the reserve from 
November 1998 to October 1999, m monthly field trips, each one with a 50-60 
hr sampling effort in the field. We established routes through the firebreaks 
that cross the reserve (Fig. 1) and covered one of them each day by driving a 
vehicle and stopping whenever vegetation of interest was seen. The routes com- 
prised the following reference points (with approximate distances): i) U2, Ul, Q, 
R, S, T U2 (51.5 km); ii) U2, V, PI, O, Ul (52.5 km); iii) O, M, N, PI (45 km); iv) y P2, 
W, Y, V (47.5 km); v) X, Zl, Z2, Z3, Z2, Zl, Y, W, X (47.5 km); vi) A, B, J, G, D, Q K2, 
Kl, A (48 km). 

We collected fertile botanical material along the pre-established routes. 
During the last two field trips, we also collected sterile material from species 
previously not found in reproductive stages. The collected material was identi- 
fied to species level by comparing it with lodged vouchers and consulting taxo- 
nomic references. The specimens were then sent to taxonomists for confirma- 


10'W),Goias State, central 


tion. The voucher material was stored in the herbaria of the Sao Paulo Botani- 
cal Institute (SP) and State University of Campinas (UEC). 

We classified the species in families according to the system proposed by 
Judd et al. (1999) and in life forms following Raunkiaer s (1934) system adapted 
by Mueller-Dombois and Ellenberg (1974). We considered the chamaephytes, 

epiphytes, hemicryptophytes, geophytes, lianas, vascular parasites, and 
therophytes as belonging to the herbaceous component and the phanerophytes 
as belonging to the woody component. We applied the chi-square test (Zar 1999) 
to verify whether the herbaceous to woody species ratio was significantly dif- 
ferent from the expected by Mantovani and Martins (1993). The results found 
were also compared to the patterns obtained by Castro et al. (1999). 

We collected 2,123 voucher specimens, representing 601 species, 303 genera, and 
80 families (Appendix I). Out of these 601 species, 571 were identified to species 
level, including one new to science, Piriqueta emasensis Arbo (Turneraceae). 
Six out of the remaining 30 species were also new to science and are currently 
being described by taxonomists: Annona sp. nov (Annonaceae), Gyrostelma sp. 
nov (Apocynaceae), Dimmerostema sp. nov (Asteraceae), Ipomoea sp. nov 
(Convolvulaceae), Hyhanthus sp. nov (Violaceae), and another Convolvulaceae 
that belongs to a new genus. There were 22 species identified to genus level, one 
identified to family level (Fabaceae sp.), and one that we could not identify even 
to family level. 

Twelve species (2.0% of the total number of species) were considered by 
Mendonga et al. (1998) as weeds that do not occur spontaneously in the cerrado. 
Of the 601 species, 149 were phanerophytes and thus included in the woody 
component, while 452 belonged to other life forms and were included in the 
herbaceous component. The herbaceous to woody species ratio was 3.03T, a 
value not significantly different (x^ = 0.014, P = 0.906) from the highest ratio 
(3:1) set by Mantovani and Martins (1993). 

The richest families were Asteraceae (88 species), Fabaceae (87), Poaceae 
(51), Myrtaceae (39), Lamiaceae (24), Malpighiaceae (23), Euphorbiaceae (20), 
Apocynaceae (19), Malvaceae (16), Rubiaceae (16), and Convolvulaceae (15), 
which together summed up 66.2% of the total number of species. In the herba- 
ceous component, the richest families were Asteraceae (85 species), Fabaceae 
(59), Poaceae (48), Lamiaceae (23), Euphorbiaceae (19), Malpighiaceae (16), 
Myrtaceae (16), Convolvulaceae (15), and Apocynaceae (14), which comprised 
65.27% of the herbaceous species. In the woody component, the richest families 
were Fabaceae (28), Myrtaceae (23), Malpighiaceae (7), Melastomataceae (7), 
Annonaceae (6), Apocynaceae (5), Vochysiaceae (5), Bignoniaceae (4), 
Nyctaginaceae (4), and Rubiaceae (4), which together accounted for 62.42% of 
the woody species. 

Seventeen out of the 149 woody species (11.4%) did not appear on the check- 
list of the cerrado woody flora established by Castro et al. (1999): Annona sp. 
nov, Aiouea trinervis Meisn., Apoclada arenicola McClure, Banisteriopsis acerosa 
(Nied.) B. Gates, Calliandra macrocalyx Harms, Dalhergia cuiahensis Benth., 
Mimosa amnis-atri Barneby, M. gemmulata Barneby, M. hehecarpa Benth., 

Myrcia hracteata O. Berg, M. camapuanensis N.F.E. Silveira, M. crassijolia (O. 
Kiaersk, Olyra taquara Sw., and Psidium laruotteanum Cambess. 

If we assume that the number of species in the cerrado ranges from 3,000 to 
7,000 (Castro et al, 1999), then the ENP contams approximately 8.5 to 20.0% of 
the cerrado flora. These figures show the importance of this reserve for the con- 
servation of the cerrado vegetation. The number of species in ENP might be 
increased by species not found in our survey Floristic surveys certainly miss a 
number of species in a given area, especially those that are not at reproductive 
stage at the time of the visit, flower sporadicaly are ephemeral, or are incon- 
spicuous, problems that particularly affect the herbaceous component of the 
vegetation (Mantovani & Martins 1993; Castro et al. 1999). 

Among the 601 species found in the ENP, seven are new to science. Although 
the cerrado is one of the most studied vegetation types in Brazil (Castro et al. 
1999), the fact that undescribed species keep on appearing in floristic surveys- 
for example, also in Brasilia, Federal District (Pereira et al. 1993)-shows that 
the cerrado has not yet been satisfactorily sampled. The affirmation that the 
tropical flora remains undercollected (Prance et al. 2000) seems to apply also 
to the cerrado vegetation. Some of the species not identified to species level in 
our inventory may be new to science as well. 

Although the proportion of weedy species in the ENP's cerrado flora was 
lower than the ca. 5% found by Mendonga et al. (1998) for the whole Cerrado 
Domain, the invasion of ruderal plants in ENP is alarming, notably the African 
grasses BrachicznadecumbensStapf and Melinis minutiflora P. Beauv Plant in- 
vasion has become a great problem in virtually all cerrado fragments (Pivello 
et al. 1999) and will grow into a serious problem in the ENP if precautions are 

The richest families in the ENP were also the richest ones m other cerrado 
sites (Mantovani & Martins 1993; Batalha et al. 1997; Batalha & Mantovani 2000). 
Exceptions were Convolvulaceae and Lamiaceae, well represented only m the 
ENP The herbaceous to woody species ratio, although not significantly differ- 
ent from the maximum set by Mantovani and Martins (1993), was higher than 
the ratios found m other surveys (Mantovani & Martins 1993; Batalha et al. 1997; 
Batalha & Mantovani 2000). This probably was a consequence of the preva- 
lence of open physiognomies Uampo limpo and campo sujo) in ENP 

According to Castro et al. (1999). surx'cys m poorly sampled regions should 
augment the cerrado woody species checklist. Indeed, the 17 species found in 
our survey that should be included m their list represent a high percentage of 
the species collected m ENP and indicate that even the woody component of 
the cerrado vegetation remains undercollected. As for the herbaceous compo- 

iilable for only aboi 
survey of tlie dom 
;tep, and it will be nece 

campo, and the aquatic ve; 

nent, this undercoUection might be even greater I 
baria and in our search for references, the absence o: 
became evident. 

Species inventories, even at the most basic level 
1% of the tropical regions (Hammond 1992). Our f 
nant cerrado vegetation in the ENP represents a fir 
sary to survey the other vegetation types existing with 
seasonal semideciduous forest, riparian forest 
etation of streams, rivers, and lakes. 

Species collected in the floristic survey of the cerrado vegetation in Emas Na- 
tional Park (17'49'-18°28'S, 5239'-5310'W), Goias State, central Brazil. Taxa were 
listed alphabetically by family, genus, and species. The format used was species 
name, authority, collection number, and life form. Authorities were abbreviated 
according to Brummit and Powell (1992). Species designated by an asterisk (*) 
were considered non-native weeds by Mendonga et al. (1998). All specimens were 
collected by M.A. Batalha (B). Life form was assigned according to Raunkiaer's 
(1934) system adapted by Muller-Dombois and Ellenberg (1974). Life form 
classes were abbreviated as: Ch = chamaephyte, Ep = epiphyte, Geo = geophyte, 
H = hemicryptophyte, Li = Liana, Ph = phanerophyte, Th = therophyte, and Vp 
= vascular parasite. 

The vascular flora of the cerrado in Emas National Park (central Brazil). 
ACANTHACEAE Annona monticola Mart., B 3620, H 


Alstroemeriagardneri Ba^ 

Froelichiaprocera (Seub.) 
Gomphrena arborescens 


Pfaffio h 




Aspidosperma tomentosum Mart., B 4028, P 
Barjonia cymosa Fourn,, B 2897, H 
Barjonia erecto (Veil.) K. Schum., B 2182, H 
Blepharodon bicuspidotum Fourn., B 245 1 , 1 
Gyrostelma sp. nov., B 2081 , H 
Hancornia speciosa Gomez, B 2 1 06, Ph 
Hemipogon acerosus Decne., B 2083, H 

/ Baker, B 303 l,Th 

(ano Baker, B 3771, H 


ep/5 Baker, B 2886, H 

s Sch. Bip. ex Baker, B; 

5204, H 

m chlorolepis Baker, B 


m megacephalum {M. 

art.) King & 

;err/mo (Veil.) Burkart, 

B 2089, Th 

r/^aeseoe H.Rob., B 33- 


eucocepha/a Gardner, 


jalida {A.DO King &H.Rob., E 
chyop/iy//a(Spr.) King & H.Rob. 

engelianum (Gardner) Cabrera 

'sperafum Blake, B 2592, H 
>ras/7/ontym Cass., B 3161, H 
't//o//um (Sch. Bip.)Blake,B 2246 



Elephantopus biflorus Less., B 31 59, H 
Elephantopus mollis L, B 31 28, H 
f/ephantopusracemosus Gardner, B 34 

Acrocom/a /ioss/er/(Barb.Rodr.)Hahn,B3828,Geo 
Allagoptera campestris (Mart.) Kuntze, B 2006, 

Allagoptera leucocalyx {Man.) Kuntze.B 221 5,Geo 
Attalea geraensis Barb. Rodr., B 2509, Geo 
Syagrus flexuosa (Mart.) Becc, B 2249, Geo 

Eremanthus erythropappus Sch. Bip., B : 
Eremanthus glomerulatus Less., B 3333, 
Eremanthus sphaerocephalus Baker, B 2 


Eupatorlumlanigerum hook.&Arn..B-: 

s Sch. Bip., B 3093, 1- 

jreo/des (Lam.) A. DC, B 3400, T 
(Spreng.) Baker, B 3377, H 

Tabebula aurea (Silva Manso) S. 
'turn (L) A. DC, B 31 55, H Ph 

ig//o//a Gardner, B 2331, H Tabebuia ochracea {Cham.) Stan^ 

a Chod., B 1 900, H BORAGINACEAE 

edosus Baker, B 2504, H oriaviicauis resen., B 381 1,1 

-tvoohvllum (A DC) Kina & H BROMELIACEAE 

Vemonia buddleiifolia Sch. Bip. ex Baker, B 2338, H Dickia tuberosa (Veil.) Beer, B 2839, H 

Vernonia compactiflora Mart. ex Baker, B 2867, H BURSERACEAE 

Vemonia desertorum Mart, ex A. DC, B 3567, H Pmtium ovatum Engl., B 1 930, Ph 

ZZ7a fel^'^LeTB 35?0°r ' CACTACEAE 

Vemonia grandiflora Less., B 2487, H Epiphyllum phyllanthus (L.) Haw., B 3878, Ep 

Vemonia herbacea (Veil.) Rusby, B 1 992, H CARYOCARACEAE 

Vemonia polyanthes (Spreng.) Less., B 2336, C 

Vemonia rubrirameaMan.,B3]9],Ch ' CELASTRACEAE 

Vemonia simplex Less., B 4014, H ' Plencl<ia populnea Reissek, B 2277, Ph 

Vemonia tomentella Mart, ex A. DC, B 2063, H Tontelea miaantha (Mart.) A.C Sm., B 1 934, Ph 

Vemonia tragiaefolia A. DC, B 2 1 72, H CHRYSOBALANACEAE 

Vemonia varroniifolia A. DC, B 3304, H Couepia grandiflora (Mart. & Zucc.) Benth. ex 

Vemonia virgulata Mart., B 2033, H Licania humilis Cham.& Schltdl., B 3034, Ph 

Viguiera sp.2, B 1960, H 

Wedella macedoi H. Rob., B 3920, H n,c„Mcyc,u u^unu .o^^,, u ^^.., ^, , 

BALANOPHORACEAE Kielmeyera grandiflora (Wawra) Saddi, B 2061 , Ph 

Langsdorffia hypogea Mart., B 4030, VP Kielmeyera rubriflora Cambess., B 3022, Ph 

BIGNONIACEAE Kielmeyera (richophora Mart., B 2035, Ch 


Anemopaegmaglaucum Mart.ex A. DC, B 3035, H Cochlospermum regium (Mart.) Pilg., B 2428, H 

^"'^^g^Sn'^^ ^'^'^^^'^^'^"''"^ ^^^^' '''' ^' ^^' ^ COMBRETACEAE 

Arrab'daea brarhvnodn (A DC ) R R 2?72 C\ Buchenavia tomentosa Eichl., B 4031, Ph 


Ipomoeasp nov,B2357,H 

Croton glandulosus Mull. Arg., B 2389, H 

Ipomoea virgata Meisn , B 341 5, H 

Croton goyazensis Mull. Arg., B 2222, H 

Jacquemontia guaranitica HassI , B 2636, H 

Jacquemontia sphaerocephala Meisn , B 2363, H 

Croton pohlianus Mull. Arg.! B 21 96^ H 

Merremia contorquens (Choisy) Hall f , B 2600, Li 

Croton sc/eroca/yx Mull. Arg., B 2885, H 

Merremia digitata Meisn , B 2140, Li 

Crofon sp.,Bl 964, H 

Turbina abutiloides (Kunth) O'Donnell, B 25 1 2, H 

Dalecharvpia humilis Mull. Arg., B 1 981 , H 

Convolvulaceae sp. 1 , gen. et sp. nov., B 1 967, H 

Dalecharvpia linearis BailL, B 2076, H 


Julocroton humilis Didr., B 2207, H 

Ceratosanthes hilariana Cogn., B 3645, Li 


Bulbostylisjunciformis (Kunth) C.B.CIarke,B 2300, 

Manihot tripartita (Spreng.) Mull. Arg., B 1908, 
Maprounea guianensis AubL, B 3930, Ph 
Phyllanthus orbiculatus Mull. Arg., B 2445,Th 
Sapium glandulatum (Veil.) Pax, B 2301 , Ch 
Sebastiania bidentata (Mart.) Pax, B 2430, H 

Bulbostylis paradoxa (Sprang.) Lindm., B 3565, H 
Bulbostylis sphaerocephala (Nees) C.B. Clarke, B 


Bulbostylis truncata (Nees) M.T. Strong, B 3971 , H 
Cyperus aggregatus (Willd.) EndL, B 2023, H 

Aeschynomene marginata Benth., B 271 6, Th 
/1nodenonr/ierafo/cota(Benth.)Speg.,B 2650,1 

Kyllinga odorata Vahl, B 293 1 , H 

^nfo ZtdialTntKB llTl'cl^ 

Rhynchosphora diamantina (C.B.Clarke) Kukenth, 

Andira laurifolia Benth., B 2342, Ch 

Andira vermifuga (Mart.) Benth., B 3718 

Rhynchosphora emaciata Boeck., B 3087, H 

Arachis tuberosa Bong, ex Benth., B 227 

Rhynchosphora exaltata Kunth, B 2466, H 

Rhynchosphora rugosa (Vahl) Gale, B 3534, H 

Bowdichia virgilioldes Kunth, B 3350, Ph 

Calliandra dysarnha Benth., B 1 979, Ch 


Calopogonium sericeum (Benth.) Ch 

Davtila nitida (Vahl) Kubitzki, B 3988, Li 

Camptosema ellipticum (Desv.) Burkart, 1 


Centrosema venosum Mart, ex Benth., B 

Dioscorea amaranthoides PresI, B 31 24, Li 

Chamaecnsta basifolia (Vogel) Irwin & 

MOOl, Mimosa n 

Chamaecrista desvauxii (Collad.) Killip, 2457, Ch 
Chamaecrista filicifolia (Benth.) Irwin & Barneby, 

Chamaecrista flexuosa (L) Greene, B 2290, Ch 

Dalbergio cuiabensis Benth., B 3214, Ph 
Dalbergia miscolobium Benth., B 3504, Ph 

Desmodium incanum (Sw.) A. DC, B 2523,Th 
Desmodium platycarpum Benth., B 3944, H 
Dimorphandra mollis Benth., B 2047, Ph 
Dioclea bicolor Benth., B 2042, Li 
Diptychandra aurantiaca Tul., B 3056, Ph 
Eriosema crinitum (Kunth) Gardner, B 2483, h 


leterophyllum Benth., B 3992, l- 
3ng//o//um Benth., B 21 94, H 

Fabaceaesp. l,B2796,Pf 

Caseana grandi flora Cam 

Galactia dimorpha Burk., B 21 30, H 
Harpalyce brasiliana Benth., B 2548, P 
Hymenaea stigonocarpa Mart., B 232( 
Indigofera gracilis Bong., B 2057, H 
Lupinus subsessiiis Benth., B 2320, H 
Machaerium acutifollum Vogel, B 21 1 





is capriariifolia Pohl ex Benth., B 3006, H 
is caudata Epiing & Sativa, B 3643, H 

Byrsonima crassa N 
Byrsonima gauithei 
Byrsonima guillemn 
Byrsonima intermec 
Byrsonima ngida A 
Byrsonima verbasal 


Juss,B2135,Ch ' 

/s enop/iy//a Pohl, B 2849, H 

CanZ. ■ ^^ 


is intenupta Pohl ex Benth., B 3076, H 

Hetc; •! ' 

' ' ''"''-' 

is lytiiroides Pohl ex Benth., B 3708, H 

Heo ■ ■: 

1 i476,f 

;5 multiflora Pohl ex Benth., B 2670, H 


1 2, Li 



' 111 '■ ."-'ai 

'is tZsa Po^hl ex^Bemh^'^B 2255 ^H ^ ^ ' ^^ 



is virgata Benth., B 1 925, H 


is sp., B 3544, H 

Byttneria obiongatc 


Marsypianthes montana Benth., B 3 1 08, H 



3336, Ph 

Pe/fodonpu5/7/us Pohl, B 3271, H 

Helicteressacaroiha A.St-H\ 

Peltodon tomentosus Pohl, B 3065, H 



Meloctiiavillosa {Mill.) f awe 


Pavonia rosa-campestris A.'. 

Peltaea edouardii (Hochr.) \< 

Cassyttia filiformis L, B 2422, VP 




Eschweilera nana (0. Berg) Miers, B 2825, Ph 



Suyclnnos pseudoguina A, St-HIL, B 3943, Ph 

Sida cerradoensis Krapov.,B 

Cupiiea carthagenensis (Jacq.) Macbr,, B 21 79, H 

Sidl rhomMolial *"b 2840 

Cupliea linarioides Koehne, B 21 36, H 

Waltheria douradinha A St 

Lofoensia pacari A. St-Hil., B 3782, Ph 

Waltheria indica L * B 2274 


Banisteriopsis acerosa (Nied.) B.Gates, B 2809, Ph 

Mcon,a fo//ax A. DC., B198f 
Miconia ferruginata A. DC., E 

Banisteriopsis amplectens B. Gates, B 2429, Li 

Banisteriopsis campestris (A. Juss.) Little, B 2346, Li 

Banisteriopsis gardneriana (A. Juss.) W. Anderson 

Miconia ligustroides (A. DC.) 

Rhynchanthera ursina Naud., B 3536 


Psidium cinereum Mart., B 221 2, Ch 

Tibouchma gracilis (Bon pi.) Cogn., B 


Psidlum firmum 0. Berg,' B 3848^ Ch 

Tibouchina stenocarpa (A. DC.) Cogr 


Psidium laruotteanum Cambess., B 2250, P\ 


Psidium multiflorum Cambess., B 2547, Ch 
Psidium rufum Mart, ex A. DC, B 21 99, Ph 

Cissampelos ovali folia Ruiz & Pav„ B 





Guapira campestris (Netto) Lund., B 3969, C 
Guapira gracilifora (Mart, ex J.A. Schmidt) L 


Guap/ranox/a (Netto) Lund., B 3552, Ph 


Myrsine leuconeura Mart,, B 2820, Ph 

Neea macrophylla Poepp. & EndL, B 3080, P 



Campomanesia adamantium (Camb 

ess.) 0. Berg, 

Ouratea acuminata (A. DC.) EngL, B 2463, P 
Ouratea castaneaefolia (A. DC.) EngL, B 357 

Eugenia aurata 0. Berg, B 2699, Ph 
Eugenia bimarglnata A. DC, B 3502, Ph 
Eugenia calyclna Cambess., B 261 8, Ch 
Eugenia complicato 0. Berg, B 3583, Ch 
Eugenia cristaensis O. Berg, B 223 1 , H 


Eugenia piloesls Cambess., I 


Habenaria obtusa LindL, B 2633, Geo 

Oxalis sellowii Spreng., B 1 91 0, H 

Mitostemma brevifilis Gontsch., B 3562 
Passiflora mansoi (Mart.) Mast., B 3066, 

crassifolia (0. Berg) Kia. 
decrescens 0. Berg, B 1 ' 
Max (Rich.) A. DC, B 3 

\ndropogon fastigiatus Sw., B 
\ndropogon leucostachys Kui 
\ndropogon selloanus (Hack.; 




m lanatum (Kunth) Nees, B 3700, H 
hrysothryx (Nees) Conert, B 2845, h 


Crumenaria polygaloides Reissek, 

Paspalum convexum Hunnb.& BonpI.ex Fleug.,B 

D/od/o feres Walt., B 2524, Th 

Ga//onrhe grand/fo/Zo E.L. Cabral, B 2944, H 

Paspalum geminiflorum Steud., B 3250, H 
Paspalum malacophyllum Trin., B 3050, H 

PalicoureaconaceaiCham.) K.Schum.,B 1903,C 
Pallcourea ngida Kunth, B 1 941 , Ph 

Z7dZ ^^iZ^ue:!, bim^m 

/^JaT2're//ans (Cham. &Schltdl.) Steud., 

Paspalum sp.],B3060,H 
Paspalum sp.2,B2938,H 

Sipanea hispida Benth., B 2485, H 

fihync/ie/yfmm repens (Nees) C.E.Hubb/,B 





Schizachyrium condensatum (Kunth) N( 

?es, B 

Honia brasiliensis Vand. ex A. DC, B 2765, Ph 

3430, H 

Spiranthera odoratissima A. St-Hil., B 2073, Ch 

Setoria geniculata (L.) P. Beau v.*, B 2495, T^ 
Sporobolusacuminatus Boechat& Longhi- 


Matayba guianensis AubL, B 3925, Ph 

Sporobolus ciliatus (Trin.) Hack., B 2986, H 
Sporobolus tenuissimus (Schrank) Kuntze.B 




Tristachya leiostachya Nees, B 2269, H 

Polygala angulata A. DC, B 2425, H 
Polygala aphylla A.W. Benn., B 391 0, H 
Po/yga/a op/no Wurdack, B 2223, H 
Polygala violacea AubL, B 2492, H 


Pouteria ramiflora (Mart.) Radlk., B 1 984, Ph 
Pourer/fl subcaeru/ea Pierre ex Dubard,Bl 899, C 


a 5chauer, B 241 5, Ch 


Simaba suffruticosa Engl., B 3730, 
Simarouba amara Aubl., B 3802, F 


Lippia lupulina Cham., B 

3642, Ch 

Lippia mc 

mulina S.Mooi' 

■e, B 3966, H 

Lippia sta 

chyoides Chan 


nerifolia Cham 


-pheta maximih 

'/an/ Schauer, 


pheta simplex \ 

Hayek, B 2369 


Styrax ferrugineum Nees & M< 

Piriqueta sidi folia (Cambess.) 

We are grateful to Fapesp, for financial support; to Ibama, for granting the re- 
search permit; to the Emas National Park staff, for their help in field; to M.B. 
Ramos-Neto, for logistical assistance; to M.F.R. Siles, for translating the Abstract 
into Spanish; to W. Mantovani, for useful discussions and assistance in species 
identification; to the taxonomists K. Althott, M. Alves, M.M. Arbo, M.C. Assis, C. 
Bestetti, R.S. Bianchini, V. Bittrich, A.C. Cervi, I. Cordeiro, C.L. Cristobal, T.S.C. 
Dorow, S.I. Elias, G. Esteves, M.A. Farinaccio, M.S. Ferrucci, R Fiaschi, E. Forero, 
A. Furlan, E. Gongalves, M.L. Kawasaki, M. Kirizawa, A. Krapovickas, J.A. 
Lombardi, M. Magenta, A.C. Marcato, R. Mello-Silva, E. Melo, M. Moraes, R.C. 
Oliveira, M.B. Paciencia, J.R. Pirani, A.P Prata, C. Proenga, N.A.T. Ranga, ER. 
Sahmena, J.U.M. dos Santos, J. Semir, M.F Simon, E.R.V. Silva, M.M. da Silva, J.P 
Souza, A. Spina, J.Y. Tamashiro, W. Thomas, R.B. Torres, M.G. Wanderley, and A. 
Zanin, for their assistance in the identification of species; and to J. Ratter and 
W. Thomas, for their useful comments on the text. 

SP). Bol. Bot. Univ. Sao Paulo 1 6:49-64. 

(Santa Rita do Passa Quatro, SR Brazil): a com- 
the woody floras. Rev. Bras. Biol. 60:1 29-145. 
Brummitt, R.K. and C.E. Powell (eds.). 1 992. Autliors of plant names. Royal Botanic Gardens, 

Castro,A.A.J.F., F.R.Martins, J.Y.Tamashiro, and G.J. Shepherd. 1999. How rich is the flora of the 
Brazilian cerrados? Ann. Missouri Bot. Gard. 86:1 92-224. 

Conservation International. 1999. A(;6es prioritarias para a conservagao do Cerrado e do 
Pantanal. Conservation International, Brasilia. 

Coutinho, L.M. 1 990. Fire in the ecology of the Brazilian cerrado. In: J.G.Goldannmer,ed. Fire 
in the tropical biota. Springer, Berlin. Pp. 82-1 05. 

FoNSECA,G.A.B.DA,R.A.MiTTERMEiER,R.B.CAVALCANTi,andC.G.MinERMEiER.2000. Brazilian Cerrado. 
In: R.A.Mittermeier, N.Myers, P.R.Gil, and C.G.Mittermeier, eds. Hotspots, Earth's biologi- 
cally richest and most endangered terrestrial ecoregions. Conservation International, 
Chicago. Pp. 148-159. 

Hammond, RM. 1 992. Species inventory. In B. Groombridge, ed. Global biodiversity. Chap- 
man and Hall, London. Pp. 1 7-39. 

JuDD, W.S., C.S. Campbell, E.A. Kellog, and RF. Stevens. 1 999. Plant systematics: a phylogenetic 

.0PPEN,W. 1948.Climatologia. Fondo de Cultura Economica, Mexico. 

enthall, J.C, S. Bridgewater, and RA. Furley. 1 999. A phytogeographic analysis of the woody 

elements of the New World savannas. Edinburgh J. Bot. 56:293-305. 
Iantovani, W. and FR. Martins. 1993. Floristica do cerrado na Reserva Biologica de Moji 

Guagu, SR Acta Bot. Brasil. 7:33-60. 
^endonca, R.C. de, J.M. Felfili, B.M.T. Walter, M.C. da Silva Junior, A.V. Rezende,T.S. Filgueiras, and 
RE. Nogueira. 1998. A flora vascular do cerrado. ln:S.M.Sano and S.R Almeida, eds.Cerrado: 
-id methods of vegetation ecology.John 

^.S., M.A. Silva, and R.C. Mendonca. 1 993. Reserva Ecologica do IBGE, Brasilia— DF: 
? plantas vasculares. IBGE, Rio de Janeiro. 
1,V.M.C.Carvalho, RRLopES,A.A. pEcciNiNi,and S. Rosso. 

Biotropica 31:71-82. 
Prance, G.T., H. Beentje, J. Dransfield, and R. Johns. 2000. The tropica 

undercollected. Ann. Missouri Bot. Gard. 87:67-71 . 
RAMOs-NETo,M.B.and V.R. Pivello. 2000. Lightning fires in a Brazilian savant 

rethinking management strategies. Environ. Managem. 26:675-684. 
Ratter, J. A., J. R RiBEiR0,and S. Bridgewater. 1 997.The Brazilian cerrado vegeta 

to its biodiversity. Ann. Bot. 80:223-230. 
Raunkiaer.C. 1 934.The life forms of plants and statistical plant geography.CI 
RizziNi, C.T. 1 963. A flora do cerrado. In: M.G. Ferri, ed. I Simposio sobre o 

Blucher and Edusp, Sao Paulo. Pp. 1 25-1 78. 

RoijRiuijFh, F.H.G. and El. A. Munhiko. 2000. Home range and activity patterns of pamp 

deer in Emas National Park, Brazil.J.Mannmai. 81:1 136-1 142. 
Sarmiento, G. 1 983. The savannas of tropical America. In: Goodall, D.W., ed. Ecosystems 

the world - tropical savannas. Elsevier, Amsterdam. Pp. 245-288. 
Sarmiento, G. 1984. The ecology of Neotropical savannas. Harvard University, Cambridgi 
UNESCO. United Nations Educational, Scientific and Culturae Organization. 2001 . Cerrado pr 

tected areas: Chapada dos Veadeiros and Emas National Parks. Paris: Unesco. http 
Zar.J.H. 1999. Biostatiscal analysis. Prentice Hall, Upper Saddle River. 

4 M, Brodo, Sylvia Duran Siiarnoff, and Sthphfn SllARNOlu^ 2001. Lichens of 
North America. (ISBN 0-300-08249-5, hbk.). Yale University Press, 302 Temple 
Street, P.O. Box 209040, New Haven, CT 06520-9040, U.S.A. (Orders: 203- 
432-0960, 203-432-0948 fax). $69.95, 795 pp, 924 color photos, distribu- 
tion maps, 9 1/2" x 11". 

parts. The first part begin: 

5 with i 

ntroductorv material inc 



iderful description of the sym- 

biotic relationships that 1 


form and moN'cs on to d 


;s luhci 

1 morphology, anatomy, repro- 

duction, physiology, chemistry, t 

ecology and bmgcograph 

y Tu 


crs describe how humans have 

chapters offering very pr. 


information on how to d- 

be and c 

lassify lichens and how to col- 

lect and preserve them foi 

information about how tc 

e maps, keys and descrij 


sin the 

second half of the book. 

The second part of t 

his vol 

ume provides easy to fol 


;eys. we 

Il-crafted descriptions, beauti- 

ful photographs, and ver) 

! usefu 

1 range maps for 805 spe> 

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is. The photographs and range 

professionals or avocatior 

lal entf 

lusiasts. A comprehensi^ 



.the back of the book provides 

a most welcome guide to 

people know very little. 


lis book is extremely wel 



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:xt tim 



xcal Research Institute oj Texas, 

Jose R. Hernandez Joe F Hennen 

Systematic Botany and Mycology Laboratory Botanical Research Institute of Texas 

USDA/AR5, Rm. 304, BO I lA 509 Pecan Street 

Beltsville, MD 20705-2350, U.S.A. Fort Worth, TX 76 1 02-4060, U.S.A 

En el presente trabajo se presentan los resultados de la prospeccion de royas (Uredinales) realizada 

pertenecientes a la region conocida como Noroeste Argentine. Como resultado de este trabajo se 
elaboraron 635 colecciones correspondientes a 51 familias de hospedantes (257 especies). Ochenta y 
cuatro de estas colecciones son nuevos hospedantes para Argentina, de las cuales 50 son nuevas citas 
como hospedantes para royas. Se identificaron 30 generos y 182 especies de royas, de las cuales 61 son 
nuevos registros para el pals y 10 nuevas especies. Las colecciones fueron depositadas en Fundacion 
Miguel Lillo (LID y U.S. National Fungus Collection (BPl). 

The fungi of Argentina have been studied by a number of mycologists, most 
notably Carlos Luigi Spegazzim (1858-1926) whose taxa were compiled into a 
comprehensive account by Farr (1973). Spegazzini's collections of rust fungi 
(Uredmales) as well as collections made by later mycologists were organized 
and studied by Lindquist (1982). Lindquist s studies resulted in numerous pub- 
lications on rust fungi, culminating in his "Royas de la Republica Argentina y 
Zonas Limitrofes" (Lindquist 1982). Although the fungi of Argentina are rela- 
tively well known, the rust fungi of Northwest Argentina are less well studied 
because few mycologists collected rusts in that region. 

In this paper we present a study of rust fungi collected on wild and culti- 
vated plants in the provinces of Catamarca, Jujuy, Salta and Tucuman in North- 
west Argentina. This subtropical region is characterized by climatic and bio- 
logical diversity and consists of a plain in the east and the Cordillera de los 
Andes in the west where valleys closely alternate with mountains. The grada- 
tions in latitude and altitude in Northwest Argentina result in a mosaic of 
bioecosystems with vegetation types varying from desert to rain forest. The di- 
versity of flora is reflected in the range of phytogeographical regions described 

for this area by Cabrera (1994). For these reasons, Northwest Argentina is an 
interesting and rich area for collecting rust fungi. 

Lindquist (1982) documented 450 species representing 28 genera of rust 
fungi from throughout Argentina of which 124 species were reported from 
Northwest Argentina. Between 1993 and 1999 the authors collected and identi- 
fied or re-examined 635 specimens of rust fungi representing 182 species in 30 
genera. These rust specimens were collected on 257 host plant species from 51 
plant families. Eighty-four of these host plants are new host records for Argen- 
tina, and 50 of these are new host records for rusts worldwide. Sixty-one rust 
species are reported for the first time from Argentina and 10 of these are new 
species. One of those, Baeodromus ranunculi].R. Hern. &J.F. Hennen, has been 
described (Hernandez 2000) and the remainder will be described elsewhere. 

The rust species reported here are presented according to host family This is 
the most useful form of organization because of the high level of host specific- 
ity of the rust fungi. Collections for each rust species are then listed alphabeti- 
cally by host species followed by the province, details of locality, date, collector(s) 
and collector's number, herbarium number and rust stage present. For the rust 
stages symbols are used as follows: = spermogonial stage, 1 = aecial stage, 11 = 
uredinial stage, and III = telial stage. The abbreviation "An" refers to the pres- 
ence of an anamorph stage for which we are uncertain whether it is function- 
ally an aecial or uredinial stage. Symbols representing spore types are X = 
amphispores, M = mesospores, and IV = basidiospores. 

New rust records are indicated in superscript following the species name 
as follows: new rust species (N), new rust record for Argentina (A), Catamarca 
(C), Jujuy (J), Salta (S), and Tucuman (T). Superscript notations following host 
names indicate: new host plant genus for this rust worldwide (G), new host plant 
species for this rust worldwide (S), new host plant species for any rust in Argen- 
tina (A), and new host plant species for which another rust was previously 
known from Argentina (AR). 

Abbreviations of authorities for plant host and fungal names follow 
Brummit and Powell (1992) and Kirk and Ansell (1992) respectively Plant host 
names were verified using W3TR0P1C0S at the Missouri Botanical Garden. 

Ill]; Las Salas, 5 May l995J.R.Heniandez 

24 Sep 1997,J.EHennen &J.R.Herndndez 

95-015 (BPI 841079) [MMII]; San Javier, 

Parque Sierras de San Javier, 23 Jul 1QQ4, 

ing area, 29 Nov 1997 J.R.Herndndez 97- 


1 1 7 (BPI) [11]; W of Catamarca, 23 Sep 1997, 


j.EHenncn & J.R.Herndndez 97-043 (BPI) 

IF.Henncn&MM.Henncn 94-07 nUL 

[III; W side of Catamarca, 23 Sep 1997, 

54703, BPI) 101. 

J.F.Hennen & J.R.Herndndez 97-035 (BPI 

841184) [II]. Salta: Dept. Santa Victoria, La 


Mision, 30 Jun 1996 J.R.Herndndez 96- 

E of La Vina, 24 Sep 1997, J.F.Hennen & 

lR.Herndndez 97-061 (BPI 841197) [11]. 

j.R.Herndndez 94-149iUL 54943, BPI) [II]! 


on Hydrocotyk sp.; Catamarca: El Rodeo, 29 

usticia sp.; Tucuman: San Javier, Parque 

Nov 1997, J.R.Herndndez 97-121 (BPI) [II]. 

Tucuman: Amberes, 5 May 1989, 

l.R.Herndndez 94-H4 (LIL 54705, BPI 


841067) [11-111], 

Dept. Trancas, Rt. 9, km 1364, 26 Oct 1995, 

es sp.^ 

J.R.Herndndez 95-052 (BPI) [111; Sanjavier, 

leo I4No\ 1999 J R Hernandez 99- 
' (BPI 841276) [0 II Salta: Dept. 

an Dept JRHcinandLz97 167 (BPI) [I| Tucumar 

on HydrocotylehonanensisC 

El Rodeo campi 

de San Ja\ lei 

27 M 

n 1993 




93 035 i 

1 Seneco cfcmc./lr;rus Mattr Catamarca 

m 19 Ap, 1926 Wentun 4138 [III] 

Dept AmbHos El Rodeo 14 No^ 19Q9 

ljunl933 A L Cflbre-a 3024 [III] 

of Catamarca Hwy 38 24 Sep 1997 


JFHennen &] R Hernandez 97 055 (BPI 


del Valle 6 May 1995 J R Hernandez 95 

I Senecio deferens Giiseh Tucuman Sier 

078 (BPI 841106) [III] Tafi del Valle, 6 Apr 

ras de San Javier 17 Nov 1949 Smge> & 

1993 JhHtnnen&JRHe,nandez93-J23 


iSeneaootopterusGnseh Catamarca El 

Puccinia conochnii Se> m in Burrill 

Rodeo 29 Nov 1997 J R Hernandez 97 

126 (BPI 841231) [II] 

I Senecio rudhttkiijobus Meyen & Wilp 

Siern 11 Mir 1969 A Krapovickas et al. 

T„cun.a„ Taf> del Valle 6 Apr 1995 

in [II] Dept Burruyacu Cerro del 

Cimpo ele\ 1500 m 29 Mar 1930 

Valle 6 Apr 1993 JFHLnntn & 


onEupatonum^p Salta Dept Rosariodela 

f 168 (BPI 841075) [III Apr 1994 

]RHt,n,ul HIIKIU ^4 H Ml on 1 k/ u nun, sp Salta Calayate, 8 Apr 

841046)1111111] Uui ini nntn MMHennen&].R. 

Salta: Dept. Santa Victoria, Los Toldos, El 
Nogalal, 29 Jun 1996 J.R.Herndndez 96- 
048 (BPl 841137) [l-lll]. 
lia proluviosa H.S.Jacks. & Holw.'^ 
I Senecio peregrinus Griseb.^; Catamarca: E 



f Catamarca, 23 Sep 1997 J. F.Hennen & 
RBerndndez 97-045 (BPl 841189) [II]. 





J.F.Hennen & J.R.Herndndez i 



84n68) [11]; San Javier, Parque 


lia hieracii(Rohl.)H. Mart. 

i62(LlL 54746, BPl) [11]; San Ja^ 

Taraxacum officinale Weber; Cai 


Sierras de San Javier, 27 

Dept. Ambatos, El Rodeo, 14 N 

ov 1999, 

J.F.Hennen, LD.PIoper6' J.R.J 

J.R.Herndndez 99-003 (BPl 8412- 

72) [111. 


Puccinia rata H.S. Jacks. &Holw.'^ 

Achyrocline sp.; Salta: Dept. Sant 

:a Victo- 

on Vernonia saltensis Hieron.''; Sal 



Cerro San Bernardo, 27 1 

96-052 (BPl 841140) [11]. 

A.LCabrera 3021 [Il-lIIl Tucu 

lia lilloanaJ.C. Lmdq.^ 


Baccharis lilloi Heenng; Tucumar 
Apr 1904, leg. HlIIl. 


Screiter 47843 [MMII]; Sar 
Colalao, 1 Apr 1994, J.F.I 


s ; Jujuy: 


Qumta prop. Laguna de la Brea 

, 31 May 

Puccinla schileana Speg var parthen 

1901, R.E.Frias s.n. [11]. Tucumai 

Jacks. &Ho]w.)J.C.Lindq. 

Molle, 17 Sep 1997, J.R.Herndnde 


on Parthemum hysterophorus L.; 

(BPl) [II-llI]; Dept. Cruz Alta, J 

M.M.Hennen &J.R.Herndn 
(LIL 54743, BPl) [11-111]; Sa 
Colalao, 2 Apr 1994, J.F. 

097 (BPl 841218) [II]. 

Tucuman J B Alberdi S of Fbcabi Di 
22 Sep 1997 ]FHenntn&]R Ha nam 
97 on (BPI 841169) [11] 

1 Vernoma squamosa Gardner Catamarca 
Balcozna 22 Sep 1997 JbHennen & 
]RHanandLz97 022 (BPI 841175) [II III] 

nf-JR Hernandez 93 

93-0K(LIL 54764, BPI)[IHII]; 
road to Escaba, 28 Nov 
Hernandez 97-096 (BPI 8412 
:es megalospermus Speg. 

Hernandez 97-115 (BPI 841227) [II]; ^ 
side of Catamarca, 23 Sep 199 
J FHennen &J R He> mindez 97-024 (B 

E of La Vina, 24 Sep 1997, 


^J.R.Herndndez 97-060 (BPI) 

[1]. Jujuy: Caimancito, 2 Apr 1993, 

& L.D.Ploper 93-110 (LIL 

54714, BPI 

) [I-ID; Parque Nacional 


Decl997, J.R.Herndndce 97- 

268) [11; Santa Clara, 12 Apr 


nen<S'LD.PIoper94-J3? (LIL 

III Saka: Dept. Santa Victo- 


i,30Jun 1996, J.R^Herndndez 

Martin, 7 Dei 

:1997.J,R.Herndnde2 97-J72 

(BPI 841262 

15 Nov 1995, 


[I]; Horco 1 

Molle, 14 Nov 1997, ].R. 


17-132 (BPI) [Il;J.B.Alberdi, 22 



Escaba, 28 ^ 

iov 1997 J.R.Herndndez 97- 


5 May 1995, 

J.R.Herndndez 95-036 (BPI 

III; Sanjavier, Parque Sierras 


:r, 27 Mar 1993. J.F.Hennen, 

LD.Ploper &J.R.Herndndez 93-027 (LIL 

54713, BPI) [ 

I-IIl; San Pedro de Colalao, 1 

Dept Santa Victoria Los Toldos, 28Jun 
1996,JR Herndfidez 96-033 (BPI 841132) 
[11] Tucuman: road to Villa Nougues, 5 
May 1995 ] R Hei nandez 95-035 (BPI 
841082) [II[, San Javier, 27 Mar 1993, 
JFHennenJ.DPlopcf &JRHerndndez 

061 (LIL 54715, BPI 841030) [II-IIIj. 
ospodium elegans (J. Schrot) Cummins-^ 

841048) [ml. 
on Tabebuia aveUanedae Lorentz ex Griseb.; 

Mision, 30 Jun 1996, J.R.Herndndez 96- 
057 (BPI) [III]. Tucuman: Cevil Pozo, 12-15 
km E of San Miguel de Tucuman, 30 Mar 
1993,J'.F.Hennen &J.R.Herndndez 93-056 
(LIL 54739, BPI) [II-III]; Dept. Capital, 1 
Nov 1995, J.R.Herndndez 95-062 (BPI 
841095) [O-U; Dept. Capital, 1 Nov 1995, 
J.R.Herndndez 95-061 (BPI) [O-I]; Dept. 




=ba, 15 Nov 1995 J. R.Herndndez 95< 
I 841112) [ii-ml. 

na sp .; Tucuman: Dept. Capital, 12 A 




1 841057) [11 

-III]. Tucuman: 


Mar 1993, 

J.F.Hennen & 


93-06J(LIL 54721, BPI) [II- 

ml; San Pedrc 

) de Colalai 

3, 1 Apr 1994, 

J.F.Hennen & 

54720, BPI 841 


Prospodium singeri Pe 


on Clytostoma call 


:ham.) Bureau 




Miguel Lillc 

R Singer sn [II 
Uropyxis rickiana Ma^ 

^nus ' 

on Macjadyenaun 


) AH Gentry 

Salta Dept 

la Frontera 

Horconeb 6 De 

'L 1997 J Rl 

153 (BPI 84125 


6 Dec 1997 ji?; 

^7-155 (BPI) [II] 

Tucuman 19 5 

km W ol ^ 

,an Miguel de 

LDPlopn &j 




torco MoUe, 17 Sep 1997J.R.Herndnd, 
7-079 (BPI) [II]; Horco Molle, 14 Nov 19^ 
R.Herndndcz 97-134 (BPI 841237) [I 
ferba Buena, 29 Mar 1993.J.R Herndnd 

Dept. Capital, 27 M 
54767, BPI) [II]; J.B./ 

28 Nov 1997, j.R.Hernm 
[II]; San Miguel de Tucui 

1 Ipomoea hieronymi (K 

hroxylum coca Lam.;Jujuy: 2 Apr 1993, 
iennen&L.D.Ploper 93-115 (.UL 5477 4, 

'-J35A (LIL54775,E 



, El Rodeo, 1 

,v 1999 J.RMerndndcz 

99-005 (BF 


xt to monastery, 3 Nc 

3V 1997, j.f 

rndnde^97-M9 (BPI 841 

246) [III]. 


Javier, Parqu 


rndnde^ 94-158 (LIL 547 


DietelinFngl &Pr- 

'tui tetraphylla (Pn 
lentes Dept Concepc; 
I Estancia bantillan. 

(Kunth)Benth'^ Salta 
1993 JFHenncn & 

odeColalaol Apr 1994 
IHtnnen 94 054 (LIL 
jan Pedro deColalao 2 

&J.R.Herndndez 97-041 (BPI 841187) [ 

1997 J.F.Hennen&J.R.Herndndez97-0 
(BPI 841185) [II-IIII. Salta: Pichanal, 2 A 
54834, BPI) [11-111]; Pichanal, 2 Apr 19^ 
J.F.Hennen & L.D.Ploper 93-108 (L 
54850, BPI) in], Tucuman: San Pedro 


Jujuy: Ledesma, between Valle ( 


Nogalal. 23 Sep 1997, J.FHennen& J 

and Rio Agua Negra, 8 Apr 1971, Lt 

^Cue^zo 7970c [O-lIll; Ledesma, fr 

cm Rio 


Agua Negra to Alba de Caiias, : 

Z9 May 

on Erj/thrznacnsta-galliL.; Buenos Aires: 

l977,Legname&Cuezzo 8247c [\ 

Plata. 29 Sep 1997. J.F.Hennen &i 

Hook. & Arn.) H.S. Irwin & Bai 


on undtn. ; Tucuman: Cevil Pozo, 12-15 kn 

Tucuman: Dept. Trancas. Cuesta C 


of San Miguel de Tucuman, 30 Mar 19' 

; Dept. 

J.F.Hennen &J.R.Herndndez 93-052 (L 

54849, BPI 841002) [Il-ml. 

S.Venturi s.n. [Ill]; Tafi del Valle 

,6 Apr 

Ravenelia spegazzinianaJ.C. Lindq.^^ 

on Acacia aroma Gillies ex Hook. & Ar 
Salta: Metan, 2 Apr 1993. J.EHennen 

Senna hookeriana Batke'\ Tucuma 

in: Tali, 


■035 (LIL 54842, BPI) [11-111]; Dept. Cruz 
:a, San Agustin, Monte Redondo, 29 
.r 1994, J.F.Hennen, M.M.Hennen & 
Hernandez 94-027 (LIL 54841, BPI) [11- 
; Las Cejas, 30 Mar 1993,J.FHcnnen & 


Apr 1994,J.FHennen (S'LD.PIoper 94-221 
(LIL 54852, BPI) [11]; Cafayate, 9 Apr 1994, 

Dept Trancab 17 Mai 1995 J R Hci 
95 070 (BPl 841102) [11] LasCii. 
Nox 1986 LDPIoper &GWui 

HernandiZ 94 JOS U IL 54859 BPl) |1 
Tucuman Dept Trancas Vipob 28 M 
1996JRHernandfc?96 002(BP1841117)[ 

34805, BPI) III-II]; San Pedro de Cola 

lao, 2 

n Allium sativum L.; Tucuman: Leaks, 15 

T lTsS bTi" n 1^1^"'"'" 

71 94- 

Aug 1992, LD.Ploper & Ramallo 


1994, j^j^j 


J.F.Hennen & L.D.Ploper 94-134 

(LIL p^^^. 


iniahetero teridis Thum ' " 

54808, BPI 841059) [IMII]. 

njanusia guaranitica (H. St.-Hil.) A. Juss.^; 



5 Apr 



(BPI) [III; W side of Catamarca, 23 Sep 

(LIL 54760, BPI 841020) [11-111], 

1997 J.F.Hennen &J.R.Herndndez97-034 


(BPI 841183) [IMII]. Tucuman: San Pedro 

Hyptis mutabihs (Rich.) Bnq,^- i 


Cafayate, 8 Apr 1994, J.F.Hennen, 


Hennen & J.R.Herndndez 94-102 


841027) [III]. 

54810, BPI) Ill-X]; Cafayate, 9 Apr 


man" C^tei 


Cruz Alta, San Agustin, 30 Mar 


n Pavonia malvacea (Veil.) Krapov. & 



Cnstoval^; Tucuman: San Javier, Parque 

94-037 (LIL 54809, BPI) [II-X 

I; El 

Sierras de San Javier, 23 Jul 1994, 

Manantial, 15 May 1996, j.R.Hernd 



jas, 30 Mar 

1993, J 




ncindc^ 93-07 

7 (LIL' 

34813 BPI 








"w of San Ml 




Mar 19 

>93, J FHenne 




(LIL 54814. BPD [II- 


on Menthc 

1 pulcgium L 

, Tucu, 

man: Vail 

c de 


ndndez 97-150 (BPh 



sp, Tucuman: 


Japual L 


1995, j- 



cinia pallic 


on Stachy, 

gil/iesn Ben, 





a, 15 Apr 1 

Cruz Alta San Agustin, 30 Mar 1994, 
JtHcnnen M MHenncn &]RHerndndez 
94-036 (LIL 54868, BPI) [HI], Cruz Alta, 


z 96 003 a 

JtHennen&]RHanandez93 06S(I II 
54871 BPI)[III] 
iSidaprosttataCiv Tucuman JB Alberdi 
Sol EscabaDike 22Sepl997 JFHenncn 
&J R Hei nandez 97 008 (BPl 841167) [111] 



\ Mercedes 15 Sep 

Hernandez 95-063 (BPI 841096) fill; San 
Javier, 27 Mar 1993 J.f^Hen ne n, L.D.Ploper 

BPI) [II]; San Pedro de Colalao, 16Jul 1992, 
j.RHerndndez & A.Wurschmidt 92-009 
(BPI) [III. 

Feb l995J.R.Herndndcz 95-065 C 
El Corte, 15 Feb 1995, J.R.Herndn 
066 (BPI 841098) [III 

between Salta and Jujuy, 7 Dec 1997, J.R. 
a Griseb,; Catamarca: ^^^"^^ Victoria, Los Toldos, El Nogalal, 29 

(LIL 54885, BPI) [O-I-IMII]. Tucuman: Sar 
Javier, Parque Sierras de San Javier, 23 Jul 
1994,;.R.Herndndez 94-164 (LIL 54887 
BPI) [I-II-III]; San Pedro de Colalao, 16 Jul 
1992, ].R.Herndndez & A.Wurschmxdl 
92-008 (BPI) [I]; San Pedro de Colalao, I 
Apr 1994, J.F.Hennen 6'M.M.Hennen 94- 

841138) [III. Tucuman: E slope of A 

Uar 1993, J.R.Herndndez93-C 
54783, BPI 840999) [III. 

s Dietel & Ellis 
1.; Jujuy: San Sah 

:>.Ploper & G.Wiirschmia 

Catalina, Oct 1941, 

J.CLindquis( s.n. (LPS 4895)[IH 

i28(LIL 54785 BPDilI llll Tucuman Dcpt Cruz Alta San Agustir 

1 CMoin polydaclyla (Lj b\\ Tucuman Montt Rtdondo 29 Mir 1994 J FHenner 

Dept Trancas Vipos 30 Mar 1995 JR MMHtnntn&JR Ha nandez 94 02i 

^ 034 (LIL 54803 BPUi 

54786 BPI Tucuman Dtpt Hpinl 23Junl996 

jRHt,,unnk^% OsJt BPI 841153) [11 III] 

iBhiiLi 1 Apr 1993 J 

^pi 1993 jtHinntn & ]R 841053)111 III] 

Z 93 126A aiL54780 BPI j [HI] Puccima purpurea Cooke' 
saeArthui' on Sorghum cajwium (Thunb ) 

91-089 (BPI 841212) [II-III]; Cevil Pozo, 12- 

on Ruprechtia laxijlora Meisn.-^; Tucuman: 

15 km E of San Miguel de Tucuman, 30 

Dept. Capital, 15 Aug 1994,J.R. Hernandez 

Mar 1993, J.F.Hennen &]R.ntrnandtz 

94-J35 (LIL 54893, BPI 841060) [III. 

93-05^ (LIL 54795, BPI) [11]; Dept. Trancas, 

on Ruprechtia sp.- Tucuman: SW ol J.B. 

Vipos, 28 Mar 1996 J. R.Herndndez 96-010 

Alberdi, 22 Sep 1997,;.EHennen &J.R^ 

Herndndez 97-004 (BPI 841163) [11]. 

1994, J.EHennen & M.M.Hennen 94-049 Puccinia polygoni-amphibii Pers.:Pers."^ 

on Polygonum acuminatum Kunth; Salta: 

ia recondita Roberge ex Desm. 

Cafayate, 9 Apr 1994, J.EHennen, MM. 

Bmmussp- Tucuman: Tafi del Valle, 15 Jul 

Hennen &]. R.Herndndez 94-116 (LIL 

1994, j.RBerndndez 94-169 (BPI) [II]; Tafi 

54892, BPI) [III Tucuman: San Javier, 27 

del Valle, 6 Apr 1993, J.EHennenc^j.R. 

Mar 1993, J.EHennen, L.D.Ploper &j.R. 

Herndndez 93-040 (LIL 54891, BPI) [II]. 

Bromus unioloides Kunth; Tucuman: El 

on Polygonum hydropiperoides Michx.; 

Manantial, 8 Oct 1995J.R.Herndndez 95- 

Catamarca: El Rodeo, 29 Nov 1997, J.R. 

Herndndez 97-123 (BPI 841229) [II[. Salta: 

Triticum aestxvum L.; Tucuman: Dept. 

Posta de Yatasto, 6 Dec 1997,J.R.Herndnde2 

97-157 (BPI 841253) [II]. Tucuman: road to 

Lapachos, 19 Sep 1999 J. R.Ucrndndez 99- 

Vi 1 la Nougues, 5 May 1995, J.R.Hernd ndez 

0.n (BPI 841279) [IHII]. 

9,5-034 (BPI 841081) [II[. 


on Polygonum sp.; Catamarca: E of La Vina, 

24 Sep 1997.J.EHennen &].R.Herndndez 

Frontera and Horcones, 11 Apr 1994, J.E 

97-059 (BPI) [II]; W side of Catamarca, 23 

Hennen&LD.Ploper 94-123 (LIL 54798, 

Sep 1997,J.EHennen &J.R.Herndndez 97- 

14 May 1995 J. R.Herndndez 95-050 (BPI 

Balcozna, 22 Sep 1997, J.EHennen & 

841091) [111]; Dept. Capital, 8 Mar 1988, 

J.R.Herndndez 97-012 (BPI) [11]. 

L.D.Ploper&G.Wurschmidts.n.iB?imi U 


El Colmenar, 14Jun 1989, L.D.Ploper & 

G.Wurschmidt 89-003 iWim-llll 

de Agronomia, 10 Oct 1999, J.R.Herndnde^ 

'Thlanthus mnarum 

Tucuman: San Javier, Parque Sierras de p^^^j^j^ leptochloae Arthur & Frommei ' 

San Javier, 23 Jul l994J.R.Herndndez 94- 


road to Parque Nacional Calilegua,' 8 De^c 

Digitaria sp.; Tucuman: Dept. Capital, 10 

I]. Tucuman: Dept. Capital, 12 Mar 1997, 

Mmnm-mr"''"'^"'^'^ ^^'^^^ ^^^' la Vega s.n. (BPI 841249) [O-I]. 


Clematis monlevidensis Spreng.-, G^rwasia imperialis(Speg.)J.C Lindq. 

Catamarca: El Rodeo, 29 Nov 1997, on Rulnis lwliviensisFocke\Tucuman: Horcc 

Alta Gracia, 19 Nov 1997, J.R.Herndndez ^'^'^^' J.F.Hennen, M.M.Hennen & J.R 

97 091 (BPl) [0 11 El Corte 15 Feb 1995 "^''"^^^^^ '^ ''' ^^IL 54902 BPl) [1 111] 

N0V1997 rRHtf nande- 97 148 (BPl) [01 Hu mincJt? 93 030A (L1L54903 BP 

J B° Alberdi roarto Esclba bank of creek 840998BJ [0 1] ban Ji\ lei Pirque bierns de 

28 Nov 1997 JR Hernandez 97 lOHBPl) Sin Jiyer 27 Mir l993jtHenntn 

10 11 NW of San Miguel de Tucuman 31 opttc^j tynamtz 
Mar 1993 J FHennen&JRHernana 
93 081 (LIL 54895 BPl) [0 H NW of S 

5 035 (BPl 841133) U 



Prunus domestica L.; Salta: Dept. San 
Dept. Capital, 10 Apr 1996 J. R.Herndndt 

841039) [II]; Metan, 2 Apr l993J.EHennen 
& L.D.Ploper 93-112 (LIL 54911, BPI) [II]. 
Tucuman: Dept. Capital, 27 Mar 1993, 
J.F.Hennen, L.D.Ploper &].R.Herndndez 
93-041 A (LIL54910, BPI) [II]; Dept. Capital, 
4 Apr 1994 J.EHennen &J.R. Hernandez 
94-081 (LIL 54913, BPI) [II]; San Pedro de 
Colalao, 3 Apr 1994, J.RHennen & M.M. 
Hennen 94-079 (LIL 54912, BPI) [Il|. 
Salix caprea L.^"; Tucuman: Dept. Capital, 
17 Dec 1995, ].R.Herndndez 95-096 (BPI 
841115) [II]. 

083 (BPI 841209) [II]; H 

4-093 (LIL 54906, BPI) [Il-Ull; San Javie 
7 Mar 1993, J.F.Hennen, L.D.Ploper i 
R.Herndndez 93-021 (LIL 54905. BPI) [II 

084 (BPI 841210) [II]; El Corte 
1993, J.F.Hennen, L.D.Plope, 

841202) [III; Santa Fe,ic 

dasycephala Cham. & S 
lan: El Corte, 15 Feb 19 
^dez 95-064 (BPI 841097; 

JRHefnandtz 94 145 (LIL 54Q24 BPl Dcpt Ti me^s 27C\i U 

841068) [III] 95-060(BPIJ[Il|SinlV 

.hyllum pampeanum (Speg J J C Lindq Mir 1994 J FHt nnt n (5 

1 Salpichwa onganijoha (Lam) Bull 045(LIL54937 BPDill 

\pT 199^, ].EHcnncn& J RHa 
?4-099 (LIL 54929, BPIj [IMIll. C 
i Apr 1994 J.FHennen &JRHei 
d4-099B (L1L54931, BPI) [II] Cal 

indez 90-002 iWl) mi 

aha and Jujuy, 7 Dec 199^ 

del Valle, 6 Apr 1993, J.F.He; 

94 033 (LIL 54936 I 

^ 091B (UL54733 E 

jspodium hppiae (Speg ) Arthui 

Tucuman Monte Redondo 31 Mir 

J R Hernandez 96 0^8 (BPI 841157) 

on Aloyiia polystachia (Cribeb) Mold 

Salta Cafayate 8 Apr 1994 JhHt 

(LIL 54946 

1996 N Canton sn 


. CruzAlta 


30 Mar 1993 J FH£ 



93 058 (LIL 5495( 





Mir 1993 J FHen 



93 080 (LIL 5494 

^ BPI) [11 11 


30 Mar 1993 J FHe 



93 062 (LIL 54951 BPI) [III 

NW of Sm 

Miguel de Tuci 


93 084 (LIL 


uss Salta 

ras de San Javier, 23Jul 1994 J.R. H 


on Lippia sp.; Salta: Dept. Santa Vict 

ona. La 

94-163 (LIL 54956, BPl 841073) 



dez 96- 

Lippiaalbaium.) N.E.Br.^; Tucuir 


062A (BPl) [III]. Tucuman: J.B. / 


Capital, 14 May 1995, J. R.Hernd 

road to Escaba, shrine. 28 Nov V 


049 (BPl 841090) [III]. 


Lippia dr. turnerijolia Cham.^;! 


on Lippia tuihinataGYisth-TucumsL 




Trancas, \'ipos, 30 Mar 1995,i.R. Her 

95-002 (BPl 841078) [III]. 

QShWIBPI 841088) [III]. 

New rust species (N): 

Aecidium sp. (on Dyschoriste sp.), Aecidium sp. (on Solanum riparium Pers., 
Solarium tucumanense Griseb., Vassohia hreviflora (Sendtner) Hunz.), 
Cerotelium sp. (on Ruprechtia apetala Wedd., Ruprechtia laxiflora iVleisn., 
Ruprechtia sp.), Endophyllum sp. (on Fuchsia holiviana Carriere), Puccinia sp. 
(on Vernoniafulta Griseb., Vernonia sp.), Puccinia sp. (on Vernonia squamosa 
Gardner), Puccinia sp. (on Ichnanthus minarum (Nees) Doll), Skierka sp. (on 
Cupania vernalis Cambess.), Uredo sp. (on Psychotria carthagenensis Jacq.), 
Ypsilospora sp. (on Inga edulisMznX 
New rust records for Argentina (A): 

Aecidium tournefortiae Henn., Catenulopsora praelonga (Speg.) Buritica, 
Cionothrix praelonga (G. Winter) Arthur, Coleosporium vernomae Berk. & M.A. 
Curtis, Desmella anemiae Syd. & P. Syd., Edythea quitensis (Lagerh.) H.S.Jackson 
& Holw., Frommeella mexicana var. indicae ].W McCain & J.F. Hennen, 
Melampsora epitea (Kunze & J.C. Schmidt) Thum., Melampsoridium 
hiratsukanum S. Ito ex Hirats. f., Phakopsora meihomiae (Arthur) Arthur, 
Phakopsora neocherimoliae (Cummins) Buritica & J.F. Hennen, Phakopsora 
nishidana S. Ito, Phakopsora zizyphi-vulgaris Dietel, Phragmopyxis degluhens 
(Berk. & M.A. Curtis) Dietel in Engl. & Prantl, Physopella compressa (Mains) 
Cummins & Ramachar, Prospodium pithecoctenii (Pazschke) Cummins, 
Prospodium singeri Petr, Puccinia aristidae Tracy var. chaetaria Cumm. & 
Husain, Puccinia bergii Speg., Puccinia calcitrapae DC. var centaureae (DC.) 
Cummins, Puccinia chaetochloae Arthur, Puccinia cnici-oleracei Pers., Puccinia 
commelinae Holw., Puccinia cordiae Arthur, Puccinia coronata Corda var. 
rangijerina (S. Ito) Cummins, Puccinia exornata Arthur, Puccinia hyptidis- 

mutahilis Mayor, Puccinia inclita Arthur, Puccinia incondita Arthur, Puccinia 
lateripes Berk. & Ravenel, Puccinia levis (Sacc.6a: Bizz.) Magnus var panici- 
sanguinalis (Rangel) Ramachar & Cummins, Puccinia ocellifera Cummins, 
Puccinia pelargonii-zonalis Doidge, Puccinia proluviosa H.S.Jacks. & Holw., 
Puccinia rata H.S.Jacks. «&r Holw., Puccinia tanaceti DC. var tanaceti, Puccinia 
thunhergiae Cooke, Puccinia triumjettae Dietel & Holw., Puccinia unicolor 
Arthur, Ravenelia cohniana Henn., Ravenelia echinata Lagerh. & Dietel var. 
ectypa (Arthur & Holw.) Cummins, Ravenelia macrocarpa Syd. & R Syd., 
Uromyces aspiliae H.S.Jacks. &r Holw., Uromyces castaneus R Syd. & Syd., 
Uromyces celosiae Dietel & Holw, Uromyces dietelianus Pazsche, Uromyces 
dolichosporus Dietel & Holw, Uromyces iresines Lagerh. ex Syd. & R Syd., 
Uromyces orbicularis Dietel, Uromyces sp., Uromyces transversalis (Thiim.) G. 

New host plant genus for this rust worldwide (G): 

Anadenanthera sp. (Ravenelia cehil Speg.), Chaetothylax umhrosus Nees 
(Puccinia justiciae Puttemans), Dyschoriste sp. (Aecidium sp.), Ruprechtia 
apetala Wedd. (Cerotelium sp,), Ruprechtia laxifhra Meisn. (Cerotelium sp.), 
Ruprechtia sp. (Cerotelium sp.). 
New host plant species for this rust worldwide (S): 

Athyrium lilloi (Hicken) Alston (Desmella anemiae Syd. & R Syd.), Baccharis 
latijolia Lem. (Puccinia colossea Speg.), Calliandra formosa (Kunth) Benth. 
(Ravenelia echinata Lagerh. & Dietel var ectypa (Arthur & Holw.) Cummms), 
Canna compacta Roscoe (Puccinia thaliae Dietel), Chamissoa altissima (Jacq.) 
Kunth (Uromyces celosiae Dietel & Holw), Clematis montevidensis Spreng. 
(Puccinia recondita Roberge ex Desm.), Clematis sericea Michx. (Puccinia 
recondita Roberge ex Desm.), Cologania ovalifolia Kunth (Phakopsora 
meihomiae (Arthur) Arthur), Coursetia hrachyrhachis Harms (Phragmopyxis 
degluhens (Berk. & M.A. Curtis) Dietel in Engl. & Prantl), Desmodium 
suhsericeum Malme (Uromyces orbicularis Dietel), Echinochloa cruspavonis 
(Kunth) Schult. (Puccinia abnormis Henn.), Grahowskia schizocalyx Dammer 
(Puccinia paradoxapoda Speg.), Iresine diffusa Humb. & Bonpl. ex Willd. var 
diffusa (Uromyces iresines Lagerh. ex Syd. & R Syd.),Janusia guaranitica (H. St.- 
Hil.) A. Juss. (Puccinia heteropteridis Thiim.), Lippia alba (Mill.) N.E.Br. 
(Puccinia lantanae Earl.), Lippia cfr turnerifolia Cham. (Puccinia lantanae 
Farl.), Lippia grisebachiana Moldenke (Puccinia lantanae Earl.), Lippia modesta 
Briq. (Puccinia lantanae Earl), Lonchocarpus lilloi (Hassl.) Burkart (Ravenelia 
lonchocarpiicola Speg.), Mimosa debilis Humb. & Bonpl. ex Willd. var. debilis 
(Ravenelia mimosae-sensitivaeHennXParapiptadeniaexcelsa(Ghseh.)Burkan 
(Ravenelia papillosa Speg.), Pavonia malvacea (Veil.) Krapov & Cristoval 
(Catenulopsora praelonga (Speg.) Buritica), Ruellia ciliatiflora Hook. (Puccinia 
lateripes Berk. & Ravenel), Ruellia erythropus (Nees) Lindau (Puccinia sp.), 

Salvia )\paia Briq, (Puccniu; farinacca Long), Senccio hravcnsis Cabrera 
{Colcosporium tus<.ilagUus (Pers.) Lev.), Senccio crcmciflorus Mattf. 
(Coleosporiumtussilaginis,{Pers.) Lev.), Senccio otoptcrusGnsehXCokosporium 
tussilaginis (Pers.) Lev), Senccio pcrcgrinits Griseb. (Puccinia proluviosa H.S. 
Jacks. & Holw.), Senna sulmlata (Griseb.) H.S. Irwin 6a: Barneby (Ravenelia 
macrocarpa Syd. & P. Syd.j, Serjania nunginata Casar (Puccinia arechavaletae 
Speg.), Simsia domheyana DC {Puccinia enceJiae Dietel & Holw var aemulans 
(Syd. & P Syd.) Parmelee), Solanu m almtiloidcs (Griseb.) Bitter & Lillo (Puccinia 

tucumanense Griseb. (Accidium sp ), Stachys gilhcsii Benth. (Puccinia 
pa/lidissima Speg.), Tarasfljorgcn.sfnn (1 M )ohnst.JKrapov (Pucciniflp/atyspora 
(Speg.) H.S.Jacks. & Holw.), Toumcfortia ruhuunda DC. (Uromyces 
dolichosporus Dietel & Holw.), Tnpogandra elongata (G. Mey.) Woodson 
iPuccxnia commelinae Holw), Vassobu! hrevijlora (Sendtner) Hunz. (Aecidium 
sp.), Vernonia fulta Griseb. (Puccinia sp. A), Vernonia remotiflora Rich. 
(Puccinia cnici-oleracci Pers.), Vernonia saltensis Hieron. (Puccinia cnici- 
oleracei Pers., Puccinia rata H.S.Jacks. & Holw), Wcdc/ia saltensis Cabrera 
iUromyces aspiliae H.S.Jacks. & Holw), Zizyphus mistol Griseb. (Phakopsora 
Zizyphi-vulgaris Dietel). 

New host plant species for any rust in Argentina (A): 

Alcca rosea L. (Puccinia platyspora (Speg.) H.S.Jacks. & Holw.), Alcea sp. 
(Puccinia platyspora (Speg.) H.S.Jacks. & Holw), Ainus acunnnata Kunth 
(Melampsondium hiratsukanum S. Ito ex Hirats. f.), Bauhinia Jorficata Link 
(Uromyces dietelianus Pazsche), Carthamus tinctorius L. (Puccinia calcitrapae 
DC. var. centaureae (DC.) Cummins), Cissus sicyoides L. (Endophyllum 
circumscriptum Whetzel & Olive var catamarcensisj.C. Lindq.), Cucurhitella 
cucumifolia (Griseb.) Cogn. (Uromyces novissimus Speg.), Cupania vernalis 
CamhessXSkierkaspXDesmodiumpurpureumHook.&nArn.iUromyces orbicu- 
laris Dietel), Desmodium purpureum Hook. & Arn. (Uromyces castaneus P Syd. 
63: SydX Dichondra sericea Sw. (Puccinia dichondrae Mont.), Duchesnea indica 
(Andrews) Focke (Fronrmeella mexicana var. indicac J.W. McCain & J.F. 
HennenXG/adioIus sp.(L/romycestransvcrsa/is(Thum.)G. Winter), Hydrocoty/e 
bonariensis Comm. ex Lam. (Puccinia hydrocotylcs Cooke), hhnanthus 
minarum (Nees) Doll (Puccinia sp. B, Puccinia inc/ita Arthur, />Mccinia levis 
(Sacc.& Bizz.) Magnus var pa n icisa ngu i nalis (Rangel) Ramachar & Cummins), 
Leptochloa mucronata (Michaux) Kunth (Puccinia leptochloae Arthur), 
Mikania micrantha Kunrh {Pucunia spegazzinii De Toni), Pelargonium 
(Prior.) Burkart (Puccinia bcrgii SpcgJ, Ptcns deXlexa Link (Desmella ancmiae 
Syd. & P Syd.), Rubus boliviensis Focke (Gerwasia imperialis (Speg.)J.C. Lindq., 
Kuehneola loeseneriana (Henn.) H.S. Jacks. & Holw), Senna birostris var. 

hookeriana (Gillies ex Hook. & Arn.) H.S. Irwin & Barneby (Ravenelia 
macrocarpa Syd. & P. Syd.), Senna hookeriana Batke (Ravenelia macrocarpa Syd. 
&r P. Syd.), Setaria lachnea (Nees) Kunth {Puccinia chaetochloae Arthur), Se- 
taria parviflora (Poir.) Kerguelen (Puccinia chaetochloae Arthur), Sorghum 
cajrorum (Thunb.) P Beauv. (Puccinia purpurea Cooke), Thelypteris 
quadrangularis (Fee) Schelpe (Desmella anemiae Syd. 61 R Syd.), Thelypteris 
sp. (Desmella anemiae Syd. & R Syd.), Tournefortia paniculata Cham. (Aecidium 
tournefortiae Henn.), Tripogandra sp. (Puccinia commelinae Holw.), 
Tnumphetta sp. (Puccinia triumfettae Dietel & Holw.), Valeriana sp. (Aecidium 

New host plant species for which another rust was previously known from 
Argentina (AR): 

Aloysiavirgata(Rmz&iPzv.)] (ippiae (Speg.) Arthur), A ristida 
sp. (Puccinia aristidae Tracy var. chaetaria Cumm. & Husain), Aspilia sp. 
(Uromyces aspiliae H.S.Jacks. & Holw.), Avena sativa L (Puccinia coronata Corda 
var. rangiferina (S. Ito) Cummins), Baccharis sp. (A) (Puccinia exornata Arthur), 
Baccharis sp. (B) (Puccinia exornata Arthur), Baccharis lilloi Heering (Puccinia 
baccharidis Dietel &c Holw), Baccharis sp. (Puccinia unicolor Arthur), Chloris 
gayana Kunth (Puccinia cacahata Arthur & Holw.), Chrysanthemum sp. 
(Puccinia tanaceti DC. var tanaceti), Clytostoma ca/listegioides (Cham.) Bureau 
ex Griseb. (Prospodium singeri Petr), Digitaria insularis (L.) Fedde (Puccinia 
oahuensis Ellis & Everh.), Elephantopus mollis Kunth (Coleosporium vernoniae 
Berk. ^ U.K. Curtis), Eupatorium macrocephalum Less. (Puccinia conoclinii 
Seym, in Burrill), Eupatorium sp. (Cionothrix praelonga (G. Wmter) Arthur), 
Hyptis mutahilis (Rich.) Briq. (Pucania hyptidis-mutahilis Mayor), Hyptis sp. 
(.Puccinia hyptidis-mutahilis Mayor), Inga edulis Mart. (Ypsilospora sp.), Pani- 
cum sp. (Puccinia levis (Sacc.& Bizz.) Magnus var panicisanguinalis (RangeD 
Ramachar & Cummins), Paspalum sp. (Physopella compressa (Mains) Cummins 
& Ramachar), Pluchea sp. (Puccinia ocellijera Cummins), Pteris sp. (Desmella 
anemiae Syd. & R Syd.), Sahx hahylonica L. (Melampsora epitea (Kunze &J.C. 
Schmidt) Thiim.), Salix caprea L. (Melampsora epitea (Kunze & J.C. Schmidt) 
Thum.), Salix humholdtiana Willd. (Melampsora epitea (Kunze &J.C. Schmidt) 
Thiim.), Setaria sp. (Puccinia chaetochloae Arthur), Tradescantia sp. (Puccinia 
commcHnae Holw). 

The first author gratefully acknowledges his advisor and friend, L. Daniel Ploper, 
National University of Tucuman, who provided support in completing his doc- 
toral thesis. Many thanks to Maria Magdalena Schiavone and her staff at the 
herbarium of the Fundacion Miguel Lillo (LIL), San Miguel de Tucuman, 
Tucuman, who organized and accessioned many specimens, and to Alberto 

Slanis for host plant identifications. The efforts of Erin McCray and David F. 
Farr in providing accession numbers and assistance m the final publication, 
respectively, are appreciated as are the encouragement and efforts of Amy Y. 
Rossman that contributed to the completion of this paper Finally, I would like 
to thank my wife, Mary E. Palm, for her invaluable motivation and support. 

Brummit, R.K. and C.E. Powfi i (Eds.). 1 992, Authors of plant names. Royal Botanic Gardens, 

Cabrera, A.L. 1994. Regiones fitogeograficas argentinas. Enciclopedia Argentina de 
Agricultura y Jardineria (1 ra reimpresion) T. II. Buenos Aires, Argentina. 

J. Cramer, Lehre, Germany. 
Hernandez, J.R. 2000. eaeodromusranunctv//, a new rust on Ranunculus from Argentina and 

a synopsis of Saeoc/romus. Mycotaxon 76:329-336. 
Kirk, RM.andA.E.ANSELL.1992.The author of fungal names. CAB International. Wallingford, 

United Kingdom. 
LiNDQuiST,J.C. 1982. Royasde la Republica Argentina y Zonas Limitrofes. Instituto Nacional 

de Tecnologi'a Agropecuaria, Buenos Aires, Argentina. 


Eric H. Roalson Thomas R.Van Devender and 

School of Biological Sciences Ana Lilia Reina G. 

f the Rio Mayo region of the Mexican state of Sonera, particularl) 
ncovered more species of Cyperaceae not previously reported fron 
in some cases, northern Mexico. Here we present 14 species of Cypera 
; not previously known to occur in the Rio Mayo region, clarify the j 
Cyperus (Cyperaceae) species, and discuss the significance of the ri 
1 new discovery This brings the total number of Cyperaceae taxa kn 

as especies de Cyperaceae q 
ora, o, en algunos casos, el no 

Gentry's 1942 Rio Mayo Plants was a major contribution to the knowledge ol 
the flora and vegetation of northwestern Mexico. The Rio Mayo region includes 
the area in southern Sonora and adjacent Chihuahua between the Rio Fuerte 
and Rio Yaqui drainages. Beginning in the 1970s, a new network of roads, espe- 
cially Mexico Federal Highway 16 (MEX 16) between Hermosillo, Sonora, and 
La Junta, Chihuahua, was completed in 1992 and provided access to many ar- 
eas of the Sierra Madre Occidental not previously explored f loristically (Biirquez 
et al. 1992). A revision of the Rio Mayo flora (Martin et al. 1998) incorporated 
recent floras from the Cascada de Basaseachi (Spellenberg et al. 1996) and 
Nabogame (Laferriere 1994), Chihuahua. The total of 2825 taxa reported for the 
Rio Mayo region included 11 genera and 77 species of Cyperaceae and two gen- 
era and 12 species of Juncaceae. 

An intensive f loristic survey of the Municipio de Yecora, an area of 3,300 
km^ along MEX 16 in the Sierra Madre Occidental in eastern Sonora, was be- 
gun in 1995 (Reina et al. 1999; Van Devender et al. in press). The current total for 
the Municipio is 1648 taxa, raising the total for the Rio Mayo region to over 3000 

taxa. The families with the most taxa in the Municipio de Yecora flora are 
Compositae (248 taxa), Gramineae (186 taxa), Leguminosae (168 taxa), 
Cyperaceae (59 taxa), and Euphorbiaceae (56 taxa). The genus Cyperus with 29 
species is especially diverse. This ongoing f loristic work in the Rio Mayo region 
has uncovered more species of Cyperaceae not previously reported from the 
region, the state of Sonora, or, in some cases, northern Mexico. Here we present 
14 species of Cyperaceae and two species of Juncaceae not previously known to 
occur in the Rio Mayo region, clarify the presence of three varieties of one 
Cyperus (Cyperaceae) species, and discuss the significance of the range exten- 
sion involved with each new discovery Specimens were determined by Eric H. 
Roalson, unless noted otherwise. Most of the noteworthy records are in cienegas 
or other moist areas from 1200 to 2100 m elevation in oak woodland and pine- 
oak forest in the Municipio de Yecora, or the grasslands and mudf low barrens 
within them (Reina G. et al. 1999). 
Bulbostylis barbata (Rottb.) C.B.Clarke 

This species is previously known from the southeast United States, Martinique, 
Guatemala, and the Old World tropics (Krai 1971). Bulhostylis harhata appears 
to be quite weedy and expanding its range in North America (Krai 1971). This is 
the first report of the species in Mexico. 

Voucher specimens. MEXICO. Sonora. Municipio dc Yecora: 14.7 km E of Maycoba, 7.7 km E of El Kipor 
on MEX 16 (KM 342), pine-oak forest, 2876'04"N 108-3336"W, 1460 m elevation, locally common herb 

Bulbostylis pubescens (J. & C.Presl) Svenson 

This species is widespread in the lowlands to the south along the Pacific slope 
of the Sierra Madre including the Mexican states of Sinaloa, Nayarit, Jalisco, 
Guerrero, Mexico, Oaxaca, and Chiapas, and south through Central America 
and northern South America (Krai 1971; McVaugh 1993). This is the first report 
of the species from the Rio Mayo region and the state of Sonora. 

oak woodland on barren volcanic hilltops, 28'22'40"N 108'56'W, 1^40 m elevation, locally common 

Carex planostachys Kunze 

This species is previously known from Texas to Guatemala, and in Mex 
known from as far northwest as Chihuahua (Hermann 1974). This is t 
report of this species in Sonora and the Rio Mayo region. 

arex thurberi Dewey 

his species is known from Arizona and localities across Mexico (Sonora, Chi- 
uahua, Jalisco, Veracruz, and Chiapas), as well as Guatemala (Hermann 1974; 
though see McVaugh 1993; Espejo & Lopez F. 1997). This is the first report of 
lis species for the Rio Mayo region. 

Jul 1998, Irauba 425-98 (WS). 
Cyperus arsenei O'Neil & Ben.Ayers 

This species is previously known from the western Mexican states of Baja Cali- 
fornia Sur, Nayarit, and Michoacan and the Districto Federal (Tucker 1994). This 
is the first record of this species in the state of Sonora and the Rio Mayo region. 

Voucher specimens. MEXICO. Sonora. Municipio de Yecora: 1 km W of Yecora on volcanic ridges/ 
outcrops, W of cemetery, E facing slope with occasional Arctostaphylos, Bouteloua, other Cyperus sp., 

dcscanlia, 28-2212"N f08-56'23"W, ca. 1600 m elevation, 5 Sep 1996, Roalson 1364 (WS). 
Cyperus manimae HBK 

In Gentry's Rio Mayo Plants (Martin et al. 1998), varieties of C manimae were 
not distinguished. Tucker (1994) recognized three varieties of C manimae, with 
only one of these (C manimae var asperrimus) known from Sonora. Cyperus 
manimae var divergens is known from Chihuahua and Coahuila south to 
Chiapas in Mexico and south to Costa Rica and C manimae var. manimae is 
known from Baja California, Sonora, and Durango to Chiapas in Mexico, south 
to Guatemala and northern South America (Tucker 1994; Espejo & Lopez F 
1997). All three varieties are present in the Rio Mayo region. This increases the 
range of var asperrimus and var. manimae into the Rio Mayo region, and the 
range of var divergens west from Chihuahua into Sonora. 
var asperrimus (Liebm.) Kiik. 

m shady slope, 27 Sep 1998, V 
s (HBK) Kuk. 

)cky stream canyon m pine-oak forest, 28-2r38"N 108-53'55"W, 1520 m 
rocky slope, 18 Sep 1998, Van De vender 98-J340(WS), with A.L. Reina 

4 108-55'38"W, 1460 m elevation, locally common perennial in mud at 

tall herbaceous perennial m moist places 30ctl9% HoresM 4969(USON Wb) withj Sanchez ca 
2km N\\ ol Recoil on old roid to banta Rosa 28 22 33 N 108 5024 W 1560 m elevation uncommon 
pcrcnni Urn moist soil 17 Aug 1998 VanDevendt r98 J009(F1U WS) with A L RemaG MEFishbem 
indC M Ui^uson ( inidil i Ventana (Aiioyo El Otio Lado) 2 5 km (by aitj ESEof Yecora rocky 
stanncm)oninpinL oik forest 28 2138 N 108 5355 W 1520 m elc\ation uncommon perennial on 
rocky slope 18Sepl998 Van Dt\endti 98 1342 (WS) with A L ReinaG rnd W Trauba Rio Ma) coba 
at MEX 15(20 5 km W ol Ma)coba 28 6 km E ol \etora) 28 2215 N 108 4530 W 1220 m ele\ation 
rare sedge at base ol clitl 26 Sep 1998 Rcuui G 98 UU (WS) with TR Van Defender load to Ric 
Yepachic from MEX 16 near Ano>o Hondo (11 5 km f oi El Kipor on MEX 16) oak woodland 28 2650 N 
108 32 20W1480mele\ation %ery common on distm bed loidsidt 27 Sep 1998 VanDeMmki98 
1808{WS) with A L ReiniG 
Cyperus sphaerolepis Boeck. 

This species is known fiom New Mexico, Arizona, and west Texas south tc 
Puebla and the Districto Federal in Mexico, and has been documented previ- 
ously from Sonora (McVaugh 1993; Tucker 1994; Espejo & Lopez F 1997). This 
species was not listed, however, for the Rio Mayo region (Martin et al. 1998). These 
specimens verify its presence there. 

300 m elevation locally 
vender 97 i538(ARIZ \ 

vn from much of the United States, south- 
western Canada, and north-central Mexico. In Mexico it is previously known 
from Chihuahua, Durango, Aguascalientes, Jalisco, Michoacan, Mexico, 
Queretaro, the Districto Federal, Oaxaca, Puebla, Hidalgo, and San Luis Potosi 
(McVaugh 1993). This is the first report for its presence in the Rio Mayo region 

» S.Gonzalez E. 

This species was described m 1985 from the state of Durango, Mexico (Gonzalez 
E. 1985) and is also known from the states of Jalisco (McVaugh 1993) and Sinaloa 
(Espejo & Lopez E 1997). These collections represent the first record of this spe- 
cies in the Rio Mayo region and Sonora and are a significant northward expan- 
sion of the species distribution from Durango and Sinaloa. 

Voucher specimen. MEXICO. Sonora. Municipio de Yecora: NW of the cemetery in Yecora, sparse 
oak woodland on barren volcanic hilltops, 28"22'40"N 108-56'W, 1540 m elevation, uncommon on 
moist rock surfaces, 23 Sep 1997, Reina G. 97-1171 (WS), with T.R. Van Devender and W. Trauba; Cienega 
de Camilo, 6.3 km E of El Kipor, 11.2 km W of Chihuahua border on MEX 16, sphagnum bog in clear- 
ing in shady riparian pine-oak forest, 28"46'43"N 1083r50"W, 1520 m elevation, abundant sedge in 
mud at edge of cienega, 27 Sep 1998, Van Devender 98-1827 (V^S), with A.L. Reina G. (Van Devender et 

Eleocharis yecorensis E.H.Roalson 

This species was described in 1999 from material collected in a freshwater 
marsh on the outskirts of Yecora (Roalson 1999). It now appears that this spe- 
cies is quite widespread, with documented localities as far south as Nicaragua 
(Stevens et al. 2001). 

28-22'02"N 108-55'47"W, 1,505 m, 16 Jul 1997, Van Devender 97-810 (ARIZ, WS). 
Fimbristylis complanata (Retz.) Link 

This species is previously known from the New and Old World tropics (Krai 
1971; McVaugh 1993), and prior to Martin et al.'s (1998) discovery of it in Chi- 
huahua, the northernmost record of this species in the New World was from 
Durango. This is the first report of this species from Sonora. This species has 
also recently been collected by Richard S. Felger and collegues N of Guaymas, 
Sonora (Municipio de Guaymas, Cafion La Balandrona, N side of Sierra El Aguaje, 
28°04.27'N, lir04.44W, 825 ft. elevation, 19 Dec 2001, Felger 01-659 (USON, 
ARIZ, MEXU), with J. Sanchez E. and P.A. West; R.S. Eelger, pers. comm.). 

the road to Agua Blanca, rocky stream canyon in pine-oak forest, 2876'27"N 108-55'29"W, 1520 m 

Fimbristylis decipiens Krai 

This species was described by Krai in 1971 "... to repress 

phology between F. dichotoma and F. annua." It is dif 

from F. d ichotoma and F. annua, but the two specimens listed below seem to fall 
within Krai's circumscription. Krai recognized this species in the southeast 
United States, but suggested it was likely in Central and South America and the 
Old World (Krai i971). This is the first report for Sonora and the Rio Mayo region. 

ash, 28"2674"N 108-33'20"W, 1400 m elevation, 25 Sep 1997, Reina G. 97-1374 (ARIZ, MEXU, NY, WS), 
with T.R. Van Devender; Arroyo El Toro on NE edge of Yecora, 2872'45"N 108-55'15"W, 1520 m eleva- 
tion, locally common sedge in mud near stream, 2 Oct 1998, Van Dewnder 98-1997 (WS), with A.L. 

Fimbristylis pentastachya Boeck. 

This species has been traditionally known from southern Mexico (Krai 1971). 
McVaugh (1993) listed F. pentastachya from southern Sonora and Martin et al. 
(1998) listed it for the Rio Mayo region in Chihuahua. This collection expands 
its range into the Sonoran portion of the Rio Mayo region. 

Lipocarpha maculata (Michx.) Torr. 

This species is previously known from the southeastern United States, in Mexico 
the states of Nayarit, Jalisco, Veracruz, and Chiapas, Central and South America, 
and the West Indies, although this species is thought to be uncommon m Mexico 
(McVaugh 1993). This is the first report of this species from the Rio Mayo region 
and the state of Sonora, with the closest documented locality to the south in 
Jalisco and Nayarit (Espejo & Lopez F. 1997). 

(Arroyo El Otro Lado), 2.5 km (by air) ESE of Yecora, rocky stream canyon in pine-oak forest, 28-2r38"N 
108-53'55"W, 1520 m elevation, locally abundant dwarf sedge on shallow soil on mud flow slope, 1 
Oct 1998, Rema G. 98-1967 (ARIZ, WS), with T.R. Van Devender and W. Trauba. 
Scleria interrupta Rich 

Two collections from the Rio Mayo region apparently fall within the circum- 
scription of Scleria interrupta (Konraed Camelbeke, pers. comm.). This is an 
extreme range extension with the closest localities of this species in Chiapas 
(Espejo & Lopez F. 1997). The two collections vary somewhat m the character- 
istics of pubescence and extent of the tubercles on the achene. Reina G. 97-1373 
has achenes with a large number of tubercles covering the surface which at 
times approach a reticulate pattern and most of the bracts subtending the spikes 
are covered in bristly hairs on the margins and midrib. This appears to fit well 
within the circumscription of S. interrupta, somewhat tending towards the 
sometimes-segregate S. pinetorum Britton which is included in synonymy of S. 

interrupta here (Core 1936; Raynal 1976; Konraed Camelbeke, pers. comm.). The 
other collection, Reina G. 97-1174, includes plants with achenes that are mostly 
smooth with a few tubercles at the achene apex and the bracts subtending the 
spikes are often near glabrous with few, scattered, bristly hairs. These plants do 
not fit as well in the circumscription of S.interrupta, and approach S. verticillata 
Muhl. (due to the lack of bristly hairs), or maybe 5. distans Poir. (due to the 
smooth achenes; Core 1936). For now these two collections are treated as 5. 
interrupta, but more detailed studies of the variation in achene ornamentation 
and pubescence in these annual Scleria species is necessary. 

Juncus dichotomus Elliott 

This species is quite widespread, occurring from Alaska, across the United States, 
Mexico, Central America, and South America (Balslev 1996; Espejo 62: Lopez F. 
1996; Brooks & Clemants 2000). This is the first report of J. dichotomus for the 
Rio Mayo region. 

ranca El Salto, pine-oak forest, 28"2ri8"N 109-0r48"W, 2000 m elevation, locally common, 14 

son, M. Kaib, and M.J. Martinez C; Arroyo Hondo, 11.5 km E of El Kipor, 4 km W of Chihua: 
der on MEX 16, pine-oak forest, 28-26'30"N 10832'30"W, 1460 m elevation, locally common hei 
js perennial m water m rocks, 25 Sep 1997, Rema G. 97-1392 (WS), with T.R. Van Devender; 2.3 
ve Restaurant Puerto de la Cruz on road to microwave tower, Mesa del Campanero, pine-oak 

G.M. Ferguson; Canada La Ventana (Arroyo El Otro Lado), 2.5 km (by air) ESE of Yecora, rocky stream 
canyon m pine-oak forest, 287r38"N 108"53'55"W, 1520 m elevation, solitary clump in moist soil in 
narrow side canyon, 18 Sep 1998, VanDevender98-1303 (ARIZ, WS), with A.L. Reina G. and W. Trauba; 
Cienega de Camilo, 6.3 km E of El Kipor, 11.2 km W of Chihuahua border on MEX 16, sphagnum bog 

Juncus liebmanii J.FMacbr. var. polycephalus Balslev 

This species is known from Chihuahua in Mexico south to Ecuador. The northern 
Mexican collections are considered part of J. liehmanii var liebmanii or var poly- 
cephalus Balslev (Balslev 1996). The Sonoran collection belongs to var polycephalus. 
This is the first report of this species for Sonora and the Rio Mayo region. 

We thank Father Bill Trauba for help and companionship in the field and pro- 
viding shelter in Yecora; George Ferguson, Mark Fishbein, Sam Friedman, and 
Richard Spellenberg for help in the field and for sharing their collections; Miriam 
C. Fritts for help with Carex identifications; and Richard S. Felger and M. Socorro 
Gonzalez E. tor helpful comments on a previous version of the manuscript. 


Balslev,H. 1996. Flora r 

leotropica. 1 

Monograph 68. Juncac 


^w York Botanical 

Garden, Bronx. 

Brooks, R.E. and S.E.Clem 




Editorial Commit- 

tee, Flora of North 

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)lume 22. Oxford Uni^ 

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s, Oxford, UK. Pp. 


BuRQUEZ, A., A. Martinez ^ 

(,and RS.M, 

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adre to the coast: 

changes in vegetatic 

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and R.H. Schmidt, ed 

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especies Mexicanas d 

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ma village o 


lua, Mexico. Phytologia 77:102-140. 

3 Plants. The University of Arizona Press, Tucso 

^(;Vaugh,R. 1993. Flora Novo-Galiciana. Volume 13. Limnocharitaceae toTyphaceae.The 
University of Michigan Herbarium, Ann Arbor, Michigan. 

:aynal,J. 1 976. Notes Cyperologiques: 27. Identification de deux Scleria de Poiret. Adanso- 
nia,ser.2, 16:21 1-217. 

:eina G., A.L.J.R. Van Devender, W.Trauba, and A. Burquez M. 1 999. Caminos de Yecora. Notes 
on the vegetation and flora of Yecora, Sonora. In: D. Vasquez del Castillo, M. Ortega N., 
de la Flora Silvestre de Zonas Aridas, Universidad de Sonora, Hermosillo. Pp. 1 37-144. 

loALSoN, E.H. ]999. Eleochorisyecorensis (Cyperaceae),a new species of spike-sedge from 

PELLENBERG, R.S.,T. Lebgue, and R. Corral D. 1 996. A specimen-based, annotated checklist of 
the plants of Parque NacionaT'Cascada de Basaseachi"and immediately adjacent ar- 
eas, southwestern Chihuahua, Mexico. Listados Floristicos de Mexico, Instituto de 
Biologia, Universidad Nacional Autonoma de Mexico, Mexico, D.F. 

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Books Received/Noticf.s 

Texas Floras/Natural HistoryAVildlife 

•ERICK R Gehlback. 2002 Messages from the Wild: An Almanac of Suburban 

Natural and Unnatural History. (ISBN 0-292-72837-9, hbk ,; 0-292-72838- 
7, pbk.). University of Texas Press, P.O. Box 7819, Austin, TX 78713-7819, 800- 
252-3206, 800-687-6046 fax). $45.00 (libk), $24.95 (pbk), 280 pp., 24 color 
photos, 12 line drawings, 5 1/2" x 9". 

/ORDS: Natural history, Texas, central Texas, environmental studies, animals, migrations of birds, 
;, ecology, life cycles. 

ED Richardson 2002 Wildflowers and Other Plants of Texas Beaches and 

Islands. (ISBN 0-292-77115-0, hbk.; 0-292-7716-9, pbk.). University of Texas 
Press, PO. Box 7819, Austin, TX 78713-7819, U.S.A. (Orders: 800-252-3206 
fax; 512-471-4032). $65.00 (hbk), $29.95 (pbk), 271 pp., 316 color photos, 1 
map, 5 3/4" x 8 1/4". 

Raymond C. Telfair II (ed.). 1999. Texas Wildlife Resources and Land Uses. (ISBN 
0-292-78159-8, pbk.). University of Texas Press, PO. Box 7819, Austin, TX 
78713-7819, U.S.A. (Orders: 800-252-3206 fax; 512-471-4032). $26.95, 416 
pp., 20 b/w photos, 39 tables, 6" x 9". 

Key Words: Texas, wildlife resources, land uses, environmental history conservation, agricultural 
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If you need information at your fingertips on Texas Parks, National Parks, National Wildlife 
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The handy book is divided into five major sections: Part 1) Perspectives on Texas Wildlife Resources; 




Carlos G.Velazco Macias Glafiro J. Alani's Flores 

) Pedro Qarza Garcia U.A.N.L, San Ni 

colds delos Garza 




jo" se ubica en el municipio de Pitiquito y Caborca, Sono 

ra, en la porcion Centre - 

3 sonorense. El complejo orografico Sierra "El Alamo'-Sierr; 


1 Prioritaria Terrestre para la conservacion en Mexico. Este 


locimiento florlstico de las familias Cactaceae y Agavac 


, "El ■ 

itrarian en el area y posteriormente se corroboraron en campo mediante recorridos en diversas 

dades. Se registraron 21 especies de cactaceas distribuidas en 10 generos y 4 especies de Agavaceas 

ceas representan un 21 % del total de las especies presentes en el estado de Sonora. Una especie, 
ictus cyU nd raceus, cuenta con categoria de proteccion. La literatura seiiala que dos de las especies 
ives son endemicas a la Sierra "El Viejo," Agave pdona y A zebra. Se sugiere continuar con estudios 

ion de las poblaciones y el area en 

;rra "El Viejo" is located in the municipality ol Pitiquito and Caborca, Sonora, in the west central 
rtion of the Sonoran Desert; the complex Sierra "El Alamo"-Sierra "El Viejo" is catalogued as a 
-restrial high-priority region for nature conservation in Mexico. The objective for this study is to 
ntnbute to the floristic knowledge of the families Cactaceae and Agavaceae within this region, 
amly in the Sierra "El Viejo." Bibliographical research was carried out and species reported were 
rroborated in the field by searching different localities in the area. Twenty-one species of cacti, in 
genera, and 4 species, and 2 genera of the Agavaceae family were recorded. A total of 25 species 

1 protect 

d ecologic 

;al work 


eded in the region that n 


upport c 




ie dist 






ipios de la 

^Studios t 

que estas familias tienen en el ambito comercial (como plantas de ornato 
altamente cotizadas, medicinales y de alimento), es necesario promover estudios 

La sierra "El Viejo" esta catalogada como una region terrestre prioritaria 
para la conservacion en Mexico (Arriaga et al. 2000; RPT Clave 16). Bajo este 
criterio, consideramos que para la region, existe un conocimiento de escaso a 
regular en cuanto a inventarios de especies, por lo que el desarrollo de listados 
f loristicos y faunisticos es de suma importancia para la correcta planeacion en 
acciones de conservacion y manejo del area. De igual forma, el mismo autor, 
considera que la region se encuentra en un excelente estado de conservacion y 
con una alta integridad ecologica, lo que permite una buena oportunidad para 
llevar con exito planes de conservacion y de manejo. 

1) Ampliar el conocimiento f lorlstico y de distribucion para las especies de la 
familia Cactaceae y Agavaceae en la Sierra "El Viejo." 

2) Aportar algunos datos sobre su etnobotanica e importancia ecologica. 
Area de estudio 

La sierra "El Viejo" se encuentra ubicada en los municipios de Caborca y 
Pitiquito, Sonora, en el noroeste de la republica Mexicana (Fig. 1). Ocupa una 
extension aproximada de 1,128 km^, constituida principalmente por lomerios 
y elevaciones montafiosas que van desde los 200 a los 1,000 msnm. El clima es 
desertico o muy seco semicalido BWhw (x'Xe'). El ciclo de lluvias es bianual y 
se presenta en verano y otono e invierno. La vegetacion de la planicie 
corresponde al matorral desertico microfilo y en la sierra la vegetacion 
corresponde al matorral microfilo sarcocaule. 

Durante los meses de enero a diciembre de 1999 se realizaron recorridos por 
diferentes localidades de la Sierra "EL Viejo" y sus planicies aledaiias. Se 
corroboraron los registros existentes en la literatura y la taxonomia de las 
diferentes especies de cactaceas que se encuentran en esta area. La identificacion 
taxonomica se hizo siguiendo los criterios de Bravo-HoUis (1978), Bravo-Hollis 
y Sanchez-Mejorada (1991) y algunas modificaciones segun Anderson (2001). 

Se encontraron un total de 10 generos y 21 especies para la familia Cactaceae y 

Estados Unidos de America 


Si bien la sierra "El Viejo" 1 

la sido explorada por diversos 


;s (Gen- 

try 1951, 1966; Turner 198C 

), 1983; Dimmitt 1983; Yatskle^ 

/ych y Fische 


1982) es considera por Arn 

iiga (2000), como un area que i 

10 ha sido rev: 


una manera detallada. El c 

onocimiento botanico en esta ; 

area podria ai: 


enormemente, teniendo > 

como ejemplo que Yatskievy 

xh y Fischer 

■ (1984) 

encontraron los siguientes nuevos registros de la tamiiia Cactaceae: 
Ancisirocatus uncinatus, Echinocactus horizontalonius, var. nicholii, 
Echinocereus scopulorum y Mammillaria lasiacantha, de estas especies se ha 
podido corroborar la presencia de todas ellas excepto de E. horizontalomusYa.r. 
nicholii, esta especie fue encontrada originalmente en la porcion norte de la 
sierra, consistiendo su colecta en un solo ejemplar Al examinar ejemplares de 
Mammilaria lasiacntha, se tiene cierta reserva al defmir esta especie, debido 

Cornegiea gigantea (Engelm.) Britton & Rose 1 908 


Echinocereus fendleri (Engelm.) F.Seitz 1870 

Echinocereus nicholii [LD.Benson] B.D.Parfitt 1987 


Echinocereus scopulorum Britton & Rose 1922 


Ferocactus covillei Britton & Rose 1 922 


Ferocactus cylindraceus (Engelm.) Orcutt 1 926 


Hamatocactus uncinatus (Galeotti ) Orcutt 1926 


Lophocereus schottii (Engelm.) Britton & Rose 1 909 


Mammillaria grahamil Engelm. 1 856 


Mammillaria thornberi Orcutt 1 902 


Mammilloria sp. {aff. lashiacantha) 


Opuntia acanthocarpa Engelm. & Bigelow 1856 


Opuntia arbuscula Engelm. 1856 


Opuntia bigelovii Engelm. 1 856 


Opuntio engelmannii Salm-Dyck ex Engelm. 1 850 


Opuntia fulgida Engelm. ]856 


Opuntia leptocaulis A.DC 1828 


Opuntia macrocentra Engelm. 1857 

Nopal n 

Pachycereus pringlei (S.Watson) Britton & Rose 1 909 


Peniocereus striatus (T.Brandegee) Buxbaum 1975 


Stenocereus thurberi (Engelm.) Buxbaum 1961 


TABLA2.Especiesde Agava 

Nombre Cientifico 

sten diferencias en el color de la f lor y la espmacion, por lo cual actualmente 
mcuentran en revision para determinar su categoria taxonomica de una 

Comparativamente, la Sierra El Viejo se puede considerar rica en especies 
cactaceas, ya que Felger (1992), al realizar una sinopsis de las plantas 
culares del noroeste de Sonora, aproximadamente 15,000 km^ (incluyendo 
■eserva de la biosfera "El Pinacate"), registra 10 generos, 26 especies y 7 
iedades (reconocidas por Anderson 2001). En la Sierra "El Viejo," area 13 veces 
nor, se senalan igual numero de generos y cinco especies menos; para la fa- 
ia Agavaceae solo existen 2 generos con una especie cada uno. 

Hernandez (1998) caracterizo la vegetacion en el area de la Sierra "El Viejo," 
y registro nueve cactaceas, ademas de Agavt sp. y Yucca sp. sin profundizar en 
su categoria taxonomica. El presente estudio arroja mas del doble de especies 
de cactaceas y determina las especies de agaves mediante la revision de literatura 
ya existente. Cabe destacar que dos especies (Agave pelona y A. zebra) son 
endemicas de la Sierra «E1 Viejo» y por lo menos A. pelona es parte importante 
de la dieta y fuente de agua para el borrego cimarron (Ovis canadensis mexicana) 
que habita la region. Precisamente es con el borrego cimarron, con quien se 
observe un fenomeno interesante de interaccion entre fauna y flora, durante la 
epoca de f loracion de A. pelona, se da un alto consume de los escapes f lorales de 
esta especie por parte del borrego cimarron y a lo largo del ano se consumen 
plantas enteras para satisfacer las necesidades tanto alimenticias como hidricas. 

Cabe destacar que del total de las especies de cactaceas solo una, Ferocactus 
cylindraceus, esta dentro de la norma NOM-059-ECOL-1994 (Diario Oficial de 
la Federacion 1994), bajo la categoria de rara y que ninguna de las especies de 
agavaceas esta en alguna categoria de dicha norma, a pesar de que dos de ellas 
son endemicas del area. 

Historicamente y de acuerdo con los pobladores locales, se ha realizado 
extraccion de Agave pelona y A. zehra. El primero para la extraccion de fibras 
vegetales y el segundo para la elaboracion, a pequena escala, de una bebida 
alcoholica. A pesar de dicho aprovechamiento (realizado en la primera mitad 
del siglo XX), las poblaciones se encuentran en un buen estado, ya que se 
observan mdividuos de Agave de todos los tamanos y clases de edades. 

La Sierra "El Viejo," area prioritaria para la conservacion en Mexico, 
mantiene dentro de sus limites un total de 21 especies de cactaceas distribuidas 
en diez generos, siendo Opuntia el genero (7 spp.) mas diverse, seguido por 
Echinocereus y Mammillaria (3 spp.). Esto representa apreximadamente un 21 
% del total de especies registradas para el estado de Sonora (Paredes et al. 2000). 
Las especies de la familia Agavaceae estan representadas por el genero Agave (3 
spp.) y el genero Yucca (1 sp.). 

Nuestro mas sincere agradecimiento a la Biol. Liliana Ramirez Freire y Erika 

■abajos de ca 

.mpo. Org 

us Gutierrez y Mar 
;anizaci6n Vida S 



Qenta que cooper 
■e A.C. por las f 






Bravo, H.H. y H. Sanchi 

DiARio Oficial de la FederaciOn. 1 994. Norma Oficial Mexicana NOM-ECOL-059.Secretaria de 
Desarrollo Social. Tomo CDLXXXVIIi. No. 1 0. Mayo 1 6 de 1 994, Mexico. 

Felger, R.S. 1 992. Synopsis of the vascular plants of northwestern Sonora, Mexico. Ecologica. 

Hernandez, C.M.C. 1 998.Caracterizaci6n de la vegetacion del predio"EI Plomito,"municipio 
de Pitiquito, Sonora, Mexico.Tesis inedita. Facultad de Ciencias Biologicas, Universidad 

Paredes A,T.R.Van Devender, y R.S. Felger. 2000. Cactaceas de Sonora, Mexico; su diversidad, 

uso y conservacion. IMADES-ASDM Press, Canada. 
Turner, R.M.,J.E. Bowers yT.L Burgess. 1995.Sonoran Desert plants: an ecological atlas.The 

University of Arizona Press, Tucson. 
Yatskievych, G.y PC. Fischer. 1 984. New plant records from the Sonoran Desert. Desert Plants 





Kay M.Fleming 

Jason R. Singhurst Walter C Holmes 

Wildlife Diversity Program Department of Biology 

3000 South tH-35, Sur, 

a bottomland hardwood forest 

d by eight distinct alliances:!) 
tonwood; 5) Sugarberry-Cedar 
id Post Oak-Bluejack Oak. The 

The Texas Parks and Wildlife Department began purchasing bottomland in 1991 
m an effort to preserve this unique and rapidly disappearing hardwood habitat. 
Among the first acquisitions was the area now known as Big Lake Bottom Wild- 
life Management Area (BLBWMA), one of the largest remaining bottomland 
hardwood tracts of its type in the central Trinity River basin. The area consists 
of approximately 1684.8 ha that lie within the transition zone between the 
Pineywoods and the Post Oak Savannah vegetational areas of eastern Texas 
(Gould 1962), about 16 km S W of Palestine. This alluvial bottomland was formed 

by the periodic erosion and sedimentation of the river as it flowed through the 
region. The poorly draining soils and uniform topography produced a forest 
dominated by hydrophytes. 

Because of its unsuitability for most agricultural pursuits, the management 
area has remained relatively free from outside disturbances. Two of the past 
major disturbances were selective harvesting of timber in the late 1930s and 
early 1940s and the clearing of about 65 ha for cultivation in the 1950s. Current 
use of the property is limited to human recreational activities. Past and present 
water control and agricultural practices have generally widened the Trinity 
River, resulting in severe erosion problems along sloughs and creeks. This ero- 
sion problem has been compounded by the use of all-terrain vehicles on the 
management area. 

Since the management area was purchased to preserve the bottomland 
habitat along the Trinity River, this systematic inventory of the vascular flora 
is imperative for proper management. Results of the study will be used by re- 
source managers in decisions concerning activities that might affect the plant 
communities and ecosystems of the area. These activities include future land 
purchases or trades, mineral exploration, granting of rights-of-way and ease- 
ments, and public use. 

Field work was conducted primarily from October 1997 to October 1999, with 
occasional visits made until July 2001 to search for taxa expected to occur but 
not previously collected. The survey included more than 50 field visits that pro- 
duced 532 field collections including ferns, gymnosperms, and flowering plants. 
Voucher specimens are deposited m the Baylor University Herbarium (BAYLU). 


BLBWMA is located within the humid subtropical zone. Temperatures rarely 
go below -12° C in winter or above 38° C in summer, with the mean temperature 
being near 20° C (Hatch et al. 1990). The soil of the study area is predominately 
composed of alluvium and fluviatile terrace deposits and a small area of car- 
bonaceous clay, silt, and ironstone on the Recklaw Formation (University of 
Texas Bureau of Economic Geology-Palestine Sheet 1993). The deep clay bot- 
tomland soils are classified as part of the Kaufman-Trinity Association while 
the more loamy and sandy uplands belong to the Axtell-Lufkin Association 
(Coffee 1970). Elevation varies from 57.9-82.3 m above sea level. 

Nixon and WiUett (1974), in the only previous study of the site located, 
described 5 bottomland hardwoods types. These were described as coalitions 
of species and not afforded names as such. Many species were included in more 
than one type, apparently based upon abundance. These types actually over- 
lapped to some extent, thus we find it advantageous to use the alliance method 
of community description. 

The BLBWMA vegetation areas can be segregated into eight alhances (Na- 
tional Vegetation Classification System (NVCS) 1997) based on dominant spe- 
cies, landscape position, and soil water content. The location, characteristic 
species, and a short description of each alliance follow. 
Overcup Oak (Water Hickory) Seasonally Flooded Forest Alliance 
The Overcup Oak (Water Hickory) Alliance is characterized by temporary 
standing water during and after flood events and consists of approximately 
818.3 ha. This alliance is found throughout the site on heavy clay soils (Fig, 1). 
Dominant trees are Quercus lyrata, Carya aquatica, Celtis laevigata var. laevigata 
and Ulmus americana. Other common woody species in this area include 
Fraxinus pennsylvanica, Ilex decidua, and Crataegus viridis. Common herba- 
ceous species include Iva annua, Xanthium strumariumjusticia ovata, Leersia 
knticularis, Saururuscernuus, Lobelia cardinalis, Diodia virginiana, Gratiola 

Willow Oak Seasonally Flooded Forest Alliance 

Willow Oak Alliance consists of seasonally flooded areas and is the smallest 
wet bottomland feature on the area at 50.2 ha. These are depressed areas of pe- 
riodic inundation in which hardwood trees dominate and form a dense canopy. 
This alliance is located north of the drainage channel of Big Lake into Keechie 
Creek, an area that has been carved out by flooding events (Fig. 1). Common 
tree species include Quercus phellos, Q. nigra, Celtis laevigata var. laevigata, 
Carya aquatica, Nyssa sylvatica, Ilex decidua, and Liquidamhar styracijlua. 
Characteristic herbs of this forest include Carexfrankii,Juncus coriaceus, and 
Trachelospermum dijforme. 

Planer Tree Seasonally Flooded Forest Alliance 

Planer Tree Alliance comprises 85.8 ha and consists of mid-story forests in 
backswamps and sloughs that are produced in soils created by rapid siltation. 
This alliance occurs along the corridor of Cedar Creek Slough and along Keechie 
Creek Slough just south of where Keechie Creek joins the Trinity River, as well 
as m a small section of the Johnson Tract (Fig. 1). Common woody plants in- 
clude Planera aquatica, Gleditsia aquatica, Forestiera acuminata, Carya 
aquatica, Cephalanthusoccidentalis, Fraxinus pennsylvanica, Salix nigra, and 
Berchemia scandens. Herbaceous plants include Saururus cernuus, Boehmeria 
cylindica, Seshania drummondii, and 5. exaltata. 
Cottonwood Temporarily Flooded Forest Alliance 

Cottonwood Alliance consists of 10.9 ha of periodically flooded forests on sandy 
alluvium primarily on developed river fronts. It is restricted to a narrow strip 
adjacent to the Trinity River bank (Fig. 1). Common tree and shrub species in- 
clude Populus deltoides, Salix nigra, Acer negundo, Gleditsia aquatica, Celtis 
laevigata var. laevigata, Betula nigra, Ulmus americana, Morus rubra, Cornus 


Vegetation Alliances 

^H Overcup Oak - Water Hickory 
iiii Willow Oak 
^^^ Planer Tree 

lllllflllli Sugarberry-CedarElm 

p showing Big Lake Bottom lA 

mondii, Symphoricarpos orhin 
and woody vines are well repre 
ens, Vitis riparia, V.palmata, 5 

I. lacu 

lulus, and Vovcsticra acuminata. Herba- 
ented. Common species include Mikania 
rilax hona-nox, S. tamnoides, Ampelopsis 

Sugarberry-Cedar Elm Temporarily Flooded Forest Alliance 

Sugarberry-Cedar Elm Alliance consists of periodically flooded, forested, flat 
areas with rich base soils. This alliance occupies 412.4 ha along Cedar Creek 
Slough and along drainages of Keechie Creek (Fig. I). Common trees include 
Celtis laevigata var. laevigata, Ulmus crassifolia, U. alata, Gleditsia triacanthos, 
Sapindus saponaria var. drummondii, Quercus macrocarpa, Fraxinus 

pennsyhanica, F. americana, and Crataegus mollis. Herbaceous plants which 
typify the area include Iva annua, Carex cherokeensis, C. crus-corvU Fanicum 
a nceps, and Leersia oryzoides. 
Bur Oak-Shumard Oak Forest Alliance 

Bur Oak-Shumard Oak Alliance occurs on bluffs and terraces with clay soils 
and rich humus layers. This alliance also occurs on two bluff areas that origi- 
nated as a result of severe changes in river direction (Fig. 1). The area occupies 
178.9 ha. This alliance was not formally described as occurring in Texas by the 
N VCS (1997) or as a series level community by Diamond et al. (1987) and conse- 
quently is mentioned here as occurring in Texas for the first time. Common 
trees and shrubs include Quercus macrocarpa var macrocarpa, Q. shumardii, Q. 
stellata, Sideroxylon lanuginosum, Sophora affinisjuniperus virginiana, 
Crataegus viridis var viridis, Celtis laevigata var laevigata, Ilex decidua, and 
Cercis canadensis. Herbaceous plants which typify the area include Carex 
cherokeensis, Scleria triglomerata, Chasmanthium latijolium, Poa autumnalis, 
Tradescantiaohiensis, Arisaema dracontiuva, Podophyllum peltatum, Dioscorea 
quaternata, Clematis pitcheri, Urtica chamaedryoides, Stachys crenata, S. 
tenuifolia, Ruellia nudiflora var runyonii, Elephantopus carolinianus, Matelea 
gonocarpos, Myosotis macrosperma, Scutellaria cardiophylla. Euphorbia 
spathulata, Galium circaezans, Oxalis violacea, and Parietaria pensylvanica. 
Post Oak-Blackjack Oak Woodland Alliance 

Post Oak-Blackjack Oak Woodland Alliance comprises 21.5 ha and consists of 
upland wooded areas on gentle sandy-loam slopes and abrupt deep sand ter- 
races. This alliance is very species rich and found on the small Johnson Tract 
and near entrances at each of the area gates (Fig. 1). Common trees and shrubs 
include Quercus stellata, Q. marilandica, Q. incana, Q.falcata, Q. nigra, Q. phellos, 
Cercis canadensis, Ulmus alata, Fraxinus americana, Callicarpa americana, 
Crataegus crus-galli, C. marshalli, Prunus angustifolia var. angustifolia, P. 
mexicana, P. serotina var. serotina, Ruhus trivialis, Rhus copallina, and 
Symphoricarpos orhiculatus. Herbaceous species include Chasmanthium 
descantia hirsutiflora, Vitis mustangensis, V. rotundijolia, Sanicula canadensis, 
Symphyotrichum patens, Podophyllum peltatum, Hypericum hypericoides, Hy- 
pericum drummondii, Passijlora incarnata, and Geum canadense. 
Sand Post Oak-Bluejack Oak Woodland Alliance 

This alliance occurs in deep sandy soils on a ridge above the f loodplain. It con- 
sists of less than 5 ha (Fig. 1) located on the east side of Big Lake. Common tree 
species include Quercus margarettiae, Q. marilandica, Q. incana, and Carya 
texana. Other woody plants are Opuntia stricta, Yucca louisianensis, Rhus 
copallinavar.latifolia,Cornusflorida,Vaccinium arhoreum. Sassafras alhidum, 

and Prunus gracilis. Herbaceous plants include Tradescantia hirsutiflora, 
Matelea cynanchoides, Eragrotis secundijlora, Loejlingia squarrosa, 
Helianthemum rosmarinifolium, Lechea tenuijolia, Cnidoscolus texanus, and 
Dalea villosa. 

The BLBWMA flora includes 459 species in 298 genera in 99 families. Average 
number of species per genus is approximately 1.5. Families with the largest 
number of species (in parentheses) are Poaceae (55), Asteraceae (54), Fabaceae 
(32) and Cyperaceae (25). Other families with a relatively large number of spe- 
cies are Lamiaceae (14), Apiaceae (12), Euphorbiaceae (12), Rosaceae (12), and 
Scrophulariaceae (10). Genera with the most species include Carex (12), Quercus 
(9), Panicum (8), Cyperus (7), Smilax (6), and Juncus (5). The number of non- 
native species is 23, which is about five percent of the total flora. 

Three plants collected on BLBWMA are considered threatened by the Texas 
Parks and Wildlife Department and The Nature Conservancy of Texas (Carr 
2001). Brazoria truncata var. pulcherrima (B. pulcherrima Lundell) is consid- 
ered sensitive and ranked G3 (globally) and S3 (state). Both it and Yucca 
louisianensis are also notable for being included in Estill and Cruzan's (2001) 
list of narrow endemics of southeastern United States. Curcurhita texana is 
also ranked G3 (globally) and S3 (state). The species was historically widely 
distributed along major rivers in the eastern half of Texas but much of its habi- 
tat has been lost to impoundments, agriculture, and urban development. An 
accurate assessment of its current status is difficult because of its ephemeral 
nature. Cyperus grayoides is ranked G3 (globally) and S3 (state). 

Other noteworthy plants were new to Anderson County Correll and 
Johnston (1970) report Dioscorea quaternata as mainly occurring in more north- 
eastern Texas. The BLBWMA record is near the most southwestern limit of the 
species distribution. Symphyotrichum eulae, a Texas endemic, also occurs in 
the area. The distribution of this species is primarily in east-central Texas, where 
it is considered rare or uncommon. Rhynchosia minima var. minima is known 
from the coastal plain and northward to Newton, Houston, Travis, Bexar, Rob- 
inson, and Grimes counties. The present record is the northernmost record for 
the species within the state. Heliotropium procumhens is mostly a coastal (and 
western) species known to occur northward to Liberty Waller, and Brazos coun- 
ties. The new record extends the known distribution within the state farther 
northward. Finally Phyllanthus pudens has been recorded in Texas as far north 
as Brazos, Madison, and Robinson counties. The Anderson County record ex- 
tends the distribution farther withm the state and provides a new county record. 

The area contains the only known occurrence of Bur Oak-Shumard Oak 
Forest alhance within the state (see above). 

The annotated checklist is divided into ptendophytes, gymnosperms, and flow- 
ering plants, which are subdivided into monocots and dicots. Family genus, 
and species are arranged alphabetically beneath each major heading. Nomen- 
clature follows that of Correll and Johnston (1970), Correll and Correll (1972), 
and Gould (1975). As needed, names were updated using Jones et al. (1997). Com- 
mon names generally follow those of Hatch et al. (1990). Collector and collec- 
tion numbers are given as: KF = Kay Fleming or JS =Jason Smghurst followed by 
a number. An asterisk (*) denotes an introduced species. 





Ophioglossum vulgatL 

ero/ceens/s Schwein., Cherokee sedgeJS 
mplanata Torr. & Hook., Sedge, KF 457. 




)reng.) Fassett, Beaked 

1 Dewey, Thinfruit sedge, KF 

, Frank's sedge, KF 452. 

& Hook,, Sharpscale sedge, KF 

Carex texensis (Torr.) Baiiey, Texas sedge, KF 286. 
Cyperuscrocec/sVahl, Baldwin flatsedge,/<F45/. 
Cyperus erythrorhizos Muhl., Redroot flatsedge, 
KF 636. 

Cyperus strigosus L.var.smgosus, False nutsedge, 

Cyperus surinamensis Rottb., Tropical flatsedge, 

Eleocharis microcarpa Jon.,Sma\\seed 

f/eoc/ior/sobfusa (Willd.) Schult.,Blunt 

Eleocharis palustris (L.) Roem.& Scliult, 

Rhynchospora corriiculata (Lam.) Gt< 

beakrush, KF 415. 
Scirpuspendulus Muhl., Small bulrush 
Scleria triglomerata Michx.,Tall nutru: 


Iris brevicaulis F 
Sisyrinchium alt 

^ flatgra 

1 langloisii Greene, Pale t 

:3 (Small) Ames, Spur 


1 Bicknell, Blue-eyed Bronn 


ichloedactyloides (Nutt.) Engelm.,Buffalogra 

Cenc/irus sp/nZ/ex A. Cav., Coastal sandbur,( 

Chasmanthium sessiliflorum (Poir.) Yates, : 

s (Retz,) 

Carolina lovegrass^Kf 660. l/u/p/oocfof/ora (Walt.) Rydb.,Common sixweeks 

•rostis secundiflora Presl.subsp. oxy/ep/s (Torr.) grass, KF 403. 

.D.Koch,RedlovegrassKfm SMILACACEAE 

/rosf/s spectabilis (Pursh) Steud., Purple smilaxbona-noxL,Sawtoolhgteenbr\ar,KF284. 


a Willd., White cutgrass, /<F 673. Typha latifolia L., Common cattail, KF 4 
e L., Perennial ryegrass, JS 6670.* DICOTYLEDONEAE 

etteqrassKFi/6. ACANTHACEAE 

jmon Schult., Maiden cane, KF runyonii (Tharp. & Barkl.) B.L Turr 

ruellia, JS 63/5. 
(VScribn.& Merrill, Ravenel's pani- /^ue///o pedunculata Torr, ex Gray, Low 


Acer negundo L van negum 

Froelichia drummondii t 

Piptochoetium avenaceum (L.) Parodi, Blackseed ^""''^ ' ^ 

needlegrass, KF 300. ANACARDIACEAE 

Poa annua L., Annual bluegrass, KF 505.* Rhuscopallina Lvar. latifolia H. Engler,Wing-ri 

bristlegrass, KF 637. APIACEAE (UMBELLIFERAE) 

Sorghum ho/epen5e(L.)Pers.,Johnstongrass,KF chaerophyllumtaintunenHook.,Cher.VS616 
Cyclospermum leptophyllum (Pers.) Sprague e 

Fndens albescens (Vasey) Wool & Standi., White g^.^^^^ RWils.,SI,mlobe celery, KF 465. 

scaleseedKf 3. 


Scaleseed, KF 287. 
Trepocarpusaethusae Nutt.ex DCJrepocarp 


Amsonia illustris Woods., Bluestar, KF 247. 
Trachelospermum difforme (Walt.) Gray,Climbi 

dogbane, Kf 420, 

Nutt.) Raf, Scratch daisy, 


) Cabrera, Greenk 

jsonnuus L, Common 51 

;' hawkweed, K 

>ides E. Sherff, 
, Small head 

f.) Chamb., Weedy dwarf dan- 
J Gaertn., Florida wild lettuce, 

,) Michx., Variable gayfeather. 


w (DC.) C.Jeffrey, Golden 


ra(L.) DC, Camphor weedKf ''"' 5chulz'"sa"nd'bittercress"f(f 4( 

//n/anus (Wait.) DC, Carolina <^Pej^^^^^ir9'ncum ., 

^'''''\ RonppapalustnsiDBesser.Bogf,. 

.ate coneflower, KF 383. ^^^.^^^ sessiliflora (Nutt. ex T. & G.; 

us Hook., Texas groundsel, JS ^^^^ yellowcress, JS 6304. 

,s L.,Tall goldenrod,/<f 632. BUDDLEJACEAE 

\it.. Downy golden rod, /<f 665. Polypremum procumbens UJunip 

lolidago radula Nutt., Rough goldenrod, KF 627. CACTACEAE 

:o//dagou/ni/fo//aMuhl. ex Willd., Noble golden- Opuntia striaa (Haw.) Haw.,Soutl 

md,KF658. cactus, KF 393. 

.olivopterosperma (Juss.) Less., Lawn burweed, (-y^LLURicHACEAE 

^^^' Viburnum rufidulum Raf, Rusty blackhaw,/<f 2^ 


Fodophyllumpeltatum L., May appe, 9. Arenaria serpyllifolia Lsuhsp.serpyllifolla.Jhyrr 

BETULACEAE leaved sandwort, KF 502.* 


BORAGINACEAE ''' Z'!!!!c\ZZZaSF475. 

mthogynus Michx,One seed croton. 
entata Michx, Toothed spurge, Kf 
'jtans Lag., Eyebane spurge, KF 597. 


p/iorbfosparhu/ofa Lam., Warty spurge, KF 469. 
y//anr/iuspudens Wheeler, Birdseed leaf flower, 

3g/a brevispica Engelm. & Gray, Heartleaf 

drew's FABACEAE 



wood, Kf 256. 
Cornusflorida L., Flowering dogwoodJS 6? 79. 


illiana Small, Nuttall's wild indigo, 
irginianum (L) Benth., Butterfly pea, 

Clitoria moriana L.,Atlantic pigeon wings, /(f 967. 
Do/eo w7/osa(Nutt.)Spreng.,Prairieclover,/<f 350. 
Desmanthus illinoensis (Michx.) MacM. ex Rob- 
ins & Fern., Illinois bundle flower, KF 6 15. 

olium DC, Sessileleaf beggar's 


Vaccinium arboreum Marsh., Farkleberr 

Acalypha gracilens Gray, Slender copp' 

_., Small bur clover, /<f 238. nemophila, KF 288 


venw.) Benth., Yellow Caryaoquar/ca (Michx) Nutt Water hickory W 

1 (T. & G.) Isely, Palmleaf Coryo fexano Buckl., Black hickory, KFJ/J. 

Juglans nigra L.Black walnut, KF 268. 

ink, Coffee senna,/<f 1022. Brazoria truncata (Benth.) Engelm. & Gray var 
(Rydb.) Cory, Rattlebush, pulcherrima (Lundell) M.W.Turner, Centerville 


ina clover, /<F237. Monarda fistulosa L.,Wild horsemint,/(F4S0. 

Arrowleaf clover, KF 340.* Physostegia intermedia (Nutt.) Engelm. & Gray, 
Vtcia ludoviciana Nutt., Deer pea,KF291. Intermediate dragonhead, /(F 4/6. 

FAGACEAE ' ' Physostegia virginiana (L.) Benth., Virginia drag- 

rrt^jMCtAt on head, JS 6697. 

Quercus falcata Michx.^Southern red oaKKF513. ^^^^.^ ,^^^^^ ^ _ ^y^^,^^^ ^^^^^ ^^ ^^^^ 

Ouercus incana Bartr., Bluejack oak, /<F 376. Scutellaria cardiophylla Engelm.&Gray.heanleaf 
Ouercus/yrafa Walt., Overcup oak, /(F 507. ri.,,iirpn /('f.i?/^ 


OuercusstellataV^ana Post oak KF 374 



Sassafras albidum (Nun.) Uees,^^ 



^'"TaxTpIgs" ^"^'^'^" ""'' ""'' 

Geranium carolinianum L. var.coro//n/anum, Caro- 

lina geranium, /<F 255. 



/^mmann/acocc/neo Rottb.,Purp 

Mynophyllum heterophyllum Michx.,Changeleaf 


parrot's feather, /(F 432. 

yt rum^aatum 




Hibiscus moscheutos L. subsp. la 


Mollugo verticillata L.Jhread ca 

Morus rubra L, Red mulberry, K 


LIgustrum sinense Lour., Chinese p 

Gaura longiflora 5pach.,Tall guara 
Qaura parviflora Dougl. ex Lehm. 


Argemone albiflora Hornem. var. texam 
Owenby) Shinners, White prickly popf 




Polygala polygama L. var. obtusa Chodat, [ 


Polygonum aviculare L, Prostrate knotwee 

Polygonum hydropiper L., Water smartwee 

Polygonum pensylvanicum (L.) Small, Penns 

nia smartweed, KF 534. 
Polygonum persicana L, Lady's thumb, KF 6 




Oenothera speaosa Nutt., 5howy ^ 

rose, /(F 270. 


hrangulacamlmiana {Walt) Gray.CaroWnabuck- 




Crofaegus cru5-ga/// L.CockspurhawthoraKf 295. 

Crateagus morshollii EggL, Parsley hawthora KF 

rie agalinis, /<F6/6. 

Crataegus mollis Scheele, Downy hawthorn, /<F475. 

Castilleja Indivisa Engelm., Texas Indian paint- 

Crataegus wnd/sL.subsp.vindis,Green hawthora 


Geum canac/eme Jacq., White avenUf 290, 

Chickasaw plum, JS 6/33. 

hedge hyssop, KF 3/3. 

Prunus gracilis Engelm. & Gray, Sand plum, KF 

Nuttallanthus canadensis (L.) Sutton, Old field 



Verbasclmtha sus L Common mullein KF 720 


Veronica agrestis L., Wayside purslane, J5 6/73. 

Veronica peregrina L.,Purslane speedwell,KF 304. 



Physalis angulata L, Cutleaf ground cherry, KF 

Benth., Buttonbush, JS 6674. 
Diodia teres Walt. var. teres, Rough buttonweed, 

F/iyso/;5 mo///s Nutt., Field ground cherry, KF 670. 
shade, KF 677 

DIodia virginlana L., Virginia buttonweed, KF530. 

Galium aparine L.Catchweed bedstraw, KF 498. 

horse nettle, KF 337 

Galium circaezans Michx., Woods bedstraw, KF 

Solanum elaeagnifolium Gay., Silverleaf night- 

Galium tinctonum (L.) J. Scopoli, Dye bedstraw, 



Houstonia pusilla J. Schopf, Small bluets, KF 229. 

F/7/a americana L. var. caroliniana (Mill.) 

0/den/and/tosc// (DC.) Chapm., Bluets, KF 228. 

Castiglioni, American basswood, KF 527. 




Geitis laevigata Willd. var. /aewgo fa, Texas sugar- 


Planera aquatica (Walt) Gmel Water elm J5 

Populus deltoides Bart, ex Marsh.subsp.c/e/to/des, 

Ulnius^alata M\chx Win ed elm KF5/2 

Sa//xn;gra Marsh., Black willow, JS 6696. 

Ulm^us Americana L, American Tl m, KF 509. 


Ulmus crassifolia Nutt., Cedar elm, KF 508. 

Cardiospermumhalicacabum L., Balloon vine,/<F 


Boehmeriacylindrica (L.) 5w.,False-nettle, KF392. 

Saplndus saponaria L. var. drummondii (Hook. & 



Carolina violet, Kf 759. 

ufr., Beaked cc 

irn salad, 



Phorodendron to 

mentosum (DC.) Engeir 



js mistletoe, KF 7/8. 

Callicarpa americana L„ / 

Vmerican beai 




Ampelopsis arbort 

=0 (L.) Koehne, Peppervir 

We want to thank Ron George and Gary Graham of the Texas Parks and Wild- 
Ufe Department and Wildhfe Division Region 3 staff for their support on this 
basehne inventory cooperative project. We are also grateful to Tom Wendt of 
TEX/LL for his assistance. Also, special thanks to Changxiang Liu, Kim Ludeke, 
Vivian Ackerson, Michelle Valek, and Duane German of the Texas Parks and 
Wildlife Department Geographic Information Systems Laboratory for their as- 
sistance in the vegetation alliance mapping. Thanks to David Riskind, Coordi- 
nator of the Natural Resource Program, Texas Parks and Wildlife Department, 
and Heather White of GREE who read and commented on preliminary ver- 
sions of the manuscript. We are also grateful to Guy Nesom and Monique Reed 
for their review of the manuscript. Their comments were exceedingly helpful. 


ted list of the G3m and rarer plant taxa ofTexas. 
Nature Conservancy ofTexas, Austin. 
Coffee, D.R. 1 970. Soil survey of Anderson County, Texas. Natural Resource Conservation 

CoRRELL, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner. 
CoRRELL, D.S. and H.B. Correll. 1 972. Aquatic and wetland plants of southwestern United 

States. U.S. Government Printing Office, Washington, D.C. 
Diamond, D.D., D.H. Riskind, and S.L Orzell. 1 987. A framework for plant community classifi- 
cation and conservation in Texas.Texas J. Sci. 39: 203-22 1 . 
Estill, J.C. and M.B. Cruzan. 2001 . Phytogeography of rare plant species endemic to the 

:ouLD, F.W. 1 962.Texas plants - a checklist and ecological summary.Texas Agric Exp. Stat. 

Publ. MP-585Jexas A&M University, College Station. 
iouLD, F.W. 1975. The grasses ofTexas.Texas A&M University, College Station. 
Iatch, S.L, K.N, Gandhi, and L.E. Brown. 1 990. Checklist of the vascular plants ofTexas.Texas 

Agric, Exp, Stat. Publ, MP-1 655,Texas A&M University, College Station, 
DNFS, S,D,, J,K.WiPFF, and RM. Montgomery, 1997. Vascular plants of Texas: a comprehensive 

checklist including synonymy bibliography, and index. University of Texas Press, Austin. 
I ATioNAL Vegetation Classification System-Oklahoma-Texas Subset, 1 997,The Nature Conservancy 

Conservation Science Department, Southeast Region, Chapel Hill, North Carolina, 
lixoN, E,S, and R,L, Willett, 1 974. (unpublished) A vegetative analysis of the floodplain of 

the Trinity River, Stephen F, Austin State University, Nacogdoches, Texas, 
Iniversity of Texas Bureau of Economic Geology-Palestine Sheet, 1 993, Map o 

Books Recrived/Notic 

Texas Floras/Natural History/Wildlife 

T Wi iiTH (Foreword by Greg W. Lasley). 2002. Birds of Northeast Texas. (ISBN 
0-58544-192-9, hbk .; 0-58544-193-7, pbk.). Texas A&M University Press, 
John H. Lmdsey bldg, Lewis Street, 4354 TAMU, College Station, TX 77843- 
4354, U.S,A, (Orders: 800-826-8911, 979-847-8752 fax,, $44.95 (hbk), $19.95 (pbk), 160 pp., 30 color pho- 
tos, map, 6" X 9". 

VoRDs: Texas, northeast Texas, birds, identification, field guide, zoology. 

Contents include an introduction followed by chapters on Geography of the Region, Birdwatching 
Areas, and Species Accounts. The color photos are very nice. 

John W. Tunnell.Jr. and Frank W.Judd (Foreword by Richard C. Bartlett). 2002. 
The Laguna Madre of Texas and Tamaulipas. (ISBN 0-58544-133-3, hbk.). 
Texas A&M University Press,John H. Lmdsey bldg, Lewis Street, 4354 TAMU, 
College Station, TX 77843-4354, U.S.A. (Orders: 800-826-8911, 979-847-8752 
fax,, $60,00, 372 pp., 83 
color photos, 87 figures, 45 tables, 8 1/2" x 11". 

e fisheries organ- 

idix of Laguna Madre researchers and mterested parties. The literature cited is 30 pages of ref- 

nar Brune (Introduction by Helen C. Bese). 2002. Springs of Texas. Second 

Editon. (ISBN 0-58544-196-1, hbk.). Texas A&M University Press, John H. 
Lindsay bldg, Lewis Street, 4354 TAMU, College Station, TX 77843-4354, 
USA (Orders 800-826-891L 979-847-8752 fax,, 
wwwtamuedu/upress) $75 00 (hbk), 608 pp., 272 b/w photos, 38 line 
drawings 9 colot maps, 8 1/2' x 11" 

£ Springs, Prairie Dog Springs, 


Barbara R.MacRoberts 
and Michael H. MacRoberts James C Cathey 

Bog Research, 740 Columbia Qus Engeling Wildlife Management A 

Shrevepon, LA, 71 104, U.S.A. Texas Porks and Wildlife Departmer 

Herbarium, Museum of Life Sciences Tennessee Colony, TX 7586 1, U.S.A. 

5avanna, flora, Texas, Gus Engeling Wildlife Management . 

I, suelos, y riqueza de especies a pequefia escala de los lu| 

During the past few years, we have been studying wetlands (muck bogs, upland 
marshes, baygalls, and seeps) in the post oak savanna region of east central Texas 
and southeastern Oklahoma (Nesom et al. 1997; MacRoberts 6a: MacRoberts 
1998b), These wetlands appear to be associated with xeric sandylands (vari- 
ously referred to as oak-farkleberry sandylands [Ajilvsgi 1979], Post Oak-Black 
Hickory Series [Diamond et al. 1987], xeric sandylands [MacRoberts & 
MacRoberts 1994, 1995, 1996], Grossarenic Dry Uplands [Turner et al. 19991 Sand 
Post Oak - Bluejack Oak Alliance [Singhurst et al. 20001 and Quercus incana 
woodland alliance [Hoagland 2000]). These deep sands act as a reservoir or 
sponge holding water that feeds adjacent seeps and springs that are the head- 
waters for the area's wetlands and ultimately the streams and rivers. These 
upslope soils are porous and drain readily; rainwater percolates through the 
sand and moves down a gradient created by underlying impermeable or slowly 
permeable clays. Eventually, water seeps laterally out of the hillside (Martin & 
Smith 1991; Jones & Carpenter 1995; Drewa 1999; Summer 1999), 

As part of our study of muck bogs, upland marshes, and their flora, we 

studied these adjacent xeric sandylands since they are cleady the water source 
for these wetlands. 

The primary objectives of this paper are to describe: 1) the distribution of 
xeric sandylands throughout the West Gulf Coastal Plain, 2) the floristics of 
this community in the post oak savanna region, 3) the small-scale species rich- 
ness of this community, and 4) the soils upon which this community occurs. 

In addition to our primary objectives for studying this community we also 
are stimulated by a recent spate of national and regional conservation assess- 
ments (e.g.. Diamond et al. 1997; Ricketts et al. 1999a, 1999b; Myers et al. 2000) 
that propose to pinpoint ecological "hotspots." These assessments are based on 
diversity, species richness, endemism, endangered species, and unique habitat 
availability. But, as is so often the case with such large-scale efforts, while some 
regions are well known, others are not. The post oak savanna region of east Texas 
is one of the poorest known. This is evident upon examining recent regional 
descriptions of the area (e.g., McNab & Avers 1994; Keys et al. 1995; Ricketts et al 
1999a), where numerous factual errors lead to inaccurate assessments of the 
region's ecological "temperature." 

In Texas, the post oak savanna region (Fig. 1), consisting of about 30,000 sq, km, 
is gently rolling and hilly with elevations from 90 to 250 meters. Rainfall ranges 
from 75 to 115 centimeters. The Carrizo Sands are virtually coterminous with 
the region (McBryde 1933). Prairies are scattered throughout, notably in the 
south where some prairies are large (Smeins & Diamond 1983). The predomi- 
nant floristic character of the region is southeastern without pines (Smeins & 
Diamond 1986). The area contains a diversity of plant communities, from hill- 
side pitcher plant bogs, peat bogs, and upland marshes to open xeric sandylands 
and oak-hickory forests and woodlands. Characteristic communities within the 
post oak savanna region also occur to the east within the pmey woods (Marietta 
& Nixon 1983; Ward & Nixon 1992). How the post oak savanna region relates 
biotically to other regions in the area has yet to be studied in detail (Monk et al. 
1990; Bryant et al. 1993; Skeen et. al. 1993); however, it appears to be f loristically 
similar to adjoining regions (MacRoberts & MacRoberts pers. obs.) and has no 
vertebrate endemism (Telfair 1999). 

McBryde (1933) conducted the only major flonstic/edaphic study of the 
region. Subsequently, very little research has been done on the post oak savanna 
except for the inclusion prairies in its southern portion (Smeins & Diamond 

Xeric sandylands are (or at least once were) very common in the Texas post 
oak savanna region, but they also occur in the pineywoods regions of south- 
eastern Oklahoma, southwestern Arkansas, western Louisiana, eastern Texas, 

and into the Coastal Bend of Texas (McBryde 1933; Jones 1977; Drawe et al. 1978; 
Taylor & Taylor 1978; Ajilvsgi 1979; Marks & Harcombe 1981; Louisiana Natu- 
ral Heritage 1988; Bridges & Orzell 1989; Harcombe et al, 1993; Foti et al. 1994; 
MacRoberts & MacRoberts 1994, 1995, 1996; Jones & Carpenter 1995; Texas Natu- 

ral Heritage Program 1995; Turner et al. 1999; Hoagland 2000). Homologues of 
xeric sandylands occur east of the Mississippi River (Stout & Marion 1993). It is 
generally assumed that oak savannas and xeric sandylands, like prairies, are 
kept open largely by fire (Smeins & Diamond 1986; Cutter & Guyette 1994). 
While encroachment of woody vegetation is ubiquitous in the absence of fire, 
there are many areas that remain open even when fire is suppressed, suggest- 
ing that edaphic conditions play an integral role. Characteristic tree species of 
xeric sandylands include Quercusincana W. Bartram, Q. margarettiae Ashe ex 
Small, Q. stellata Wangenh., and Carya texana Buckley, and a variety of fidel 
herbaceous species (see Methods for list). The area appears to be rich in plant 
endemism (Some & Weakley 2001), and a large number of West Gulf Coastal 
Plain endemics are associated with this community, for example, Brazoria 
truncata (Benth.) Engelm. & A. Gray, Palajoxia reverchonii (Bush) Cory, Parony- 
chia drummondii Torr. & A. Gray, Pediomelum hypogaeum (Nutt. ex Torr. & A. 
Gray) var. suhulatum (Bush) J. W Gnmes, Penstemon murrayanus Hook., 
Polanisia erosa (Nutt.) H.H. litis, Rhododon ciliatus (Benth.) Epling, 
Tetragonotheca ludoviciana (Torr. & A. Gray) A. Gray ex Hall, Tradcscantia 
reverchonii Bush, and T. suhacaulis Bush. 

In order to develop an objective idea of the distribution of xeric sandylands, we 
selected 42 fidel species from the total list of species occurring in this commu- 
nity. We mapped these by county and parish over their ranges (Fig. 2). In the 
map, the number of fidels per county or parish is indicated in the legend. Par- 
ishes or counties with fewer than 10 species are left blank. 

The 42 species chosen were Astragalus leptocarpus Torr. & A. Gray, A. 
soxmaniorum Lundell, Berlandiera pumila (Michx.) Nutt., Brazoria truncata. 
Clematis reticulata Walter, Cnidoscolus texanus (Muell.-Arg.) Small, Coreopsis 
intermedia Sherff, Crataegus uni/IoraMuenchh.,Crotonargyran(:hemusMichx., 
Cyperus grayioides Mohlenbrock, Dalea phleoides (Torr & A. Gray) Shinners, 
D. viHosa (Nutt.) Spreng., Eriogonum longifolium Nutt., E multiflorum Benth., 
Froelichia floridana (Nutt.) Moq. (not distinguished from F. gracilis t(Hook.) 
Moq.l), Hymenopappus artemisiifolius DC, Lithospermum caroliniense (Walter 
ex J.R Gmel.) MacMill, Loeflingia squarrosa Nutt., Matelea cynanchoides 
(Engelm.) Woodson, Minuartia drummondii (Shinners) McNeill, Palajoxia 
hookerianaTorr.&i A.Gidiy, P. reverchomi,Paronychia drummondii, Pediomelum 
digitatum (Nutt. ex Torr. & A. Gray) Isely, P. hypogaeum var. suhulatum, Penste- 
mon murrayanus, Phacelia strictijlora (Engelm. &r A. Gray) A. Gray, Phlox 
drummondii Hook., Physalxs mollis Nutt., Polanisia erosa (Nutt.) H.H. Ihis, 
Polygonellaamericana (Fisch. & C.A. Mey.) Small, PrunusgracihsEngelm. & A. 
Gray, Rhododon ciliatus, Selaginella arenicola Underwood ssp. riddellii (Van 
Eselt.) R. M. Tryon, Scutellaria cardiophylla Engelm. & A. Gray, Streptanthus 

hyacynthoides¥iook.,Stylisma pickeringii (Torr.ex M.A. Curtis) A. Gray, Talinum 
rugospermum Holz., Tetragonotheca ludoviciana, Tradescantia reverchonii, 
Yucca louisianensis Trel, and Zornia hracteatajF. Gmel. Sources for this infor- 
mation consisted of extensive lierbarium searches, notably LSU, TAMU (both 
on line), ASTC, BRIT, Corpus Chnsti Museum of Science and History LSUS, 

SBSC, SHST, TEX, VDB, WWF, atlases (e.g., Smith 1988; Thomas & Allen 1993- 
1998; Turner m press), and a variety of literature (e.g., Jones 1977; Taylor & Tay- 
lor 1978; Singhurst 1996; Turner 1996; Nesom & Brown 1998) and unpublished 
sources (Billie Turner pers. comm.; Bruce Hoagland pers. comm.). We also made 
field searches throughout the West Gulf Coastal Plain to look for the species 
and the community in situ. 

We surveyed the flora of xeric sandylands at Gus Engeling Wildlife Man- 
agement Area (GEWMA), Anderson County, Texas, in the center of the post 
oak savanna region. The GEWMA occurs in the central part of the Trinity River 
drainage along Catfish Creek (Telfair 1988). This property consists of approxi- 
mately 44 sq. km, of vv^hich 18 are the sand post oak-bluejack oak community 
(Singhurst et al. 2000). We established a single study plot measuring 50 m x 
100 m (0.5 ha), within which was another single plot measuring 20 m x 50 m 
(0.1 ha) with two nested 3.16 m x 3.16 m plots (0.001 ha) and 6 nested 1 m x 1 m 
plots (0.0001 ha) m an opening m this community (see Peet et al. 1998 for plot 
design). The larger plot ran parallel to the topographic gradient and the adja- 
cent downslope bog and was about 100 meters north of the edge of the bog and 
a few meters higher than the bog (see MacRoberts & MacRoberts 1998b, 1999; 
Singhurst et al. 2000 for a description of Andrew's Bog, GEWMA). We surveyed 
this area monthly from March to October 2000 and listed all plant species in 
each plot. This gave us information on both total floristics and species/area 
measures for comparison with similar measures from other plant communi- 
ties. We estimated ground cover and measured vegetation height in the 0.0001 
ha plots. We also surveyed several other xeric sandylands in other parts of the 
GEWMA. Plant nomenclature throughout this paper follows Kartesz and 
Meacham (1999). Voucher specimens are deposited at TEX. 

We collected soil samples from the upper 15 cm of the two 0.001 ha plots 
and from the 0.1 ha plot for comparison with this community elsewhere in its 
range. These were analyzed by A & L Laboratories, Memphis, Tennessee. The 
soil at the study site is described as deep, well-drained acidic sandy Pleistocene 
terraces (Arenosa series, Typic Quartzipsammeents, Entisols) (Coffee 1975). 

Figure 1 shows the location of the Post Oak Savanna region of East Texas in 
which xeric sandylands are located. Figure 2 shows the frequency of occurrence 
of the 42 xeric sandyland fidel species m parishes and counties in the West 
Gulf Coastal Plain. Counties and parishes with fewer than 10 species are left 
blank. Table 1 lists the species found within the 0.1 ha plot. Table 2 gives addi- 
tional species in the 0.5 ha plot. Table 3 shows the number of species occurring 
m the 0.0001 ha, 0.001 ha, 0.1 ha, and 0.5 ha plots. Table 4 gives the ground cover 
and vegetation height in the six 0.0001 ha plots m May and August. Table 5 gives 
the soil information from the study plots. 



dendron radicans (I.) Kunlze 

virginiana (L.) Benth., Chamaecrista fasciculota 

APIACEAE: Spermolepis divarlcata (Walter) Raf. 

(Michx.) Greene, Dalea phleoides (Torr. & A. 

ex Ser., S. mermis (Nutt. ex DC.) Mathias & 

Gray) Shinners, Indigofera miniata Ortega, 


Mimosa nuttallii (DC.) B.L.Turner, Pediomelum 

AQUIFOLIACEAE://e;< wm/for/o Sol. in Alton 

digitatum (Nutt. ex Torr. & A. Gray) Isely 

ASCLEPIADACEAE: Matelea cynanchoides 

FAGACEAE: Quercus incana W. Bartram, 0. 

(Engelm.) Woods 

margareff/oe Ashe ex Small 

ASTERACEAE: Croptilon divoricotum (Nutt.) Raf., 

HYDROPHYLLACEAE: Phacelia strictiflora 

Evax Candida (Torr. & A. Gray) A. Gray, 

(Engelm.& A.Gray) A.Gray 

Helianthus debilis Nutt, ssp. cucumerifolia 


(Torr. & A. Gray) Heiser, Krigia virginica (L.) 

LAMIACEAE: Brazoria truncata (Benth.) Engelm, 

Willd., Palafoxia reverchonii (Bush) Cory, 

& A. Gray, Monarda punctata L, Rhododon 

Pyrrhopappus carolinianus (Walter) DC, Sene- 

c/7/afus (Benth.) Epling 

cio ampullaceus Hook., Thelesperma fili folium 


(Hook.) A.Gray 

MOLLUGINACEAE:/Wo//ugo verticillata L, 

BRASSICACEAE: Lepidium virginicum L., 

ONAGRACEAE: Oenof/iera laciniata Hill 

Streptanthus hyacinthoides Hook. 


CACTACEAE; Opunf/a humifusa (Raf.) Raf 

CA, Mey. 

CAPPARACEAE: Po/on/s/o erosa (Nutt.) H, H. litis 

POACEAE: /\ndropogon ternanus MicUx., Anstida 

CARYOPHYLLACEAE: Loeflmgia squarrosa Nutt., 


Paronychia drummondii Jon.& A. Gray 

\iott, Dichanthelium acuminatum (Sw,) Gould 

aSJACEAt.Helianthemum georgianunn Chapm., 

Lechea mucronata Raf., L tenuifolia Michx. 

Paspalum setaceum M\chx.,Triplasis purpurea 

COMMELINACEAE: Commelina erecta L, Trades- 

(Walter) Chapm„l/u/p/ae///oteo (Raf.) Fernald. 

canr/a reverchonii BushJ. subacaulis Bush 

POLYGON ACEAE:fr/ogonum multiflorum Benth,, 

CONVOLVULACEAE: Stylisma pickeringii (Torr. ex 

ftumex^flstafu/us Baldwin, 

M.A.Curtis) A. Gray 

PORTULACACEAE: Talinum rugospermum Holz, 

CRASSULACEAE: Sedum nuttallianum Raf 


CUPRESSACEAEJun/perus virgmiana L 

RUB! ACEAE:aod;a feres Walter 

CYPERACEAE:Carexcep/io/op/ioraMuhl.ex Willd., 

SCROPHULARIACEAE: Nuttallanthus canadensis 

Cretroflexa MuhI.ex Willd., Cyperu5 grayioides 

(L,)D,A, Sutton 


50LANACEAE:P/7y5a//s heterophylla Nees 

EUPHORBIACEAE: Cnidoscolus texanus (Muell.- 

Arg.) Small Croton argyranthemus Michx., C 



Xeric sandylands occur from southwest Arkansas and southeastern Oklahoma 

to central Louisiana and the Coastal Bend of Texas. The distribution of species 
numbers shown in Figure 2 results in part from differential collecting: Caddo 
Parish and Anderson County have been well collected while most of east Texas 

Bulbostylis ciliatifolia (Elliott) Fernald Schizachyrium s 

Carya texana Buckley Scleria triglomef 

Corydalis micrantha (Engelm. ex A. Gray) A^ Gray Trichostema dici 




Vn,s rotund 


son Count 





ilife Management Area (Ander- 



Ncofplots species (range) 

has not. Better sampling would undoubtedly fill m the picture, but the outlines 
are clear We have observed the community in situ in Miller County, Arkansas; 
Bienville, Caddo, Natchitoches, Vernon, and Winn parishes, Louisiana; Atoka, 
Choctaw, and Pushmataha counties, Oklahoma; and Anderson, Angelina, 
Aransas, Bastrop, Caldwell, Cass, Cherokee, Colorado, Franklin, Gonzales, 
Guadulape, Hardin, Henderson,Jasper, Lee, Leon, Marion, Milam, Nacogdoches, 
Panola, Rusk, San Augustine, San Patricio, Shelby, Smith, Tyler, Upshur, Van 
Zandt, Wilson, and Wood counties, Texas. We have little experience with the 
Coastal Bend xeric sandylands (Drawe et al. 1978), but this community appears 
to have affinities with the more northern and eastern xeric sandylands and 
needs further study. Xeric sandlylands also occur in Hopkins, Navarro, Rains, 
and Williamson counties and appear to have once occurred on the boundary 
of Fannin and Grayson counties, Texas (Jason Smghurst, pers. comm.). 

There were 74 species in the 0.1 ha plot and 90 species in the 0.5 ha plot. 
Other taxa in xeric sandylands at GEWMA that did not occur m our plots in- 
clude Aphanostephusskirrhohasis(.DC)Tre\.,ApocynumcannabinumL.,Asdepias 
amplexicaulis Sm., Berlandiera pumila, Bouteloua hirsuta Lag., Delphinium 
carohnianumWalter,Descuraineflpinnata (Walter) Britton,Eragrostissecundi/7ora 
J. PresL, Eriogonum longijolium, Hymenopappus artemisiifolius, Liatris degans 
var. suhulatum, Physalis turhinata Medik., Polygonella americana, Scutellaria 
cardiophylla, Selaginella arenicola ssp. riddellii, Sideroxylon lanuginosum Michx., 


Tabif. 4. Ground cover and vegetation height in six 0.0001 \ 

2 (August) 



:ics of sample plots at the ( 

:uJsEngeling Wildlife Manage 




Exchangeable ions (ppi 

Ca Mg 

OrganK Matte* 

1 4.9 26 2/ 172 18 1.0 

Stillingia sylvatica Garden ex L, Tetragonotheca ludoviciana, Tragia urticifolia 
Michx., Iriodanis perjoliata (L.) Nieuwl, and Vicia ludoviciana Nutt (see also 
Smghurst et al. 2000). 

The ground cover varied from about 15 percent to 40 percent throughout 
the growing season. Non-vegetated areas always prominently showed and veg- 
etation was never tall. In general, biomass was low and sunlight was directly 
on the ground. 

Species richness can be measured at many scales. At scales of 0.01 ha and 
larger, tropical rainforests are the most species rich. However, at scales below 
0.001 ha and often 0.01 ha, temperate grasslands and open savannas of the south- 
eastern United States are the most species rich. Values of between 20 and 40 
species per 0.0001 ha occur but are very uncommon (Peet et al. 1983; Walker & 
Peet 1983; Peet & Allard 1993; Brewer 1998; Piatt 1999). Peet et al. (1983) found 
that for a broad range of forest and woodland types, no community type ex- 
ceeded 17 species per 0.0001 ha and none averaged over 13. Even tallgrass prai- 
ries, which were the highest, averaged only 18. 

1-scale informat 



apland pine sava 


s average 

et al. 1988; Ma 


3berts & 

Coastal Plani plant communties, the scant small 
indicates that wetland pine savannas, bogs, and u] 
around 20 species in 0.0001 ha plots (Allen . 
MacRoberts 1991, 1998a; Carr 2000). 

In the light of these figures, it is interesting that the GEWMA xeric 
sandyland plots show higher species richness at small scales than virtually all 
other plant communities so far measured in the southeastern United States. 
These xeric sandylands also show a low "z" value (about 0.19), meaning that 
there is basically a species doubling for every 100~fold increase in area. In the 
present case, a 0.0001 ha plot contains roughly one-half of the species found in 
a 0.01 ha plot, and one-quarter the species found in a 1.0 ha plot (see MacArthur 
& Wilson 1967; Harris 1984 for discussion of "z" value). 

The reason for such high species counts in these small-scale plots is not clear 
except that, in this case, most species are relatively diminutive with a variety of 
growth forms (about 60 percent of the species are perennials), and many have 
very brief above-ground life histories. Species packing is therefore no problem. 

The role of fire in maintaining plant communities is well understood (Piatt 
1999). In the absence of fire, many plant communities (e.g., prairies) are rapidly 
invaded by shrubs and trees (Packard & Mutel 1997). Oak-hickory savannas 
and xeric sandylands appear to require fire for natural maintenance (Cutter & 
Guyette 1994). We noticed in this study that, because of the long drought that 
the West Gulf Coastal Plain has been experiencing, there has been a significant 
die-off of woody vegetation, notably in the dryer areas. Fire, therefore, may not 
be the only important force preventing woody invasion. Periodic droughts may 
be another factor keeping xeric sandylands, as well as other communities, open. 

Soils at GEWMA are virtually identical to soils tested from xeric sandyland 
sites in Louisiana and east Texas (MacRoberts & MacRoberts 1994, 1995, 1996). 
They are nutrient-poor and acidic. Soil conditions per se may also preclude 

We have emphasized the f loristics of xeric sandylands largely to develop 
baseline information, and we note that the species richness of this community 
is considerably greater than generally recognized. The hydrologic and geomor- 
phologic properties of these xeric sandhills also are significant, as the water 
they supply underlies the existence of adjacent wetland communities— bogs, 
marshes, baygalls, and seeps. 

Bruce Hoagland, Connie Taylor, Robert Kalinsky Suzanne Walker, and the staff 
of the Gus Engeling Wildlife Management Area and all herbaria where we worked 
aided with this research. Special thanks to Billie Turner for allowing us to use his 
unpublished distribution maps for Texas plants. The work was supported by a 
Texas Parks and Wildlife Section 6, Wildlife Diversity Program Grant: "Status sur- 


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Eric L. Keith and N. Ross Carrie 

^itcher plant bogs require frequent fires to prevent encroachment of woody vegetation. Little : 
nation is available on the relative effectiveness of grovv'ing-season burns for maintaining plant 
nunities in bogs and reducing woody encroachment. In this study the effects of growing-seaso 
lormant-season burns on plant communities were assessed in two pitcher-plant bogs on the P( 
lidge Military Reservation in western l_ouisiana. Measurements were collected every month 

orb and woody plant density were determined in both bogs. Both bogs had been burned on 2 
otations in the dormant season for approximately 20 years. One bog, hereafter Conef lower boj 

)ther bog. Woodpecker bog, was burned during the growing season in August 1996 and is apj 

;pecies in Coneflower bog and 90 of the 98 species in Woodpecker bog. No species were elimii 
rom either bog after the fires. In addition, all species that were present after the growing seaso 

/o Ha), Texas pipe wort 
; yellow-eyed grass O 

r la invasion de vegetacion lefiosa. Esta disponible 

Pitcher plant bogs are unique ecosystems found in the southeastern United 
States. In western Louisiana, they occur in Beauregard, Natchitoches, Rapides, 
Sabine and Vernon parishes (Allen et al. 1988; Correll & Correll 1972.) Similar 
pitcher plant bogs can be found in adjacent counties in East Texas as well as 
areas m southeast Louisiana and eastward to Georgia and Florida (Folkerts 1982; 
Nixon & Ward 1986). Pitcher plant bogs typically contain saturated sandy soils 
underlain by an impervious sandstone or clay layer. Water that percolates 
through the sandy soil and reaches the impervious layer seeps to the surface. 
Mudstone from the Catahoula Formation usually forms the impermeable layer 
in bogs m the Peason Ridge area (Hart &r Lester 1993). 

Pitcher plant bogs are declining in quantity and quality throughout their 
range. Approximately 97% of pitcher plant bogs have been destroyed or severely 
altered since European settlement (Folkerts 1982). Public lands are virtually 
the only areas that this ecosystem can be managed effectively Pitcher plant 
bogs require frequent fires to prevent woody encroachment (Folkerts 1982). The 
absence of fire results in eventual elimination of bog species (Folkerts 1982). 
Until recently, the flora of pitcher plant bogs was relatively unknown and httle 
information is available on the relative effectiveness of fire for maintaining plant 
communities in bogs and reducing woody encroachment (Barker & Wilham- 
son 1988), Krai (1955), MacRoberts and MacRoberts (1988, 1990, 1991, 1992, 1993), 
and Nixon and Ward (1986) have recently conducted valuable research on the 
f loristics of bogs in this region, but information is scarce on the effects of grow- 
ing-season burns on pitcher plant bogs m the West Gulf Coast Plain. This project 
assesses the effects on vegetation of a growing season fire on a pitcher plant bog 
compared to a bog burned during the dormant season. We will also comment 
on several rare and interesting plants encountered during our study 

STUDY site/methods 
The study area was located on Peason Ridge of the Fort Polk Military Reserva- 
tion at 31" 20' 05"N 9 J 17' 00" W. The area surrounding the study sites is an open, 
sandy longleaf pine forest with gently rolling hills and is dominated by herba- 
ceous vegetation in the under story. The two bogs, hereafter Woodpecker bog 
and Coneflower bog, are located approximately 100 m apart and are separated 
by a narrow upland longleaf ridge. They are both open and relatively flat (2% 
slope) with a few scattered old growth longleaf pines and are composed mainly 


of herbaceous vegetation with a few scattered small trees, shrubs, and vines. A 
small creek runs along the eastern side of both of the bogs. Both bogs were vis- 
ited monthly from March through November 1996 and from March through 
October of 1997. The two bogs were chosen because of their close proximity, 
similar slope and aspect, and because no bog f loristic studies have been con- 
ducted on Peason Ridge. The nearest study conducted on bogs was 20 km to 
the northeast of the study site in the Kisatchie District of the Kisatchie Na- 
tional Forest (MacRoberts & MacRoberts 1988.) Woodpecker bog is approxi- 
mately 0.7 ha, and Conef lower bog is approximately 0.8 ha. Ten 1 x 1 m plots 
were spaced in a random pattern in each of the bogs (Fig. 1). Herbaceous and 
woody plant measurements were collected in each of the ten plots. Measure- 
ments included species frequency of occurrence and density. 

Five 5 X 5 m randomly selected quadrates were also used to measure woody 
stem density in Woodpecker bog (Fig. 1). Measurements of woody stems were 
collected in May of 1996 before the summer burn and again m May of 1998 after 
the burn to allow for a full growing season. Woody stems over 1 m tall were also 
counted to record the number of mature plants. The growing-season burn in 
Woodpecker bog was conducted on 20 August 1996 in the afternoon with south- 
erly winds and was effective in completely eliminating most herbaceous and 
leafy material. Conef lower bog was burned in December 1995. 

We follow MacRoberts (1989) and Allen (1992) for scientific nomenclature. 
Voucher specimens for some species were deposited at the University of Louisi- 
ana-Monroe and the Botanical Research Institute of Texas (BRIT). 

Burn Affects 

We recorded a total of 98 species in both bogs, 94 species in Coneflower bog 
and 90 species in Woodpecker bog. In Table 1, we list the species present in both 
bogs; "C" indicates species found only in Coneflower bog, "W" indicates species 
found only in Woodpecker bog, no letter indicates a species that was found in 
both bogs. White-topped sedge (Dichromena latifolia), Red milkweed (Asd- 
epias rubra), and Longleaf milkweed (Asclepias longijolia) were the only spe- 
cies found exclusively in Coneflower bog. Species more abundant in Coneflower 
bog were Bog-buttons (Lachnocaulon anceps), Drum heads (Polygala cruciata), 
and Sabine coneflower (Rudheckia scahrifolia). Texas pipewort (Eriocaulon 
texense), Narrow leaved-sunflower (Helianthus angustif alius), and Rough- 
leaved yellow-eyed grass (.Xyris scahrifolia), and red-top panicum (Panicum 
rigidulum var. combsii) were the only species found exclusively in Woodpecker 
bog. In addition, Rose pogonia (Pogonia ophioglossiodes) and Meadow beauty 
(Rhexia petiolata) were much more abundant in Woodpecker bog (Table 2). 

Several species numbers were significantly affected by each of the burn 
treatments (Table 2). Pitcher plant (Sarracenia alata) numbers were similar 

Woodpecker Bog 

n both bogs. 

before and after the grow 
dia were present in ttie te 

ings of Barker and William: 

urn. Before the fire, a total of 392 phyllo 

in the ten one meter plots. One month after the fire, 383 phyl 

;ed, and m the next growing season, 395 phyllodia were present 

:au-ied the number of individuals present, contrary to the find- 

They mdicated a 24% increase in the 

ber of phyllodia present in the growing season after a winter fire. A 3% increase 
in the number of phyllodia occurred in Woodpecker bog after this burn. How- 
ever, in Coneflower bog, 463 phyllodia were present in the ten plots immedi- 
ately after the fire and only 321 phyllodia were present the next growing sea- 
son representing a 31% decrease in the number of phyllodia present between 
subsequent growing seasons. This decrease in phyllodia appears to be due to 
increased competition from other vegetation that returned after the fire. 

Our study and previous studies suggest frequent fires are necessary to main- 
tain the density of certain species over tnne. Barker and Williamson (1988) 
indicated an annual decay rate of 36% in Sarracenia alata phyllodia in an un- 
burned bog in St, Tammany Parish, Louisiana. At this rate of decay pitcher plants 
would lose 95% of their foliage in only 6 years (Barker & Williamson 1988). Bog 
buttons {Eriocaulon decangulare) displayed similar results in our study We 
found 41 plants before the growing-season burn and 40 plants after the burn. 
However, in Coneflower bog, this species declined 59%, from 51 to 21, in just one 


Apiaceae— fryng/L/m integrifolium Walt.; 

(Chapm.) Lloyd & Underw.; Lycopodium 

Oxypolis rigidior (L.) Raf.; Ptiimnium costatum 

carolinianum L. 

(Ell.) Raf. 

Osmundaceae— Osmunda cinnamomea L 

Aquifoliaceace— //ex cor/ocea (Pursh.) Chapm. 

Pmaceae^Pinuspalustris P.Mill.;P/nu5 taeda L. 

Asclepiaceae— ^sc/ep/as longifolia Michx.,(C).; 

Asclepias rubra L, (C). 

Burmanniaceae— eurmann/o capitota (Walt.) 

Asteraceae— /\sfer dumosus L.; Cacalia ovata 


Walt.; Chaptalia tomentosa Vent.; Coreops/s 


//n;fo//o Nutt.; Eupatorium leucotepsis (DC.) 

latifolia Baldw. ex. Ell., (C); Eleocharis 

Torrey & Gray; Eupatoriurm rotundifolium L.; 

tuberculosa (Michx.) Roem. & Schult.; Fuirena 

Helianthus angustifolius L, (W); Heterotheco 

Caprifoliaceae— Wbumum / 

(Michx.) Adams & Robson. 

Droseraceae— Drosera capillaris Poi 

Eucaceae—Vaccinium corymbosa L 

Orchidaceae— Ca/pogon tuberosus (L.) B.S.P.; 

Euphorbiaceae— Crotonops/s e///pr/ca Willd. 

Pogonia ophioglossoides (L.) Juss.;Spfranf/-ies 

Gentianaceae — Bartonia paniculata (Michx.) 

cernua (L.) L.C. Rich. 

Miihl -SabatiagentianoidesBl 

Lamiaceae — Scutellaria integrifolia L. 

enantia rufa (Ell.) Schult.;/^nr^oenonf/o viltosa 

Lentibulariaceae— P/ngu/cu/o pumila Michx.; 

(Michx.) Beauv.; /\r/sf/da pa/u5fr/s (Chapm.) 

Utricularia cornuta Michx.; Utricularia 

Vasey; Eragrostis refracta (Muhl.) Scribrn.; 

subulata L. 

Loganiaceae— Cynocfonum sessilifolium (Walt.) 


Ell.; Panicum rigidulum Nees. var. corvbsii 

Magnoliaceae— /Mogno//o virginiana L. 

(Scribn. & Ball), (W); Panicum scabriusculum 

Melastomataceae— /?/iex/o /uteo Walt.; /?/iex/o 

Ell.; Pan/cum tenerum Beyr.; Pon/cum 

mariana L.; /?hex/a pef/o/afo Walt. 

verrucosum Muhl.; Panicum virgatum L.; 

Myricaceae— /Myr/co cerifera L.; Myr/ca 

^'coparrmTM^ hxT N^ash; ^ 

NyssacIafCys^sasy/i^af/ca Marsh 

fenerum Nees; Tridens ambiguus (Ell.) 

Onagraceae— ^uc/iY/g/o hirtella Raf. 


Polygalaceae— Po/yga/o cruc/afa L.; Polygala 

Xyridaceae— /yr/sam6/guoBey.ex.Kunth.;Xyns 

ramosa Ell, 

Pogonia ophioglossiodes 
Polygala cruciata 
Polygala ramosa 

growing season Pitcher plants and bog buttons appear to require frequent fires 
to maintain their presence. 

Other species that also increased immediately after fire and declined after 
one growing season were Colic root (Aletris aurea), Cynoctonum sessifolium, 
milkworts (Polygala cruciata and Polygala ramosa), Rose pogonia (Pogonia 
ophioglossiodes), Meadow beauty (Rhexia petiolata), and Primrose-leave violet 
(Viola primulijolia) (Table 2). Unlike these species, Club moss (Lycopodium 

number of stems continued to increase, from 26 to 179, a year after the dor- 
mant-season burn in Coneflower bog. Perhaps this moss-like prostrate species 
requires more than one growing season to maximize the number of reproduc- 
tive shoots that it produces. 

The total number of woody stems increased after the growing-season burn, 
however, woody stems greater than one meter tall decreased. (See Table 3 for 
woody stem counts before and after the growing-season fire.) This increase in 
stems after the fire can be attributed to the large number of root sprouts from 
the base of the small hardwood trees, shrubs, and vines. Several small stems 
replaced one large stem that had been top killed by the fire. However, many 
shrubs were completely killed by the fire with no root sprouting. Sweet bay 


(Magnolia virginiana) and Loblolly pine (Pinus taeda) were the only species 
that decreased after the fire. In addition, Poison sumac (Rhus vcrnix) and Blue- 
berry (Vaccinium corymhosum) were not present in the plots before the fire, but 
six plants of each species were found in the plots after the fire. 
Species Encountered 

Rudheckia scahrifolia was common in the wetter areas of both bogs. This site is 
the northernmost record for this species. MacRoberts and MacRoberts (1993) 
did not record this species in Natchitoches Parish and did not find it in their 
study sites in the Kisatchie National Forest (MacRoberts & MacRoberts 1988, 
1990, 1991, 1992). However, our study site is only 2.5 kilometers south of the 
Natchitoches Parish line suggesting Rudheckia scahrifolia could be found in 
this parish and elsewhere on Peason Ridge. 

Eriocaulon texense was frequent in Woodpecker bog, but not present in 
Conef lower bog. This plant was the only species that was found commonly in 
one bog and was absent in the other. This occurrence could also be the north- 
ernmost location for this species. MacRoberts and MacRoberts (1993) noted that 
they did not find this species m the Kisatchie District of the Kisatchie National 
Forest. The location of this species is approximately 20 kilometers north of any 
other known location. 

Panicum rigidulum var. comhsii was very uncommon in Woodpecker bog 
with only two small clumps found, and it was completely absent from Cone- 
flower bog. This occurrence is only the second record for this distinctive vari- 
ety m Louisiana (Allen 1992). 

Panicum tenerum was common in both bogs. This plant was not found on 
Peason Ridge in a f loristic survey (Hart & Lester 1993) and is rare in bogs and 
flatwoods m the southern part of the state (Allen 1992). 

Rhynchospora chalarocephala was abundant in both bogs. MacRoberts and 
MacRoberts (1993) found this species in two bogs in the Kisatchie District and 
one bog in the Vernon District. They suggested that this species could be more 
common than previously believed. We have seen this species quite commonly 
in many bogs on Fort Polk and on Reason Ridge. 

Xyris drummondii was common in wet, bare ground in both bogs as well 
as many other bogs on Reason Ridge and Fort Polk. This species seems to be 
more common than previously documented. 

Xyris scahrifolia increased drastically after the growing season burn. We 
found 1 stem in our plots and three stems outside the plots before the burn. Af- 
ter the burn, we found 9 stems in our plots and 50 stems outside the plots. 
Godfrey and Wooten (1979) suggest that this is the rarest entity of Xyris in the 
southeastern United States, known only from Florida and Georgia. It is pos- 
sible that this species could be more common than previously thought. In ad- 
dition, it has been found recently in several locations on Fort Polk (Hart & Lester 

The goal of this study was to collect observational data before and after 
prescribed fires in two pitcher plant bogs. Although it would have been opti- 
mal to collect data in many bogs over several years burned in different growing 
seasons, logistical problems and other duties prevented such a comprehensive 
study Based on data collected through observations over two growing seasons, 
we conclude numerous bog species respond drastically to fire. The life history 
and habits of those species suggest frequent burns may be necessary to main- 
tain the unique bog ecosystem. We predict future fire ecology studies will fur- 
ther support the necessary role of fire in the bog ecosystem. 

This research was supported in part by appointments to the postgraduate en- 
vironmental management participation program administered by the Oak 
Ridge Institute for Science and Education, We would like to thank Charles Stagg, 
Stephen Parris, Stephanie Stephens, Kenneth Moore, Chris Melder, and Kevin 
Mundorff of the Fort Polk Environmental and Natural Resources Division and 
Michael and Barbara MacRoberts of Bog Research for all their help and advice 
on this project. We would also like to thank Charles Allen at the University of 
Louisiana-Monroe, Larry Brown at Spring Branch Science Center in Spring 
Branch, Texas and Guy Nesom at the Botanical Research Institute of Texas for 
their assistance m identifying specimens; and William Keith of Lufkin, Texas 
for Spanish translation of the abstract. 

:.M.,C.H.STAGG,and S.D.Parris. 1 988. Analysis of the vegetation in pitcher plant bogs 
/vo baygalls at Fort Polk in west central Louisiana. Proc. Louisiana Acad. Sci. 50:1-6, 
N.G. and G.B. Wiluamson. 1988. Effects of a winter fire on Sarracenia data and S. 

_,D.S.and H.B.Correll. 1972. Aquatic and wetland plants of the southwestern United 
es. Environmental Protection Agency, Washington, D.C. 
s,G.W. 1982.The Gulf Coast pitcher plant bogs. Amer. Sci. 70:260-267. 
Y, R.and J.W. Wooten. 1 979. Aquatic and wetland plants of the southeastern United 
es. Volume I, Monocotyledons. University of Georgia Press, Athens. 
..L. and G.D, Lester. 1 993. Natural community and sensitive species assessment on 
ouisiana Dept. of Wildlife and Fisheries, Ba- 



ilitary Reservation, 


1 Rouge, 

LA in cooperation ^ 

11955. A 


plant bogs.Phytologia 68:271-275. 
MacRoberts, B.R. and M.H. MacRoberis. 1 
Phytologia 70:135-141. 

ana with observations on species/area relationships. Phytologia 73:49-56. 

.cRoberts, B.R. and M.H. MacRoberts. 1 993. Floristics of a bog in Vernon Parish, Louisiana, 

iA/ith comments on noteworthy bog plants in western Louisiana. Phytologia 75:247- 

Bull. Mus. Life Sciences, Nos. 7-9, Louisiana State University-Shreveport. 
:oN,E.5.and J.R.Ward. 1 986.Floristic composition and management of east Texas pitcher 
plant bogs. In Wilderness and Natural Areas in the Eastern United States: A Manage- 
ment Challenge, eds.D.LKulhavy and R.W.Conner. Center for Applied Studies, School 
of Forestry, Stephen F. Austin State University, Nacogdoches, TX. Pp. 283-287. 

Books Received/Notices 

Texas Floras/Natural HistoryAVildlife 

AND H. Wauer and Carl M. Fleming. 2002. Naturalist's Big Bend: An Intro- 
duction to the Trees and Shrubs, Wildflowers, Cacti, Mammals, Birds, Reptiles 
and Amphibians, Fish, and Insects. (ISBN 0-58544-156-2, pbk.; 0-58544- 
155-4, hbk.). Texas A&M University Press,John H. Lmdsey bldg, Lewis Street, 
4354 TAMU, College Station, TX 77843-4354, U.S.A. (Orders: 800-826-8911, 
979-847-8752 lax,, $29.95 
(hbk), $15.95 (pbk), 208 pp., 15 color photos, 28 b/w photos, 90 hne draw- 

DavidJ Slhmidiy (forewords by Andrew Sansom and Robrt J Potts, afterword by 
Clyde Jones). 2002. Texas Natural History: A Century of Change. (ISBN 0- 
89672-469-7, hbk.). Texas Tech Press, Box 41037, Lubbock, TX 79409-1037, 
U.S.A. (Orders: 800-832-4042,, $39.95 (hbk), 
576 pp., 144 b/w photos, 43 maps, 6" x 9". 


Tom S. CooPERRiDER, Allison W. Cusick, and John T Kartesz (eds). 2002. Seventh 
Catalog of the Vascular Plants of Ohio. (ISBN 0-8142-0858-4, hbk.; 0-8142- 
5061-0, pbk.). The Ohio State University Press, Columbus, OH U.S.A., (Orders: University of Chicago Distribution Center, 
11030 S Landley Ave., Chicago, IL 60628, U.S.A.; 800-621-2736). $29.95 (pbk), 
195 pp., 8" X 9 1/2". 



Robert F.CNaczi 

Claude E.Phillips Herbarium 
ent of Agriculture and Natural Res 


Ronald L.Jones, 

F.Joseph Metzmeier, 

Mark A. Gorton, 

and Timothy J. Weckman 

Department of Biological Sciences 



ar plant., all 

the fir 

nana, C. oxylepis var. puhesccns, C. wanen: 
rex huxbaumii and Cseona. 









Recent field and herbarium work have resuhed in the discovery of several spe- 
cies of native flowering plants new to Kentucky or very rare in the state. These 
findings will update Kentucky distribution records as listed by Beal and Thieret 
(f 986), Browne and Athey (1992), and Medley (1993). Kentucky rarity status- 
Special Concern, Threatened, Endangered, or Historical, is based on KSNPC 
(2000, 2001). Additional information on Clinton County records is provided 
by Gorton (2000). Terminology for physiographic regions is based on Browne 
and Athey (1992), and herbarium abbreviations follow Holmgren et al. (1990). 


Bromus nottowayanus Fernald (POACEAE). Though Kentucky is well within 
the geographic range of this species as provided by Pavlick (1995), this record is 
apparently the first published for the state. Almost certainly, earlier Kentucky 
collections exist for this woodland grass. 

Carex buxbaumii Wahlenb. (CYPERACEAE). This is the second county in Ken- 
tucky documented for this species. Earher records for this species are from 
Baliard County, in the westernmost portion of Kentucky, in the Coastal Plain 
region (Medley 1993). These records are over 20 years old, and the species is cur- 
rently listed in Kentucky as Historical. This collection therefore provides a 
redocumentation of the species for the state, and is also the first record from the 
Mississippian Plateau region of the state. Kentucky is near the southern limit 
ol this species; east of the Mississippi River, it ranges south to North Carohna 
and Ten nessee (Radford et al. 1968; Gleason & Cronquist 1991; Chester et al. 1993). 

Carex corrugata Fernald (CYPERACEAE). These specimens represent the first 
record of the species from the state. Kentucky is the northern limit of this sedge, 
which is most common on river f loodplains in the Coastal Plain of the south- 
eastern U.S.A. (Naczi 1992). Most of the collections cited here are from sites along 
the Ohio or Mississippi rivers. 

Carex oklahomensis Mack (CYPERACEAE) These specimens represent the first 
1 Lcord of the species hom the state This species also occurs in the midwestern 
USA fiom Missouri and Kansas south to Texas and m the eastern U.S.A. in 
Mississippi North Citohnr Ttnncssct md Virginia (Jones & Reznicek 1995; 
Brysonetal 1996 Wicboldtet il 1998) 

\ouLhti bpLUimns Calloway Co uu I Uloxv liJd SmpL Cieek Embiyment, 5 Jun 1989J.Grubbs 
hOnUUR) McCrackenCo \va duchts on tlu Ithiuidimn Ro^d 24 May 1970. R. Athe)/ 932 (KNK, 
MURJ Marshall Co wet woods Shiip Eh i Roid oli US b8 10 Mi> 1972 R Athey 1666 iMVR). 

Carex ouachitana Krai, Manhart & Bryson (CYPERACEAE). This recently de- 
scribed sedge is known from sites in western Arkansas and eastern Oklahoma 
(Krai et al. 1987; Naczi & Bryson 1990), and a single site in Clay County, Tennes- 
see (McNeilus 1992). This collection from Clinton County, Kentucky, only a few 
km from the Clay County site, represents a new record for Kentucky, and the 
second record of the species from east of the Mississippi River 

Voucher specimen: Clinton Co.: dry-mesic slope N of Albany, 11 Apr 1998, M. Gorton 98-J47(KKY); 21 
Apr 1999, M. Gorton 99-109 (EKY). 

Carex oxylepis Torr. & Hook. var. pubescens J.K.Underwood (CYPERACEAE). 
Though C oxylepis var. oxylepis is known from Kentucky (Medley 1993), this 
collection appears to be the first of C. oxylepis var. pubescens from the state. 
Additionally, the population documented by this record represents a portion 
of the northern range limit of this variety This taxon appears to be rare; litera- 
ture reports are from only one county in Alabama (Bryson et al. 1992), ten coun- 
ties of Arkansas (Hyatt 1998), one county of Illinois (Basinger & Shimp 1999), 
one county of Mississippi (Bryson et al. 1992), and four counties of Tennessee 
(Chester etal. 1993). 

Voucher specimen: Clinton Co.: mesic deciduous-juniper forest NE of Albany, 20 May 1999, R. Naczi 
7982 (DOV, EKY, KNK). 

Carex roanensis EJ.Herm. (CYPERACEAE). This is the first record of this sedge 
from Kentucky. It was previously known from only a few populations in Geor- 
gia, North Carolina, Tennessee, Virginia, and West Virginia, mostly from high 
elevation sites (Wieboldt et. al 1998). It was found at several sites on Black Moun- 
tain, at elevations between 1100-1150 m, but these sites are threatened by strip 
mmmg activities. It has recently been added to the Kentucky list of rare plants 
as Endangered. 

1998, R.Jones and]. Metzmeier 8737k 8757 (EKY). 

Carex seorsa Howe (CYPERACEAE). These records add two counties to the two 
previously reported, Bath and Graves (Ousley & Risk 1998; McKinney et al. 
2000). This species is most common on the Coastal Plain, and populations out- 
side of that physiographic province are very few (Reznicek & Ball 1980). It is 
currently listed as a Special Concern taxon in Kentucky. 

Eleocharis tuberculosa (Michx.) Roemer & Schultes (CYPERACEAE). This col- 
lection represents a state record for Kentucky It ranges mostly on the Coastal 
Plain, and it is quite rare outside of that province (Svenson 1937), though it does 
occur nearby in Fentress County Tennessee (Chester et al. 1993). 

i Griseb. (GENTIANACEAE). This collection is apparently 
the first documented record of this species for Kentucky. Browne and Athey 
(1992) cited specimens from the Mississippian Plateau and the Bluegrass re- 
gions, but these citations were apparently based on a specimen from Mammoth 
Cave (J. Bruna s. n., 23 Oct 1965, EKY!), which has now been annotated by R.Jones 

as Gentiana saponaria L, and on a specimen once housed at the University ol 
Kentucky Agricukural Herbarium— this specimen was searched for but could 
not be located. The collection cited by Medley (1993) as a voucher for Kentucky, 
(Short s.n., 1837, MICH!) lacks locality data. The "Lexington, Kentucky" pre- 
prmted on the label was Short's residence at the tune (Stafleu & Cowan 1985), 
and not the place where the plant was collected. Short's habit was to write the 
collection locality on the label, in the space above pre-printed information. Since 
his label lacks such an inscription, it must be regarded as of unknown prov- 
enance. Pringle (1967) cited no Kentucky specimens of G. andrewsii, and states 
that this Campbell County collection is the first he has seen for the state (J. Prin- 
gle, pers. comm.). The site of the collection is at or near the southern range limit 
for G. andrewsii, which is known from nearby southern Indiana and southern 
Ohio (Pringle 1967). At the Campbell County locality, the gentians were locally 
common in an open, wet-mesic, deciduous floodplain forest dominated by 
young Acer rubrum and Quercus palustris. 

Voucher specimen: Campbell Co.: Silver Grove, floodplain of Ohio River, 13 Oct 1997, R. Naczi 7059 
(KNK), dupl. det. by James S. Pringle; 29 Sep 1998, R. Naczi 7735 (KNK). 

Vaccinium erythrocarpum Michx. (ERICACEAE). The mountain cranberry is 
restricted to the southern Appalachians, from West Virginia to Georgia, mostly 
above 1000 m. Medley (1993) reported a possible sterile specimen of this spe- 
cies collected from Harlan County (L. Pounds 85-138) deposited at the Univer- 
sity of Tennessee Herbarium, but a recent search failed to locate any Kentucky 
collection (B. E. Wofford, pers. comm.). This fruiting specimen substantiates the 
presence of the species in Kentucky. The site, however, is likely to be destroyed 
by strip mining activities. It has recently been added to our state list of rare 
plants as Endangered. 

Voucher specimen: Harlan Co.: S slope of Black Moui 
Left Fork of Fugitte Creek, elevation 900 m, 14 Aug : 

Viburnum lantanoides Michx. (CAPRIFOLIACEAE). This is the first record of 
Viburnum lantanoides for Kentucky Earlier reports were unsubstantiated or 
based on misidentifications (Weckman 1999). The species is common m some 
high elevation areas of the southern Appalachians, usually above 1000 m. It 
has been documented from several western Virginia counties, including con- 
tiguous Vv^ise County (HarviU et al. 1986), but is not yet known from Tennessee 
counties adjacent to the Kentucky border (Chester et al. 1997). A single popula- 
tion of about 50 individuals was discovered in the Razor Fork stream head, grow- 
ing with Betula alkghaniensis and Magnolia fraseri. Persistence of this Ken- 
tucky population is threatened by potential valley fill associated with proposed 
strip mining on Black Mountain. It has recently been added to the Kentucky 
list of rare plants as Endangered. 

o.: Black Mountain, at head of Razor Fork, elevs 
:r8804 (EKY); 5 Sep 1998, 1 Weckman 4349 (EKY 

:AL. E.O. and J.W.Thieret. 1 986. Aquatic and wetla 

ind plants 

of Kentucky. Kentucky NatuP 

Preserves Connmission, Scientific and Technic 


dumber 5, Frankfort. 

«)WNE, E.T.Jr. and R. Athey. 1992. Vascular plan- 

ts of Kent 

ucky: an annotated checklis- 

University Press of Kentucky, Lexington. 

7son, C.T., J.R. Macdonaid, R. Carier, and S.D. J 

ones. 199( 

X Noteworthy Carex, Cyperui 

Eleocharis, Kyllinga, and Oxycaryum (Cyperac 

eae) fron- 

1 Alabama, Arkansas, Georgic 

Louisiana, Mississippi, North Carolina,Tenness 


■xas.Sida 17:501-518. 

^YSON, C.T., R.F.C. Naczi, and S. McDaniel. 1992. r 

^otes on 

noteworthy records of Care 

(Cyperaceae) from the southeastern United States. Sida 1 5:1 25-1 35. 

Chester, EW., B.E. Wofford, R. Kral, H.R. DfSei m, and A.M. Evans. 1 993. Atlas ofTennessee vascu- 
lar plants.Vol. 1 .Misc. Publ. No. 9.The Center for Field Biology Austin Peay State Univer- 
sity, Clarksville,Tennessee. 

Chester, E.W., B.E. Wofford, and R. Kral. 1 997. Atlas ofTennessee vascular plants.Vol. 2. Misc. 
Publ. No. 13. The Center for Field Biology, Austin Peay State University, Clarksville,Ten- 

Glfason, H.A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada, 2"^^ ed.The New York Botanical Garden, Bronx. 

Gorton, M. 2000. The vascular flora of Clinton County, Kentucky. M.S. thesis, Eastern Ken- 
tucky University, Richmond. 

Harvill A.M., Jr.,TR. Bradley, C.E.STEvtNS,TRWieboldt,D.M.E. Ware, and D.W.Ogle. 1986. Atlas 
of the Virginia flora. 2'^^ ed., Virginia Botanical Associates, Farmville. 

herbaria of the world, 8th ed. New York Botanical Garden, Bronx. 

Hyatt, RE. 1 998. Arkansas Carex (Cyperaceae): a briefly annotated list. Sida 1 8:535-554. 

Jones, S.D. and A.A. Reznicek. 1 995. Carex conjucta (Cyperaceae) verified for Arkansas, and 
notes on the range of Carex oklahomensls. Sida 1 6:772-774. 

Kral, R., J.R. Manhart, and C.T Bryson. 1987. A new Carex sect. Oligocarpae (Cyperaceae) 
from western Arkansas and eastern Oklahoma. Ann. Missouri Bot. Gard 74:440-442 

[KSNPC] Kentucky State Nature Preserves Commission. 2000. Rare and extirpated biota of Ken- 
tucky. J. Kentucky Acad. Sci. 61 :1 1 5-1 32. 

[KSNPC] Kentucky State Nature Preserves Commission. 2001 . Rare and extirpated biota of Ken- 

McNeilus, V.B. 1 992. Carex ouachitana (Cyperaceae) new to Tennessee. Sida 1 5:1 54-1 55. 
McKiNNEY,L.E.,G.LiBBY,and R.MEARS.2000.New and noteworthy records of Corex (Cyperaceae) 

from Kentucky. Castanea 65:221-224. 
Medley, M. 1 993. An annotated catalog of the known or reported vascular flora of Ken- 

University of Michigan, Ann Arbo 

Naczi, R.F.C, and CT.Bryson. 1990, Nc 

teworthy records of Core 

1 the 

southeastern United States. Barto 

iia 56:49-58. 

OusLEYj.R.and A.C. Risk. 1 998. Prelimin 


r flora oft 



Seeps, Bath County, Kentucky. J. K€ 

ntucky Acad. Sci. 59:97. 

Pavuck, L.E.I 995. firomusL. of North /^ 

merica. Royal British Cour 

nbia Muse 



Pringle, J.S. 1 967.Taxonomy of Gentian 

3, section Pneumonanthae 

n eastern 



Brittonia 19:1-32. 
Radford, A.E., H.E. Ahi fs, and C.R. Bell. 1 968. Manual of the vascular flora of the ' 

The University of North Carolina, Chapel Hill. 
REZNtCEK,A.A.andP.W. Ball. 1980.The taxonomy ofCarexsectionSre//u/otoe in Nort^ 

north of Mexico. Contr. Univ. Michigan Herb. 14:153-203. 
Stafleu, F.A. and R.S. Cowan. 1985.Taxonomic literature, 2"^' ed., Volume V: Sal-' 

SvENSON.H.K. 1937. Monographic studies in the genus Eleochans. IV.Rhodora 39 

(Caprifoliaceae) in Kentucky. M.S. thesis. Eastern Kentucky University, Richmc 


Pluchea yucatanensis Nesom. When first described (Nesom 1989), P. 
yucatanensis was known in the USA from only one collection in Hancock Co., 
Mississippi. It is now recorded from four contiguous counties in Alabama (first 
report here) and Mississippi and appears to be an established element of the 
flora in that area of the Gulf Coast. Its larger range is in Central America (Belize) 
and Mexico (Veracruz, Campeche, and Quintana Roo). 

VMA. Mobile Co.: just N of jet. Co. Rd 59 and Ala. Hwy 188, sandy edge of salt 

Jordan River, edge of mixed woods with Serenoa, Sabal Nyssa, Taxodium, 30 May 1967, Jones 12656 
(TEX); S of Kiln, low sandy soil along Jordan River, 6 Jul 1967 Jones 24255 (NCU). Harrison Co.: SW of 
Wool Market, low sandy soil along Biloxi River, R lOW T 7S Sec 8, 23 Jul 1967, Jones 14818 (GH). Jackson 
Co.: Ocean Springs RO., low wet areas, 2 Jul 1952, Demaree 32262 (BRIT-SMU, GH); E of Ocean Springs, near 
borrow pit on Hwy 90. 25 Jul 1966, Deramus 548 (VDB); Ocean Springs, 30 Jul 1896, Pollard 1129 (GH). 

Pluchea yucatanensis is similar in habit and general appearance to P.foetida 
(L.) DC. and P. rosea Godfrey and has been misidentified as both. The rose-tinted 
phyllaries and florets are more similar to P rosea, but the glabrous, slightly thick- 
ened, shiny leaves and glabrous phyllaries are unambiguous recognition char- 
acters for P yucatanensis. 

Pseudognaphalium luteoalbum (L.) Hilliard & Burtt. A recent report expanded 
the known range of this species into Texas and New Mexico (Nesom 2001). It 
probably should be expected in Oklahoma and other states eastward. It is al- 
ready documented in Florida by many collections. 

Symphyotrichum puniceum (L.) A. & D. Love var scabricaule (Shinners) Nesom. 
Variety scahricau le previously has been recorded from localities in Texas, Loui- 
siana, and Mississippi (Nesom 1997). 

The occunence ot Symphyotrichum punitcum m Alabama was documented 
by Mohr (1901), who noted that the only collection was by FS Fade from Au- 
burn m Lee County m the east^cential section of the state (immediately adja- 
cent to Chambers Counts ) Mohi noted that the plants giew m swampy bor- 
ders ol woods" in"metainoiphic hiUs ol the mountain iLgion Both Alabama 
locahties are at the southwestern extieme ot the southwaid Appalachian ex- 
tension of the range of 5 pumceum through Geoigia Some ol the distinctly 
montane Geoigia populations are typical m moiphology(vai pu?i iccum) while 
others (eg Haialson Co KnU 77I82-VDB) show features of var scahricauk, 

scahnauk \\ isLonsidcicdaCuH coastal plain endemic (Nesom 1997). Cham- 
bers and Lee coiintiLsan shghtU north of the coastal plain (as maiked by the 
boundaiy between C ict leeous nid older Paleozoic foimations, eg, see Sorrie& 
Weakley 2001) Lee Count), howevei, is included m foui of the geogiaphic pat- 
terns of coastal plain endemism shown Soi i le and Weakley 

Kaitesz (1999) includes Symphyotnchiim punucum for Alabama as var. 
punitcum, but the moi phole\gy of the plants suggests that identification as var 
scabncau/e is more apptopnatc Thcleowei leal suilaeesaie about the same color 
as the upper surfaces without a tetieulum ol daik\eins the main veins of the 
upper leaf surfaces are shallow l\ mipiessed gi\ iiig a slightly rugose appear- 
ance, and the leaves of the capitulescenee aie maikedly leduced m size com- 
pared to the lower. 

I am gratefu 

.ARTESz, J.I. 1999. A synonymized checklist and atlas with biological attributes tor the 
vascular flora of the United States, Canada, and Greenland (ed. 1 ). In: Kartesz, J.T., and 
C.A. Meacham. Synthesis of the North American flora, Version 1 .0. North Carolina Bo- 
tanical Garden, Chapel Hill. 

Pluchea (Compositae: Inuleae). Phytologia 67:1 58-167. 

Gulf coastal plain. Phytologia 82:300-315. 
Jesom, G.L. 2001. New records in Pseudognaphalium (Gnaphalieae: Asteraceae) for the 

United States. Sida 1 9:11 85-1 1 90. 
ORRiE,B.A.and A.S.WEAKLEY.200LCoastal plain vascular plant endemics:Phytogeographic 

patterns. Castanea 66:50-82. 


Allison W.Cusick 

Ohio Department of Natural Resources 

vasion of indigenous floras by non-natn 
It early appearances of alien species. This 

.e element 
paper foci 

species were first noted in 1976 and 1981, the remainder in 2000. None of these 
species are included in the checkhst of Iowa vascular plants by Eilers and Roosa 
(1994). Single populations of two species were first reported in Norris et al. 
(2001). This paper documents additional populations. Although these species 
are not widespread as yet in the state, there probably is no barrier to their spread. 
Most are weeds of ornamental plantings and are spread primarily via horticul- 
tural materials. Most previous studies of horticultural introductions have fo- 
cused on the spread of woody landscape plants (Vincent & Cusick 1998). The 
present work concerns the incidental introduction of herbaceous species. One 
shrub discussed below, Tamarix parviflora DC, potentially is a serious pest spe- 
cies, however. Careful searching should yield additional Iowa populations of 
all six species. 

Specimen citations are abbreviated and full details are available from the 

Lnethum graveolens L. (APIACEAE). Culinary dill is a sporadic waif in North 
»imerica. Apparently, this is the first report of this herb in the Iowa flora. 

les Moines Co.: bank of Mississippi River, US Hwy. 34 bridge, Burlington, 15 Jul 2000, Cuiich 35548 

■atoua villosa (Thunb.) Nakai (MORACEAE). Mulberry-weed is an annual spe- 
ies from Asia which now is common throughout the southeastern United 
tates. It typically grows in nurseries and amid ornamental plantings. It also is 
common adventive in greenhouses. The Iowa population is a northwestern 
ange extension from southern Missouri and central Ohio (Wunderlin 1997). In 
he north, Fatoua villosa is restricted to protected situations, such as walls and 
outh-facing slopes. The climate at the Muscatine County site is moderated by 
he nearby Mississippi River Mulberry-weed likely will be found at other places 

Muscatine Co.: nursery on terrace of Mississippi River, St, Hwy. 22, Muscatine, 21 Jul 2000, Cusick 
35602 (ISC, MICH, MU. OSH). 

Oxalis corniculata L. (OXALIDACEAE). This species is adventive from tropical 
America. In the southern U.S., it is widespread in ornamental plantings. It is 
common in greenhouses as well (Lourteig 1979). As with Fatoua villosa, it ap- 
parently spreads as a contaminant with potted ornamentals, based upon the 
author's personal observations of nursery material. 

2000. Cusicle 35603 (ISC). 

Sagina procumbens L. (CARYOPHYLLACEAE). European pearlwort was first 
reported in Iowa from Story County where it was discovered in 2000 (Norris et 
al. 2001). It is common throughout the northeastern and northwestern U.S., but 
sporadic in Midwestern states. This perennial species typically grows in crev- 
ices in walls and sidewalks and as a weed in ornamental plantings. The similar 
annual, Sagina decumhens (Ell.) T&G., grows m comparable situations and oc- 
curs near Iowa m northcentral lUinois (Crow 1978). 

Cerro Gordo Co.: garden store, 4th St. SW & S Polk Ave., Mason City, 19 Jul 2000, Cusick 35588 (ISC); 
Muscatine Co.: nursery on terrace of Mississippi River, St. Hwy. 22, Muscatine, 21 Jul 2000, Cusick 
35601 (ISC). 

Tamarix parviflora DC. (TAMARICACEAE). Salt cedar is a widespread inva- 
sive species of riparian corridors throughout the western United States. It is a 
serious pest, often forming a monoculture that reduces natural biodiversity 
(Weisenborn 1996). The plants are isolated at the Des Moines County station, 
with no planted material nearby. Its presence in the Iowa flora should be closely 
monitored. Extirpation of the shrubs at the single known site probably is war- 

n Slough Rd., S of Burlington, 15Jul 2000, Cusick 

Viola arvensis Murray (VIOLACEAE). A single previous collection of European 
field pansy from Story County, Iowa in 1998 has been reported (Norris et al. 
2001). Unlike most violets, this is an annual species that may bloom in any 
month of the growing season. 

Cerro Gordo Co.: nursery, St. Hwy . 122, Mason City, 17 Jul 2000, Cusick 35585(150. Des Moines Co.: 

flowerbed, 1326Griswold St, Buriington, 4 Jul 1976, 1 Lammers 876 (ISCl 


I thank Diana H. Horton of the Ur 

iversity of Iowa and Deborah Q. Lewis of 

Iowa State University for their assist 

ance with this article. My research was sup- 

ported in part by the Division of ^ 

atural Areas and Preserves, Ohio Depart- 

ment of Natural Resources. 

:row, G.E. 1978. A taxonomic revision of Sagina (Caryophyllaceae) in North / 

Rhodora 80:1-91. 
iLERS, L.J. and D.M. Roosa. 1 994.The vascular plants of Iowa. Univ. of Iowa Press, lo 
ouRTEiG, A. 1979.0xalidaceae extra-austroamericanae. II. Oxc?//s section Cornicuk 

Pfiytoiogia 42:57-1 98, 
JoRRis, W.R., D.Q. Lewis, M.R Widrlechner, J.D.Thompson, and R.O. Pope. 2001 . Lessons 

inventory of the Ames, Iowa, Flora (1 859-2000). J. Iowa Acad. Sci. 1 08:34-63. 

Veisenborn,W. 1996.Tamarisk. In: J. M.Randall and J. Marinelli,eds. Invasive plants. V^ 

the Global Garden. Brooklyn Bot. Gard. Handbook 1 49, Brooklyn, NY. Pp. 43-4'^ 
VuNDERLiN, R.R 1 997. Moraceae. In: Flora of North America Edit. Comm. (eds.). Flora ( 

America. Vol. 3. Oxford Univ. Press, New York and Oxford. Pp. 388-399. 


Bruce G. Galdwin, Steve boyd, Barbara]. Erttf.r, Robert W Patteson, Ti iomasJ. Rosatti, 
and Dieter H. Wii_ken (eds), Margriet Wetherwax (managing ed). 2002. The 
Jepson Desert Manual. Vascular Plants of Southeastern Califonria. (ISBN 
0-520-22775-1, pbk.). University of California Press, Berkeley, CA 94720, 
U.S.A. (Orders: California Princeton Fulfillment Services, 1445 Lower 
Ferry Road, Ewmg, NJ 08618, U.S.A., 609-883-1759, 609-883-7413 fax, 
wwvv^ $35.00, 624 pp., 128 color photos, many line drawings. 

R Lhsica. 2002. A Flora of Glacier National Park. (ISBN 0-87071-538-0, pbk.). 
Oregon State University Press, 101 Waldo Hall, Corvallis, OR 97331-6407, 
U.S.A. (Orders: 800-426-3797, 541-737-3170 fax,, $32.95, 480 pp., 8-page color insert, 
60 color photos, 351 line drawings, 6" x 9". 

viD H. Engel and Suchart Phummae 2002. A Field Guide to Tropical Plants of 

Asia. (ISBN 0-88192-542-X, pbk.). Timber Press, Inc., 133 S. W Second Avenue, 
Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:, 
800-327-5680, 503-227-2878, 503-227-3070 fax). $19.95, 280 pp., app.rox. 
390 color photos, 8 figures, 6" x 9". 

' Words: Asia, tropical plants, field guide, flora, Identification, descriptions, local names, over 300 
The flora is broken down by trees, shrubs, vines, groundcovers. and bedding plants. Under each 

EricSundell R.DaleThomas 

Department of Biology 

Carl Amason 

The authors provide a list of 41 taxa representing additions 
to the Arkansas flora as well as noteworthy range extensions within the state. 
Herbarium abbreviations are taken from Holmgren et al. (1990). 

Alternanthera paronychioides St. Hilaire. Thomas discovered large populations 
of this prostrate chaff -flower around the edge of borrow pits inside the Missis- 
sippi River levee, in Chicot and Lincoln counties, in August and September, 2000. 
Duplicate specimens were confirmed by Kenneth R. Robertson of the Illinois 
Natural History Survey in Champaign. 



Bowlesia incana Ruiz and Pavon is a decumbent annual of low woods, cleai 

ings, lawns, and other moist, weedy places, occurring sporadically in Louisian 

and eastern Texas (Thomas and Allen 1996, Diggs et al. 1999). We record its fin 

appearance in Arkansas from Chicot County in the southeast corner of the staD 

along a road bank as well as in alluvial woods along Indian Creek south c 


Voucher specimens: Chicot Co.: Thomas 164,552 (NLU)J64,589(NLU,UAM). 

Sanicula smallii Bickn. Smith (1988) documents this cryptic black snakeroc 

from two c 

ounties m 



itams. We add collections from the Oua- 

chita Mountain 


he Coastal Pla 

n and suggest that the species might be 
iguished in the field from the widespread 

succulent leaves 
has been recogn 
woodland specie 


In addi 

haracters: sessile, slightly larger fruit 
s, thickened roots are only helpful afie 
tion, in at least one of our populatio 

with S. canadensis much the more CO 

r the plant 
ns the two 

Voucher speci 





U). Lafayette Co.: Sundell, Thomas, & A 



Co.: Ed 


71,826 (U AM). Ouachita Co.: Thomas&Doffitt 163,219 (Nim 



s glabra L 

Smith (1988) documents smooth cat's ear, a 


ounties m south Arkansas. We supply voucher specimens for 
orts ("R" for Ashley and Bradley counties) and add eight addi- 
) the record 

nwayCo.:EJvvin/JSmifh4032(UAM).LafayetteCo.: Thomas.SundelI(^Amason 
Co.: Thomas 160,883 (NLU). Nevada Co.: Thomas & Slaughter 104,341 (NLU, 
rhomas&Do[fiUl69,040{NL\Jl Sexier Co.-.Thomas.Sundell&Amason 166,304 
^dell, Thomas. & Amason 8210(11 AM). 

. In May, 1999, lavender-flowered Barbara's buttons 
own in Arkansas from eleven occurrences recorded in the Arkansas 
i Heritage Commission data base. During a field study commissioned 
ieritage Commission, ten of those populations, scattered in glades and 
parian communities m the Ozark and Ouachita highlands, were relo- 
id several voucher specimens made. 

pcL imens IM. cacspitosa Nutt van caespitosa): Carroll Co.: Su nde! ;& Wallace J2,6I7 (UAM). 
Co.: nalton i^ now la ^UARKJ Montgomery Co.: Sundell & Wallac 

Tragopogon pratensis L. lacks the inflated peduncles of T. dubius. 

goat's beard o 

the roadsides of north Ark 

ansas. I. pratensis is w 

dely estab 

lished in the n 

orthern U.S., extending south 

to Tennessee (Cronqui 

1 1980), bu 

has not previously been reported from Arkansas. 


Nandina domestica Thunb. A candidate for the most popular ornamental shrub 
in south Arkansas, "heavenly bamboo" is not documented as a naturalized spe- 
cies of the state flora. It both persists after cultivation and escapes (presumably 
dispersed by birds) to open areas and wood margins where it is sufficiently com- 
petitive to reach reproductive maturity. 


Cynoglossum zeylanicum (Hornem.) Thunb. ex Lehm. This previously unre- 
ported hound's tongue grew as an aggressive pasture weed in Sevier County in 
the foothills of the Ouachita Mountains. In addition, a second collection is re- 
ported from sandy soil in Union County. The species is known in the Southeast 
from Texas (Jones et al. 1997; Diggs et al. 1999) and Louisiana (from Claiborne 
and Union parishes along the Arkansas border) (Thomas and Allen 1996). The 
burs (mericarps) are 3-4 mm long, smaller than those of C. virginianum and C. 


Wahlenbergia marginata (Thunb.) DC. was first reported for Arkansas (Thomas 
et al. 1991) from a road bank m Union County. The species was rediscovered m 
2000 along railroad tracks on the south side of Camden. 


Murdannia keisak (Hassk.) Hand.-Mazz. We add two more documented occur- 
rences of this easily overlooked, rambUng, succulent-stemmed herb, previously 
known in Arkansas from two locahties (Sundell et al. 1999). 


I Jacq. The pink bindweed treated by Fernald (1950) and 
nosa L. forma purpurata Fern, is recognized by 
Vustin (1978) as a natural hybrid between I. lacunosa and I. trichocarpa Ell. (I. 
ordatotri loha Dennst.). We add a third county record to the two cited in Smith, 
.11 three in the Mississippi Delta of southeast Arkansas. 

Ipomea cordatotriloba Demist. We add three county records, from agricultural 
sites, to the documented Arkansas distribution (Lafayette County Smith 1988) 
of this weedy purple morning glory. 


Citrullus lanatus (Thunb.) Matsum. & Nikai var. lanatus. Six collections from 
southeast Arkansas document the status of watermelon at least as a waif in the 
state flora. 

Voucher specimens: Bradley Co.: Thomas & Amason J4J,965(Nm UAM). Calhoun Co.: Sundell etal. 
12,536 (UAM). Chicot Co.: Thomas 164.414 (NLU). Lincoln Co.: Thomas & Sundell 167,480 (NLU). Oua- 
chita Co.: Thomas & Doffin 167.999 (NLU). Union Co.: Thomas 111.808 (NLU). 

Eleocharis montevidensis Kunth is known from northern Louisiana (Thomas 
and Allen 1993) and included in Smith's Keys to the flora o/Ar/?ansas(1994) as a 
possible addition to the state flora. We document its occurrence with a collec- 
tion from Lake June in Stamps, in southwest Arkansas. 

Websteria confervoides (Poir.) Hooper (ScirpusconfervoidesPoir. in Lam.). Walker 
and Campbell (1997) first reported this submersed, aquatic sedge in Arkansas 
from a Jefferson County collection. We add two collections from Union County 
The species is rather broadly distributed in Louisiana (twelve parishes, four 
along the Arkansas border; Thomas & Allen 1993) and probably more wide- 
spread in southern Arkansas than we know. 
Voucher specimensjeffcrson Co.: Walkerl206961{VAM).VnionCo.: I homas& Amason H.3„5J2(NLU, 


Bergia texana (Hook.) Seub. ex Walp. Collections from the banks of the Red 
River and the Arkansas River represent the third and fourth confirmed occur- 
rences of this species in Arkansas. 

Voucher specimens: Lafayette Co.: Thomas 120.776 (N LU). Lincoln Co.: Thomas (^ Su ndel n67,42J (NLU). 

Caperonia palustris (L.) St.-Hil. is a wetland weed of tropical American origin 
known to occur in the U.S. in south Florida, Louisiana and southeast Texas 
(Godfrey & Wooten 1981). Smith (1988) has seen specimens from three Missis- 
sippi Delta counties in southeast Arkansas to which we add a fourth, where 
plants were collected from a rice field. 

Arkansas, its stems ascending above a mat of E maculata L. The species is pre- 
viously unreported for Arkansas. 

Voucher specimens: Bradley Co.: Sunddl 15.130 (BRIT-SMU, NLU, NY, UAM, UARK). 

Phyllanthus tenellus Roxb. was reported (Sundell et al. 1999) as a tentative addi- 
tion to tlie Arkansas flora from weeds escaping their containers at a Drew County 
nursery We add a second locality El hs's Nursery in Fountain Hill, Ashley County 
where again plants arriving as weeds m containers from Louisiana and Texas 
have spread aggressively throughout the nursery's grounds and greenhouses. 


Lathyrus aphaca L. Previously documented only from Miller County in south- 
west Arkansas (Smith 1988), this unique Eurasian peavine (the stipules function- 
ally replace the leaflets) was collected on a roadbank in central Arkansas. Estab- 
lished on the West Coast (Isely 1998), the species is apparently taking hold in the 
Southeast as well, (t was first reported for the region from Tennessee in 1972 
(Beardsley and Browne) and recently from Texas in 1997 (Jones and Reznicek). 
Thomas and Allen (1998) map collections from four Louisiana parishes. 

nctive Old World clover natural- 
: Texas (Isely 1990). It is reported 
w to Arkansas from Grandview Prairie, near Columbus, in Hempstead 
he Arkansas Game and Fish Commission is successfully restoring this 
. prairie site to its original vegetation. Plants were collected in a re- 
a dominated by compass plant {Si Iphium laciniatum) and purple and 
irie clovers {Dalea purpurea, D. Candida)— a spring alien among sum- 

Trifolium nigrescens Viviani. Ball clover is a sprawling, weedy annual from 
southern Europe and southwest Asia recently introduced into the southeast- 
ern United States (Isely 1998). Two collections from ruderal sites document its 
occurrence and extend its range as a naturalized species in Arkansas. Ball clo- 
ver IS easily mistaken for alsike clover (T. hyhridum) but distinguished by the 
smaller umbels and scarious (rather than membranous), more subulate stipules. 


s shrubby St. John's 
•t from Drew County m southeast Arkansas. We here confirm his report (no 
cher specimen was seen) of the species' presence in Ashley County with 
collections, and we document two additional counties. 


Melissa officinalis L. Smith (1988) records lemon balm as an adventive from 
several Ozark and Ouachita Mountain counties. We here make note of a collec- 
tion from the Coastal Plain. 

Voucher specimen: Grant Co.: Thomas & Sundell J64,430 (NLU). 

Hypoxis curtissii J. Rose (H. leptocarpa (Engelm. & Gray) Small) grows in 
swamps and f loodplain forests of the Atlantic and Gulf coastal plains (Godfrey 
and Wooten 1979). We record it as new to the state from Bodcaw Creek, in south- 
west Arkansas, where it occurred with Styrax americana, Planera aquatica, 
Fraxinus caroliniana, and Crataegus opaca under an overstory of Quercus lyrata. 

Lycoris radiata (LHer.) Herb. With its leaves appearing in autumn after the flow- 
ers, red spider lily is nicely adapted to the annual lawn mowing rhythms of 
most homeowners and has become a common lawn weed in southeast Arkan- 
sas as well as an occasional waif in other disturbed sites. We note its presence in 
the Arkansas flora with three collections. 

Thomas & Doffitl J67,947(NLU). 

Najas minor Allioni. Northeast Arkansas is included by Haynes (1979) withm 
the range of this distinctive Old World naiad, but neither localities nor speci- 
mens are cited. Thomforde's collection from the shallow margin of a goldfish 
pond m central Arkansas (21 August 2000) reconfirms the species' occurrence 
in the state. Recent north Louisiana collections unknown to Haynes (Thomas 
& Allen 1993) suggest that Najas minor, which apparently invaded North 
America around 1930, may still be extending its range to the west and is likely 
more widespread in Arkansas than we know. 


Mirabilis jalapa L. Reported by Leslie (1986) but excluded by Smith (1988) as a 

naturalized element m Arkansas, four o'clock is reinstated with roadside cok 

lections from the Ozark Highlands and the Coastal Plain. At no: 
did plants appear to be merely persisting after cultivation. Fi 
mapped by Thomas and Allen (1998) from six of eight of Louisia 
tier of parishes. 

Voucher specimens: Bradley Co.: Thomas & Amason 142,066 (NLU, UAM). Newto 
685 (NLU). Union Co.: Thomas & Amason 111,373 (NLU, UAM). 

Andropogon ischaemum L. var. songaricus Rupr. ex Fisch. & Mey iBothriochloa 
ischaemum (L.) Keng var songaricus (Rupr.) Celerier & Harlan), an intrusive, 
pernicious Eurasian bluestem, is common to the roadsides and pastures of the 
southern Great Plains (Gould 1975, Great Plains Flora Association 1986). It has 
become widespread in Louisiana (Thomas & Allen 1993) and an occasional 
weed east to Florida (Wunderlin 1998). We add three counties to its documented 
occurrence (Franklin Co.) in Arkansas. 

Ouachita Co. Thomas & Doffitt 168,023 (NLU). 

Eleusine tristachya (Lam.) Lam., a diminutive goosegrass of African origin, is 
known as a waif in North America from a few scattered localities (Hitchcock 
and Chase 1950, Kucera 1998) and is apparently naturalized in California (Smith 
1993). We report it as a new record from northeast Arkansas, where it occurred 
as a lawn weed. 


Fagopyrum esculentum Moench. Buckwheat spreads easily from culti 
waste places but seldom persists (Great Plains Flora Associati^ 
Steyermark 1963). Collections from a small population at the edge of £ 
parking lot mark the species' first documented occurrence as a waif ii 

Baldwin. Godfrc 
of this widespread sn 


and Wooton (1981) include Arkansas 
rtweed, and Smith (1988) made note of 
possible addition to the state flora based on its occurrence in 
la parishes along Arkansas' southern border (MacRoberts 1988). 
presence with voucher specimens from three counties. 
Chicot Co.: Thomas 163,358 (NLU).Hot Spring Co.: Sunden&Cranf2Ja733(UAM). 
Union Co.: Thomas 112,374 (NLU). 

Diodia dasycephala C.&S. Especially in the field, "perennial poorjoe" so closely 
resembles Spermacoce glabra (rather than either of the common, congeneric 

buttonweeds, D. teres and D. virginiana) that it has perhaps been overlooked 
and misidentified as often as any taxon in southern Arkansas. The two can be 
distinguished vegetatively. Leaves of the Diodia are shorter and dry greener; 
those of the Spermacoce are more prominently vemed beneath. Spermacoce 
glabra bears numerous, whitish scalelike hau's at the stipular sheath summit 
beneath and between the bristles while D. dasycephala possesses a (more or 
less) prominent vein that traverses the broader summit of the glabrous stipular 
sheath just below its bristle-bearing margin. Diodia dasycephala is documented 
from six parishes in north Louisiana (Thomas & Allen 1998) and two counties 
in southeast Arkansas (Smith 1988). We add three more counties to the record, 

Voucher specimens: Chicot Co.: Thomas 164,170 (NLU). Faulkner Co.: Sundell & Thomas 15,954 (NY, 


Lindernia Crustacea (L.) E von Muell. is a wetland species, originally from south- 
ern Asia, that occurs sporadically in the coastal plain of the Southeast (Godfrey 
& Wooten 1981). Our collection represents the second documented occurrence 

Scoparia dulcis L. Thomas and Amason collected this tropical American weed 
in a clearcut. It is known regionally from the Gulf Coast states (Correll & 
Johnston 1970) but unreported for Arkansas. 

Veronica hederifolia L. is confirmed by Smith (1988) for two northwest Arkan- 
sas counties. We add four more counties to the record of this relatively distinc- 
tive speedwell. 

Veronica pohta LiKs ippc us sul 1 icicnth distUKtlmm V dgJCstlb to be recog- 

nizedasascpn UespcLics SpccmunssLLn h\ Smith(csptcially atLJARK), who 
meiges the two m the Aikansas Atl(;is(1988) should piobably be reexamined 
foi additional mateiial ol the toimei species We document the presence of V. 
pohta m Arkansas hom four counties 

Vouchei specimcnb Baxter Co Hyalt ]0630j{UAM) Drew Co Sundell ]0755{VAM) Garland Co.: 


Verbena montevidensis Spieng Thomas and Allen (1998) recoid this slender 
vervain from three parishes m central and eastern Louisiana. Four collections 
from south Arkansas suggest that it may have become established in this state 

(NLU). Chicot Co.: Thomas 166,906 (NLU). Union Co.: Thomas & Amason J].I,853 (NLU). 


Zannichellia palustris L. Horned pond weed is documented from three counties 

in the Ozark Plateau of northeast Arkansas (Smith 1988). Two collections of the 

submersed aquatic from central and southwest Arkansas extend its range to 

the Mississippi River Alluvial Plain and West Gulf Coastal Plain. 


glish abstract into Spanish. Thanks als( 
Extension Service for his efforts in colle^ 
Cynoglossum zeylanicum, to Hugh Thomforde of the Lonoke Agricultural Cen- 
ter for his collections of Najas minor and Zannichellia palustris, and to Judy 
Griffith of Ninestone Land Trust in Carroll County for leading us to a splendid 
population of Marshallia caespitosa in full bloom. 

AusiiN, DT. 1978. The Ipomoea batatas complex-l. Taxonomy. Bull, Torrey Bot. Club 105: 

ica L new to Tennessee and the South- 

search Foundation, Renner. 

University of North Carolina Press, Chapel Hill. 

of north central Texas. Botanical Research Institute of Texas, Fort Wor 

College Center for Environmental Studies, Sherman, Texas. 
FtRNALD, M.L 1 950. Gray's manual of botany, ed. 8. American Book Compan; 
Godfrey, R.K. and J.W.Wooten. 1 979. Aquatic and wetland plants c 

States. Monocotyledons. The Univ. of Georgia Press, Athens. 
GoDFREY,R.K.and J.W.Wooten. 1 981 .Aquatic and wetland plants of the southeastern I 

States. Dicotyledons. The Univ. of Georgia Press, Athens. 
Gould, F.W. 1975. The grasses ofTexas.Texas A & M University Press, College Statior 
Great Plains Flora Association. 1986. Flora of the Great Plains. University Press of H 

Haynes, R.R. 1979. Revision of North and Central American Najas (Najadaceae). : 

States Department of Agriculture, Miscellaneoi 
Goverr^ment Printing Office, Washington, D.C. 

Holmgren, P.K., N.H. Holmgren, and L.C. Barnei r. 1 990. 1 
of the world, ed. 8. New York Botanical Garden, Bronx. 

IsELY, D. 1 990. Vascular flora of the southeastern United States.Vol. 3, Part 2. Leguminosae. 
The University ofNorth Carolina Press, Chapel Hill. 

lsELY,D. 1998. Native and naturalized Leguminosae (Fabaceae)ofthe United States. Monte 
L. Bean Life Science Museum, Provo, Utah. 

Jones, S.D. and A.A. Rfznicek. 1 997. Lathyrus aphaca (Fabaceae), previously unreported for 
Texas. Phytologia 82:1-2. 

JoNES,S.D.,J.K.Wipff,and P.M.Montgomery. 1997.Vascular plants ofTexas.UniversityofTexas 
Press, Austin. 

Kucera, C.L 1 998.The grasses of Missouri, rev. ed. University of Missouri Press, Columbia. 

Lesue,S,A. 1 986. A preliminary survey of the vascular flora of Bradley County Arkansas. M.S. 
Thesis, Northeast Louisiana University [University of Louisiana at Monroe]. 

MacRoberts, D.T 1 988. A documented checklist and atlas of the vascular flora of Louisiana. 
Part III. Dicotyledonae. Fagaceae to Zygophyllaceae. Louisiana State University in 

Smith, E.B. 1 988. An atlas and annotated list of the vascular plants of Arkansas, ed. 2. Pub- 
lished by the author, Fayetteville, Arkansas. 

Smith, E.B. 1 994. Keys to the flora of Arkansas.The University of Arkansas Press, Fayetteville. 

Smith, J.R, Jr. 1993. Poaceae. In: J.C.Hickman, ed.TheJepson manual: higher plants of Cali- 
fornia. University of California Press, Berkeley. 

Steyermark,J.A. 1963. Flora of Missouri.The Iowa State University Press, Ames. 

SuNDELL,E.,R.D.THOMAS,C.AMASON,R.L.STucKEY,and J. Logan. 1999. Noteworthy vascular plants 

Thomas, R.D., E.B. Smith, E. Sundell, RE. Hyatt and C. Amason. 1 991 . Additions to the flora of 

Arkansas. Sida 14:483-491. 
Thomas, R.D.and C.M.Allen. 1993. Atlas of the vascular flora of Louisiana. Vol. I. Louisiana 

Department of Wildlife and Fisheries, Baton Rouge. 
Thomas, R.D. and CM. Allen. 1 996. Atlas of the vascular flora of Louisiana. Vol. II. Louisiana 

Department of Wildlife and Fisheries, Baton Rouge. 
Thomas, R.D. and CM. Allen. 1 998. Atlas of the vascular flora of Louisiana. Vol. III. Louisiana 

Department of Wildlife and Fisheries, Baton Rouge. 
Walker, S.A. and J. Campbell. 1 997. A floristic survey and annotated checklist of the Pine 

Bluff Arsenal. J. Arkansas Acad. Sci. 51:1 78-1 87. 
Wunderlin, R.R 1998. Guide to the vascular plants of Florida. University Press of Florida, 




Alexander Krings 

ierbarium, Department of Botany 
North Carolina State University 

Michael G. Burton 

Department of Crop Science 

North Carolina State University 

Raleigh, NC 27695-7620, US.A. 

Alan CYork 


rtment of Crop Science 

Raleigli, NC 27695-7620, U.S.A. 


tion of the federal noxious weed Commelina benghalens 
. in a soybean field in North Carolina. Voucher specime 

ns are cited and an updat 






A common paleotropical weed, Commelina henghalensisL. (Tropical spiderwort, 
Benghal dayf lower) has been previously collected in the southeastern United 
States only in Florida, Georgia, and Louisiana (Faden 1993; Thomas & Allen 
1993). The taxon has not been reported for North Carolina by Radford et al. 
(1968), Krai (1981), or Faden (1993, 2000). Federally listed as a noxious weed in 
1983 (Lasseigne 1983), C. henghalensishas primarily been found in citrus plan- 
tations, fields, yards, and other cultivated and disturbed sites in the southeast 
as far north as Georgia, as well as in California (Faden 1993). During a field sur- 
vey, a population of this weed was identified in a soybean field in Goldsboro, 
Wayne County, in the Coastal Plain of North Carolina. The infestation occurs 
over an area of less than 2 ha, north of SR 1008 and near the Southern Railway 
The density is quite high at the edges of the crop plots and lower in neighbor- 
mg (intensively managed) cotton and soybean plots. The weed was first no- 
ticed about three years ago, but was thought to be a different species (Commelina 
camli niana Walter occurs in a neighboring field). Representative material was 
prepared for deposit at NCSC, US, and USCH in Oct 2001. 


C, Dissection of spathe with buds, D, Dissection of spathe, si 

Perfect flower; G, seed from below ground fruit, H, Dissection of seed f 

(NY), C-H based on Seydel203 (MO) Illustration by Cathy Pasquale, o 


henghalensis L (Fig. 1) can be distinguished from its southeastern 
lerics by the often clustered, tnfundibuliform spathes, the relatively broad 
often bearing reddish hairs on the sheath apex, and the basally-borne, 

sometimes subterranean cleistogamous flowers in addition to chasmc 
flowers. An updated key, largely adapted from Faden (2000) follows: 

f sheaths not auriculate;! 


3etal bl 

ue to lilac or 

;ome locules 2-seeded, 

/ith erect to ascending 

stems; lami 

na lane 




3us flowers absent 

. Spathes uniformly colored, without contrasting veins;proxir 
colored as the others; capsules 3-locular; seeds reticulate o 

. Spathes typically distinctly falcate; distal cyme in larger spathes usually well- 
developed, 1 -several-flowered; seeds reticulate Commelina diffus 

. Spathes not falcate to only slightly so; distal cyme in larger spathes usually 
vestigial (rarely well-developed, 1 -flowered); seeds smooth to somewhat 

We thank: Jamie Hinton and Steve Hoyle (Nortfi Carolina State University) fo 
their attention to detail; Gene Cross and David Patterson (NC Dept. Agric. & 
Consumer Services) for assistance in obtaining necessary transport permits 
Rodney Young (APHIS) and Robert Faden (US) for confirming our identifica 
tion; Robert Faden and an anonymous reviewer for thoughtful comments oi 

R.B. 1993. The misconstrued and rare species of Commelina (Comnnelinaceae) in 

eastern United States. Ann, Missouri Bot. Card. 80:208-21 8. 

IB. 2000. Comme//na. In: Flora North America Editorial Committee, eds. Flora North 

?rica, Vol. 22. Oxford University Press, New York. Pp. 1 92-1 97. 

1981 .Some distributional reports of weedy or naturalized foreign species of vas- 

r plants for the southern states, particularly Alabama and middle Tennessee. Cas- 

■a 46:334-339. 

U.S. Dept. Agric, Washington, D.C. 
Radford, A.E., H.E. Ahles, and C.R. Bell 1 968. Manual of the Vc 

The University of North Carolina Press, Chapel Hiil. 
Thomas, R.D. and C.lVl. Ailen. 1993. Commelina benghalensi 

hyalina Boott (Cyperaceae),and C 

Louisiana. Phytologia 74:336-338. 


Arely Nava-Rojo Manuel Gonzalez-Ledesma 

and MariCela Gomez-Sanchez^ centra delnvestigaaonesBiologicas 

nd illustrations are provided, 

Agrostis is a genus of grasses that was described by J.F. Gmelin in 1791. The ge- 
nus is included in the Aveneae tribe and Alopecurmae subtribe (Pohl & Davidse 
1994; Valdes-Reyna & Davila-Aranda 1995). As presently circumscribed, the 
genus includes about 125-220 species (Acosta-Castellanos 1990; Beetle 1983; 
Clayton & Renvoize 1986; McVaugh 1983; Watson & Dallwitz 1992) which grow 
in temperate and cold regions from both hemispheres. In the tropics, they are 
mostly restricted to the high altitude mountains. Currently, the most recent 
list of species for Mexico (Espejo-Serna et al. 2000) shows that 23 species are 
registered for Mexico. While conducting an aquatic flora project of the Queretaro 
state Mahinda Martinez made a collection of subaquatic grasses from the 
Huimilpan and Amealco municipalities that proved to be Agrostis avenaceaJ.F. 
Gmel. This collection represents the first record of A. avenacea from Mexico 
and increases the number of species to 24. 

Agrostis avenaceaj h CjIT 
GmeU Bech.XandolL 
Blumea 37:230. 1992. 


Plants annual, cespitose. Culms erect, thin, breakable, 20-60 cm tall, 1-2 mm 
broad. Leaf blade flat, 5-20 cm long, 1-3 mm broad. Ligule 2-5 mm long. Panicle 
open, lax, 15-30 cm long; branches thin, slender, mferiors as longer as inflores- 
cence axis, spikelets on the tips of the branches. Glumes acute, 2-5 mm long, 1- 
nerved, scabrous on the keel. Lemma membranous, pilose, 1.4-1.6 mm long, 5- 
nerved, with a mid-dorsal once-geniculate awn. Palea membranous, 1.1-1.3 mm 
long, nearly as long as lemma, 2-nerved, tip bifid, callus pubescent. Chromo- 
some number: 2n = 28. (Fig. lA-F) 

Habitat —Agrostis avenacea grows on dry to moist soils, with abundant clay 
and from 15 to 2350 m in elevation. In Mexico, it is grows around small streams, 
temporary and permanent ponds, and at sites at 2280-2350 m in elevation, it is 
found as a member of the subaquatic vegetation (Fig 2B). 

Distribution.-This species is native to Austraha (Canberra, New South 
Wales, Queensland and Tasmania) and New Zealand. In the Missouri Botanical 
Gardens herbarium and VAST nomenclatural database (W^ TROPICOS) 
(, there are records of its introduction in South Africa, Argen- 
tina (Buenos Aires, depression of El Salado, Entre Rios and Santa Fe) and Chile 
(Isla de Pascua) (Nicora & Rugolo 1987), as well as, some temperate regions of 
the United States of America (California, Texas and Ohio), and the fiawaiian 
Islands (Hawau, Kauai, Molokai, Oahu) (Hitchcock 1950). Recently it was re- 
corded from San Jose, Costa Rica (Davidse 1998). In Australia, it is an abundant 
weed of inundated places (Vickery 1941). Nicora and Rugolo (1987) and Zuloaga 
et al. (1994) recognize this taxon in the segregate genus Lachnagrostis. 

In Mexico, Agrostis avenacea was collected from the municipalities of 
Huimilpan and Amealco, in the southern portion of the state of Queretaro (Fig. 

Fig. 1 . Agrostis avenacea J.F. Gmel. {M. Martinez 2936, QMEX). A. Habit. B. Floret. C. Caryopsis. D. Spikelet. E. Inflorescence 
branches. F. Indument of inflorescence branches. G-H. Leaf biade anatomy. G. Detail of transverse section (M. Martinez 
2936, QMEX). H. Abaxial epidermis as seen in surface view [M. Martinez 2953, QMEX). Stippling indicates the costal re- 

2A). This species has not been previously reported in any f loristic study from 
Mexico. This taxon was introduced to U.S.A (Hitchcock 1950; Davidse et al. 1998). 
In California and texas, plants of A. avenacea act like tumble weeds and its dried 
panicles are carried widely by wind and m the pioccss dispctsing seeds It is 
unknown whether the populations of A avcnatLd f lom Mexico is the lesult of 
long range wind dispersion orb) some othei mannci such as by vehicles pass 
mg thiough wetlands and picking up spikekts and panicles It is probable that 
this taxon is glow mgmothei states of Noi them Mexico as well 

12 S4 I N 100 oiD 1\\ 24Q0inMim 2Djulig% Mnii/ic. ,i:l )>)> Mpio Auk ilto 1 m5cirretera 
Amciico Sm IklLllon.o ihuco temponl 20 1080 LN 100 540 l\\ 2540 msnm 5 Nov 1996 
Ma,tuuzj667 (QME\ duplicates lor distribution to lEB ind MEXU) 

Agrostis avenacea is distinguished by its slender, thin and lax inflorescences. 
The spikelets of mature inflorescences are easily dispersed by the wind. 
Leaf blade anatomy 

The abaxial epidermis and inner structure of the leaf blade were studied. The 
epidermis and transverse sections were obtained following Gomez-Sanchez and 
Koch (1998). 

Ahaxial epidermis.— Differentiation of the costal and intercostal zones incon- 
spicuous. Long cells, fusiform, longer than wide. Stomata abundant, 37-39 ^m 
long, parallel-sided subsidiary cells, guard cells hidden by interstomatal long 
cells Papillae absent Prickles a few m the costal zone Sihca bodies, miciohairs 
and macrohairs absent (Fig IH) 

Transverse section —Outline of the blade U-shaped nodular Abaxial and 
adaxial sui lace with i ounded i ibs associated with all \ asculai bundles, abaxial 
and adaxiallui lows bioad and deep Vasculai bundles nine usually fust order 
vasculai bundles Outei \asculai bundle sheaths absent Sclcienchyma costal 

Agio\t I s avL nana is cleaily a mcmbei of the Pooidcae m its anatomical tea- 
tut es It IS a Non-Kianz species with C3photosynthetic metabolism (Hattersley 
i!sr Watson 1975) It has a t)pical 'pooid anatomy, chaiacteristic of those spe- 
cies that glow m tempeiate climates The leaf blade is nodular with iibs and 
deep fuuows m the abaxial and adaxial surlaces The limited development of 





abaxial and adaxial sclerenchyma explains the presence of the breakable and 
lax culms and suggests that Agrostis avenacea is adapted to wet habitats. 

Several taxonomic treatments of Agrostis exist (Acosta-Castellanos 1990; 
Pohl 1980; Pohl & Davidse 1994; Davidse 1998), however these treatments are 
partial and taxonomic affinities of the species are uncertain. The genus Agrostis 
urgently needs a world revision (Pohl & Davidse 1994; Davidse 1998). In addi- 
tion to the morphological, anatomical, micromorphological work, the develop- 
ment of molecular studies will be a very important contribution to circum- 
scription of Agrostis species. 

We are grateful to the curators of the ENCB, lEB, MEXU and QMEX herbaria 
who so kindly accommodated our visit, to Mahinda Martinez who provided 
the collections, to Clara Tinoco who prepared Figure lA-E and to Robert Jones 
and an anonymous reviewer for helpful comments. Research was supported by 
CONACyT through 29106-N project. 


Acosta-Castellanos, S. 20Q] . Agrostis. In: Rzedowski, J. and Graciela C. de Rzedowski, eds. 
Flora Fanerogamica del Valle de Mexico. Institute de Ecologia, Centre Regional del 

Hidraulicos- COTECOCA. Mexico. 
Clayton, W.D. and S.A. Renvoize. 1986. Genera graminum. Grasses of tfie World. Kew Bull 

Add. Ser. 13. London. 

Version Internet (W^FM). Universidad Nacional Autonorrra de Mexico. Mexico. 
Espejo-Serna, a., A.R. LOpez-Ferrari, and J. ValdEs-Reyna. 2000. Poaceae. In: Espejo-Serna A. anc 


de Refrencia. Parts IX-XI.Consejo Nacional de la Flora de Mexico. Mexico. Pp. 8-14. 
GOmez-Sanchez, M. and S.D. Koch. 1 998. Estudio anatomico comparativo de la lamina folia 

de Eragrostis (Poaceae: Chloridoideae) de Mexico. Acta Bot.Mex. 43:33-56. 
HATTERSLEY,PW.and L.Watson. 1975. Anatomical parameters for predicting photosynthetic 

pathways in grass leaves. Phytomorphology 25:325-333. 
Hitchcock, A.S. and A. Chase. 1 95 1 . Manual of the grasses of the United States. U. S. Dept. o 

Agric, Misc. Publ. No. 200: 1 05 1 pages (Second edition, revised by A. Chase) 

Arbor. Missouri Botanical Garden: 
NicoRA, E.G. and Z.E. Rugolo de Agrasar. 1 987. Los generos de gramineas de America Austral 

Llemisferio Sur. Buenos Aires, Argentina. 
Pohl, R.W. 1 980. Flora Costaricensis. Family # 1 5. Gramineae. Fieldiana Bot, n.s. 1 4:1-607. 

Valdes-Rlyna, J. and P. Davila-Aranda. 1995. Clasificacion de los generos de gi 
Mexicanas.Acta Bot.Mex. 33:37-50. 

Watson, L. and M.J. Dallwitz. 1 992. The grass genera of the World. C.A.C. Inter 

ZuLOAGA, F.O., E.G. NicoRA, Z.E. RuGOLo DE Agrasar, 0. MoRRONE, J. pENSiERO, and A.M. 

1994. Catalogo de la Familia Poaceae en la Republica Argentina. Monogr. 

Missouri Bot.Gard. 47:1 -178. 

Books Received/Notices 


Ancii.i.e Gloudhn and Cicely Tobisch. 1995. Orchids of Jamaica. (ISBN 976-640- 
002-4, pbk.). The University of West Indies Press, la Aqueduct Flats, Kingston 
7, Jamaica, WI (Orders: University of Oklahoma Press, 4100 28th Avenue, 
Norman, OK 73069-8218;, 405-307-9048 fax). $26.00, 
222 pp., numerous color photos, 6" x 9". 

Fritz HochstAtter. 2000 Yucca I (Agavaceae). (Dehiscent-Fruited Species) in the 
Southwest and Midwest of the USA and Canada with Seed Morphology by 
Dr. Gunter Hentzschel. English Translation by Chris Holland. (ISBN 300- 
005946-6, hbk.). Fritz Hochstattcr RO. Box 510201D-68242 Mannheim, 
Germany; Hamlyn Valley Box 11, Modena, UT 84753, U.S.A. (Orders:;;;; 0621-79-00332 fax). Price not 
given, 256 pp., 189 figures, with many color photos, line drawings, and maps, 
6 3/4" X 9 3/4". 


Barbara Perry Lawton. 2002. Mints: A Family of Herbs and Ornamentals. (ISBN 
0-88192-524-1, hbk.). Timber Press, Inc., 133 S.W. Second Avenue, Suite 450, 
Portland, OR 97204-3527, U.S.A. (Orders:, 800-327- 
5680, 503-227-2878, 503-227-3070 fax). $27.95, 272 pp., 61 color photos, 5 
b/w photos, 46 line drawings, 6" x 9". 


Ray Neyland 

Department of Biological & Environmental Sciences 

This paper updates and revises the original vascular Ilora survey of Calcasieu Par- 
ish, Louisiana of Neyland et al. (2000). Specifically the purpose of this study is to 
list new collections from the Parish and to correct errors from the original survey 

Since the publication of the original survey by Neyland et al. (2000) until 
present, plant collections from Calcasieu have continued. Voucher specimens 
vv^ere prepared using standard herbarium practices and are housed at the 
McNeese State University (MCN) Herbarium. Additionally specimens from 
other herbaria have been examined. Nomenclature follows Kartesz (1999). 

The following entries are new additions to the vascular flora of Calcasieu Par- 
ish. These taxa have not been reported previously as occurring in the parish. 
Specimen references follow the same format as in the original survey (Neyland 
et al. 2000). Authorities are abbreviated according to Brummitt and Powell 
(1992). Following the collection data is an abbreviation for the vegetative region 

where each species typically occurs: prairie (Pr); longleaf pineland ( 
though not a vegetation region, disturbed areas such as pastures, fallow 
ditches, urban areas and roadsides are collectively abbreviated (D). 


The following entries have been previously reported in the parish but were not 
collected during the original survey of Neyland et al. (2000). Therefore, these 
collections provide recent evidence for their presence in Calcasieu Parish. 

e hirta (L.) Millsp. (MCN) Neyland 1939 D 
Scutellaria drummondii Benth. (MCN) Neyland 1927 D 

Oxalis violacea L. (MCN) Neyland 1945 D, Pi, Pr 

The following entries were misidentified in the original survey. Subsequent 

annotations have resulted in the noted corrections. 

1. The specimen originally reported as Asdepias variegata L Neyland 1763 
Pi is Asdepias obovata Ell. Therefore, this species is removed from the survey 

2. The specimen reported disjuncus elliottii Chapm. Brooks 609 (MCN) D, 
Pi, Pr is Juncus acuminatus Michx. However, the species does occur in the par- 
ish as evidenced by the specimen designated asju ncus dlwttu Chapm. GUtner 
37(MCN)D, Pi, Pr. 

The most notable addition to the flora of Calcasieu Parish is the discovery of a 
specimen of Platanthera Integra m the Oakes Ames Herbarium at Harvard. 
This specimen was collected by R.S. Cocks in September, 1906; however, addi- 
tional collection information is absent from the herbarium sheet. P. integra is 
ed by The Natural Heritage Program of the l..ouisiana Department 
^" ""■■ 'hich is defined as rare and local throughout 

of Wildlife and Fishen 

the state or found locally m a restricted region of the state. This apparently is 
the only known specimen that has ever been collected in Calcasieu Parish. Be- 
cause suitable habitat for this species largely has been destroyed in the parish 
(Neyland et al. 1998), it may be extirpated. 

As a result of these revisions to the original vascular flora survey of 
Calcasieu Parish, the number of confirmed specific and subspecific entries for 
the parish is increased to 1,153. The number of reported but unverified entries 
for the parish is reduced to 131. 

1 thank Sara Thames, University of Louisiana at Monroe, Monroe, LA, John 
Miller, Lake Charles, LA, Bruce A. Some, Longleaf Ecological, Whispering Pines, 
NC and Lowell E. Urbatsch, Louisiana State University Baton Rouge, LA for their 

Brummitt, R.K. and C.E. Poweil. 1 992. Authors of plant names. Royal Botanic Gardens, Kew. 

and Greenland. 1 st. ed. In: Kartesz, J.T and C.A. Meacham, eds. Synthesis of the North 
American flora, Version LO. North Carolina Botanical Garden, Chapel Hili. 

Neyland, R., H.A. Meyer, and H. Harrington. 1 998. Woody vegetation of longleaf pine com- 
munities in Calcasieu Parish, Louisiana. Phytologia 85:51-60. 

NEYLAND,R.,B.J.HoFFMAN,M.MAYnELD,andLE.URBATSCH.2000. A vascular flora survey of Calcasieu 

Books Receivhd/Noticcs 


3RAN M. Whitelock. 2002. The Cycads. (ISBN 0-88192-522-5, hbk.). Timber Press, 
Inc., 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. 
(Orders:, 800-327-5680, 503-227-2878, 503-227-3070 
fax), $59.95, 532 pp., 505 color photos, 13 line drawings, 8 1/2" x 11". 

This is one encyclopedia of information on cycads. The 505 color plates are exceptionally nice, 
you are into cycads then you will want this book. 

HN Bryan. 2002. Bulbs, Revised Edition. (ISBN 0-88192-533-0, hbk.). Timber Press, 
Inc., 133 S.W Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. 
(Orders;, 800-327-5680, 503-227-2878, 503-227-3070 
fax). $89.95, 896 pp., 1171 color photos, 10 line drawings, 2 charts, 8 1/2" x 11". 

NE McGary (editor). 2001. Bulbs of North America. (ISBN 0-88192-517-9, pbk.). 
Timber Press and North America Rock Garden Society (Orders: Timber Press, 
133 SW Second Ave., Suite 450, Portland, OR 97204-3527, U.S.A.,, 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". 

Contents include: 1 ) The Genus Allium: 2) Amaryllidaceae of North America; 3) The Brodiaea 
[iance: Bloomena, Brodiaea, Dichehuemimu and Tritdeia; 4) The Genus Calochortus; 5) The Ge- 


Guy LNesom and Robert J. O'Kennon 

Study of previous collections brings to light records for two species (Asteraceae, 
Caryophyllaceae) previously unreported for the state of Texas. 

Anaphalis margaritacea (L.) Benth. 

Anaphalis margaritacea occurs over most of the western and northern United 
States and Canada— it has not been recorded from Texas, Oklahoma, and states 
of the southeastern USA (Kartesz 1999). In Mexico it is known from localities in 
northern Baja California (Wiggins 1980). It is reported from eight primarily 
montane counties m New Mexico (Martin & Hutchms 1981), where it occurs at 
7000-10,000 feet elevation, and from several prairie counties in Colorado (Great 
Plains Flora Association 1976). The record from the Texas panhandle reported 
here is a southeastward 'outlyer' from the main range but it is a relatively small 
step eastward from the known localities in New Mexico and Colorado. The 
Brewster Co. collection is disjunct southward about 350 kilometers from the 
closest localities in Lincoln Co., New Mexico. Other UNM collections (fide Jane 
Mygatt, UNM Collection Manager) confirm that Ora Clark was botanizing in 
Brewster County in the first week of August 1931, adding a degree of confidence 
that the Anaphalis collection did originate from there. 

Hale Co.: Plainview, H of Co-op Cotton Compress on side of railroad, sandy soil, 3350 ft, 11 Sep 1967, 

Gypsophila elegans M. Bieb. 

Gypsophila elegans (var. elegans) is native to Russia (Black Sea region), Crimea, 
Caucasus, eastern Turkey, and Iran (Caspian Sea region) and is adventive in 
western Europe and North America (Barkoudah 1962), where it is grown as a 
garden plant. Gypsophila elegans var. latipetala Barkoudah is known only from 
"Turkey and Transcaucasia" (fide Barkoudah 1962). Variety elegans is known 
from scattered localities in Canada and the USA (including California, Utah, 
Colorado, Kansas, Illinois, North Carolina (Kartesz 1999). The central and north- 
central Texas localities reported here are the southernmost known stations in 

A key in the treatment of Gypsophila for California (Hartman 1993) sepa- 
rates G. elegans from G. paniculata L and G. scorzonerifolia Ser. Gleason and 

Cronquist (1991, p. 127) included a brief description and comment regarding G. 
elegans, following the description of G. muralis L., noting that it "occasionally 
escapes from cult, in our range." It is possible that the Texas plants are from 
direct roadside seeding— the species is distributed as part of 'Texas/Oklahoma,' 
'Mountain,' 'Northeast,' Gulf Coast/Caribbean,' and 'Southeast' wildf lower seed 
mixes (see www.prairiefrontiercom/pages/pfmixpg/regional.html). Another 
website touts the species as "One of the easiest seeds to grow. This plant literally 
pops out of the ground in any soil, ready to bloom." A search for the Tarrant 
County plants at the Keller site in 2002 was unsuccessful, however, suggesting 
that the species is not successfully reproductive in north central Texas. 

The common name of Gypsophila elega ns is Annual Baby's Breath or Showy 
Baby's Breath. Many photos of the species are available on the internet, includ- 
ing an excellent one of a herbarium specimen ( 

of Co. Rds. 379 and 380, in ditch on NW side of liighway, with 'oats, wheat, bermudagrass, rescuegrass 
and bluebonnets,' 15 Apr 2001, Kunselman 23 (TAG). Tarrant Co.: along railroad tracks in township 
of Keller, NE of Fort Worth, Apr 2001, 0'Kennon (sight record only). 

The new records reported here are from among specimens in loans to BRIT from 
WIS, TAG, and UNM. The Tarrant County collection of Gypsophila was inde- 
pendently identified by Richard Rabeler Review comments of Lindsay Woo- 
druff are greatly appreciated. Jane Mygatt provided helpful information on Ora 
Clark and his collecting schedule. 



Gleason, H.A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada (ed. 2). New York Botanical Garden, Bronx. 

Great Plains Flora Association (R.L McGregor, coord. ;T.M.Barkley,ed.). 1976. Atlas of the flora 
of the Great Plains. Iowa State University Press, Ames. 

Hartman,R.L. 1993.Caryophyllaceae(exceptS/7ene).ln:J.Hickman,ed.The Jepson manual: 
higher plants of California. Univ. California Press, Berkeley. Pp. 475-497. 

lar flora of the United States, Canada, ai 

il Garden, Chapel Hill. 

I.e. and CR. Hutchins. 1 981 . A flora of Ne\ 

.L. 1 980. Flora of Baja California. Stanforc 

I Greenland (Ed. 
1 flora. Version 1.0. Nc 

Larry E.Brown I.Sandra Elsik 

Houston Community College Dept^ of Ecology and Evolutionary Biology MS 
1300 Holman, Houston, TX 77007, USA. P. 0. Box 1892 

and Spring Branch Science Center Herbarium Rice University 

8856 Westview Drive Houston, TX 7725 1, U.S.A. 

us raphanistrum, Calycanthusfloridus var. glaucus, Lyonia lu- 
crum, Lythrum lineare, and IMyrtus communis. Significant new 
dght: Hypochaeris glabra, Cyperus cephalanthus, Rhynchosia 

axa are listed alphabetically by family, genus, and species with 

Flonstic work m the Houston area and other east Texas counties, coupled ^ 
plants sent to the SBSC herbarium for identification, has revealed a numbs 
plants new to Texas and significant new records for others. 

Unless otherwise indicated, all collections cited in this paper are in 
Spring Branch Science Center Herbarium (SBSC), Houston, Texas. Duplic 
of some collections are at the other indicated herbaria. Herbarium acron 
follow Holmgren et al. (1990). 

Aster laevis L. var purpuratus (Nees) A.G.Jones.— Aster laevis has been repre- 
sented in Texas by the western variety gejeri in the Guadalupe Mts of Culberson 
Co. (Nesom 1993). Here we report variety purpuratus, the narrow leaved 
southeastern entity, from two east Texas counties. Thomas and Allen (1996) 

mapped it in six, mostly north central, Louisiana parishes. Kartesz (1999) 
mapped it, as Symphyotrichum laeve (L.) A. & D. Love var purpuratum (Nees) 
Nesom, in Georgia, Alabama, Mississippi, Louisiana, Arkansas, and Texas. At 
the Polk County site, A. laevis var purpu ratus is distinguished from the similar 
and more numerous plants of A. ookntangiensis Riddel by smooth leaves in 
contrast to the harshly scabrous leaves of A. ookntangiensis. 

intersection wkh i^M 92, 12 Jun 2000,./, Uggio s.n. 

Bidens alba (L.) A. DC. var radiata (Sch. Bip.) Ballard ex T.E. Melchert.-Ballard 
(1986) divided the B. pilosa complex into three mostly tropical American spe- 
cies, namely B. alba, B odorata Cav, and B. pilosa L. He mapped United States 
records of B. alba var radiata in Florida and B. pilosa in California. Wunderlin 
(1998) reported B.alba var radiata as common and distributed nearly through- 
out Florida. Gandhi and Thomas (1989) reported B. pilosa. in Louisiana but in- 
dicated that Ballard's description of Bidens alba var radiata was in concurrence 
with their description of B. pi losa. Thomas and Allen (1996) maintained B. pi losa 
in Louisiana and mapped it in thirteen parishes. We report B.alba var. radiata 
(sensu Ballard) from four Texas counties. There is a photograph of the taxon in 

Tveten and Tveten (1993). 

the bridge over the Trinity River i 
2000, Brown 24801. Harris Co.: wee( 

Hypochaeris glabra L.— Diggs et al. (1997) reported this European weed new to 
Texas from Williamson County. Since then additional collections have been 
made in other regions of the state. 

Tridax procumbens L.— Kartesz (1999)reported this exotic and federal noxio' 
weed in Florida and fiawaii. Wunderlin (1998) reported it frequent in the ce 
tral and southern peninsula of Florida. John Tveten recently sent a collectk 
of it from the Valley Nature Center in Hidalgo County for identification. R 
garding the history of the plant in Texas, the following communication w 

received from Ken King and Martin Hagne of the Valley Nature Center; "Seed 
was collected m the Brownsville area by a native grower several years ago. That 
grower is no longer in the area and we do not know the exact Brownsville loca- 
tion. The plants at the Valley Nature Center were planted here from those seed- 
Imgs. This comp is growing in empty lots throughout Weslaco and has been in 
the area for at least fifteen years. We have seen several local establishments 
around town. There are at least two south Texas counties that have communi- 
ties of this plant; Hidalgo and Cameron." Alfred Richardson (1995) did not in- 
clude this taxon for the Rio Grande Delta area. 


Raphanus raphanistrum L.— Rollins (1993) distinguished the genus Raphanus 
by a segmented fruit with the lower segment inconspicuous and seedless. The 
petal colors vary from white, yellow, lilac, to violet. Rollins (loc cit.) reported 
two introduced weedy species in North America, R. raphanistrum and R. sa- 
liva. He indicated that in R. raphanistrum the fruits are linear, longitudinally 
grooved, and strongly constricted between the seeds. The petals are yellow and 
fade to whitish or white. In R. sativus the fruit body is widest at the base, smooth 
or very slightly grooved, and is not or only poorly constricted between the seeds. 
The petals are usually purple. The petal veins are prominent in yellow flow- 
ered R. raphanistrum, which can distinguish it from yellow petaled plants of 
Brassica and Sinapsis, if one does not notice the small lower fruit segment. 
Kartesz (1999) mapped it in Texas fide Gould (1975a) but no voucher was pro- 
vided. It is not listed in Correll and Johnston (1970) nor any of the recent check- 
Texas counties. Thomas & Allen (1996) mapped R. raphanistrum in numerous 
northern and southern Louisiana parishes. Two of these parishes are adjacent 
to Texas. In contrast, R. sativus is known from only seven parishes. Wunderlm 
(1998) found R. raphanistrum more common in Florida than R. sativa. A reex- 

Voucher specimens: Fort Bend Co.: yellow flowered weedy plant along a gravel road above a drainage 
ditch at extreme SW edge of Barker Reservoir, ca. 1/2 mi E of Mason Rd and ca. 1/2 mi N of FM 1093, 
10 Mar 1995, Brown 18435. Harris Co.: Brays Bayou at Buffalo Speedway in Houston, 18 Mar 2000, D. 
Johnson s.n. San Jacinto Co.: collected along roadside of Hwy 945 near the Evergreen Cemetery in the 


Calycanthus floridus L. var. glaucus (VVilld.) Torr & A. Gray— At least fif 
plants of this shrub are present in woods adjacent to the David and Pat: 
Lewis residence in Newton County. In a personal communication, the Lew 
indicated that these plants were here when they established a residence or 

site. Thomas and Allen (1996) were unsure of the native status of a collection 
from West Feliciana Parish, Louisiana. Likewise we are uncertain of their na- 
tive status here; however, they are well established and naturalized shrubs and 
should be considered a member of the Texas flora. 

Voucher specimens: Newton Co.: plants on a mesic slope woodland above Screw Pm Branch adjacent 
to CR 3062, W of Hwy 87 N of Bleal<wood. 22 Jul 2000, Brown & ELsi I? 24437 (SBSC, BRIT); same site, 
13 Apr 1999, Lewis, Lewis & Eisik 6169 (SBSC, TEX). 


Cyperus cephalanthus Torr. & Hook.-Carter and Mclnnis (1993) reported the 
type locality of this sedge from somewhere near Galveston Bay The type col- 
lection was made in 1835 and the species has not been recollected in Texas since. 
We here report a recent collection from Harris County Richard Carter (VSC) 
has verified the identification. 

dred feci SF of the Texas Chiropractic College, on the S side of Spencer Blvd, a few hundred feet E of 
Beltway 8 in Pasadena, 29-39'3g"N and 95-08'50"W, 29 May 1999, Brown 23076 (BRCH, SBSC, TEX). 

This site is owned by the Harris County Flood Control District and is soon to be 
formed into a detention basin for downstream flood control along Armand 
Bayou. However, Ralph Taylor of the Flood Control Environmental Services has 
removed all plants from the site and replanted them in their nursery near Lake 
Houston. Plants have been given to the Mercer Arboretum and Botanic Gar- 
dens and to the Armand Bayou Nature Center for estabhshment on their lands. 


Lyonia lucida (Lam.) K, Koch.— Specimens of these shrubs, with flowers in March 
and fruits in summer, have been recently collected in Newton County. The 
shrubs were on the property of David and Patricia Lewis when they set up a 
residence north of Bleakwood. There was no evidence of a prior human occu- 
pation of the site. David has discovered six populations with well over a hun- 
dred total plants (per comm.) along Screw Pin Branch, Fetterbush is native and 
common in the adjacent Sabine Parish of Louisiana (personal observation by 
the second author, Thomas and Allen 1996). It is our opinion that this is also a 

Brown & Elsife 2H36. (SBSC, BRIT), 


Rhynchosia tomentosa (L.) Hook. & Arn.-Turner (1959), Correll and Johnston 

(1970), Johnston (1990), Hatch et al. (1990), Jones et al. (1997), and Isely (1998) 

all reported this legume for Texas. Turner (1959) reported a collection by Charles 

Wright that lacks locahty data. Isely (1998) did not map it in the state. It is also 
not mapped in the state by Turner et al. (in press). We here report an authentic 
Texas collection from what is now the Little Rocky Preserve of The Nature Con- 
servancy of Texas. 

Voucher specimen: Jasper Co.: Little Rocky Nature Conservation area, 13 mi N of Jasper on Hwy 96 


Quercus arkansana Sarg.— Hunt et al. (1995) first reported this oak as new to 
Texas. They found two small populations in Cass County and an historical 
record for Jasper County. We here report recent collections from Jasper and 
Hardin counties. Jensen (1997) mapped it in the extreme northeast and south- 
east portions of the state. 

Jasper m the headwaters of Hog Creek, 3 Jul 2001, E. Keith Uh. 


Hyptis mutabilis (Rich.) Briq.-Kartesz (1999) reported this exotic weed in seven 
southeastern states from Virginia to Louisiana. Thomas and Allen (1998) 
mapped it in three parishes east of the Mississippi. River We here report a col- 
lection for Texas. 



Lilium longiflorum Thunb.-In the summer of 2000, we found this tall white 
flowered plant to be common and conspicuous along roadsides near Newton. It 
is also found as an escape from cultivation in Alabama, Florida, and Utah 
(Kartesz 1999). We identified it to this species with the key m Wundedm (1998). 
Thomas and Allen (1993) reported L.formosum in Louisiana, but our plants have 
the totally white flowers of L. longiflorum, in contrast to the red-purple outer 
perianth of L.formosum. 

N of jet Hwy 190/FM 256 W of Woodville, 3 Aug 1991, SchuUz 061. 


Cuphea viscosissima Jacq — Correll and Johnston (1970) reported Blue Wax weed 
in Texas but without a specific locality. Graham (1975) eliminated it from Texas, 
as did Johnston (1990), Hatch et al. (1990), Jones et al. (1997), Kartesz (1999), and 
Turner et al. (in press). We here report a voucher to confirm its presence in Texas. 

Lythrum lineare L.-Correll and Johnston (1970), Graham (1975), Johnston (1990), 
Hatch et al. (1990), Jones et al. (1997), and Turner et al. (in press) did not report 
this species in Texas. However, Kartesz (1999) listed Texas within its distribu- 
tion. We provide two vouchers to support its presence in the state. The identifi- 
cation was verified by S. Graham (KE). 

Voucher specimens: Chambers Co.: Anahuac National Wildlife Refuge, brackish marsh in the Jackson 
Ditch Unit, 2935'24"N and 94"25'09"W, 3 Oct 2000, Brown 24774; Anahuac National Wildlife Refuge, 
brackish marsh in the East Unit, 2939'N atid 94-28'W, 20 Jul 2000, Brown 24399 (SBSC, KE). 


Myrtus communis L.— Kartesz (1999) did not report this cultivated ornamental 
plant as occurring wild in the United States. The first author found many well 
developed shrubs scattered across a salt dome in coastal Texas. These plants 
had mature fruits and were rather large, suggesting they have been here for some 
time. The salt dome is isolated and surrounded by a coastal prairie and is for the 
most part undeveloped except for some old machinery and buildings associ- 
ated with prior oil extraction. 

Voucher specimen: Brazoria Co.: shrubs with fruits along gravel road to the center of Hoskins Mound, 
Galveston-Brazoria Key Map 861 13 and H, near 29"08'26"N and 95'13'26"W, 6 Aug 2000. Brown 24588 


Oenothera elat 


1 subsp. hirsut 


(A. Gray e> 

S. Wats.) W. Dietr.- 

Dietrich et al.(1997) im 


this taxor 


y m Trans- 

Pecos Texas with 


collection from 

the Pa 

nhandle. They 

also m 

apped thre 

2 populations m 


Texas in Anders 

on, Bra 

zos, a 

nd Leon c 


tchmg the key tr 


and description 

of this 


have bee 

1 colle 

cted m Har 

ris and Liberty cc 


ties. They are m 

ore CO 


on sandy soil 

n Liberty County These are 


plants with larg 

e yellov 


ers whose 


as are elevated above the stam 


Voucher specimens: 

Harris C 

a.: tall 

herb along H 



River bridge, 28 Jul 


Gregg s.n. Liberty Co 

: plants 


on sands ale 

ng the Trinity River at 

he Hwy 105 bridge w 

St of 

serve, 30"28'59"N and 94-5019"W, 25 Aug 2000, Brown 24680a; tall plants with rather large 1 lowers on 
deep sands along the Trinity River, Trinity River Nat Wildlife Refuge at Sevenmile Bend on the Davis 
Hill 7.5'quad. 30 Aug 1996, Brown 19528; large flowered primrose on dry sandy soil along CR 2252 
near the Davis Hill Baptist Church E of Cleveland and N of Hwy 105, 15 Sep 1996, Brown 19693; a 
single tall plant at edge of Hwy 146, ca 3 mi N of Moss Hill, 21 Aug 1994, Brown 18124; tall weedy 


Leptochloa dubia (H.B.K.) Nees.— Gould (1975b) reported this grass from all re- 
gions of the state except for the east Texas pineywoods and the post oak savan- 
nah area. We here report collections from two east Texas counties. 

at the intersection of Underwood St. and Fairmont Parkway in Deer Park, 29 Sep 1984, Brown 8100. 

We wish to thank Tom Wendt (TEX) and Leslie Landrum (ASU) for the identi- 
fication of Myrtus communis, Richard Carter (VSC) for the verification of 
Cyperus cephalanthus, and S. Graham (KE) for the verification of Lythrum 
lineare. Michael MacRoberts provided a hterature source and Guy Nesom (BRIT) 
helped with some nomenclatural questions. Justin Williams (SHST) and Mo- 
nique Reed (TAMU) reviewed the paper and provided many helpful suggestions. 
The first author is thankful to the Houston Community College for a sabbati- 
cal leave during the spring semester of 2002 that allowed him to complete this 
project. Publication funds were provided by Rice University. 

Ballard, R. 1 986. Bidenspilosa complex (Asteraceae) in North and Central America./ 

Bot. 73:1452-1465. 
Carter, R. and N. McInnis. 1 993. Final status report for Cyperus cephalanthus T & G. L 

lished report. U.S. Fish and Wildlife, endangered species office. Jackson, MS. 
CoRRELL, D.S. and M.C. Johnston. 1 970. Manual of the vascular plants of Texas. Te> 

search Foundation, Renner,TX. 
Dietrich, W.,W.L.WAGNER,and RH. Raven. 1997. Systematics of Oenothera section Oer 

subsection Oenothera (Onagraceae). Syst. Bot. Monogr. 50:1-234. 
DiGGs, G.M., R.J. O'Kennon, and B.L Lipscomb. 1 997. Hypochaeris glabra, a new Asterac 

Gandhi, K.N. and R.D.Thomas. 1 989. Asteraceae of Louisiana. Sida Bot. Misc. 4:1-202. 
Gould, F.W. 1975a. Texas plants —a checklist and ecological summary. Revised e 

Texas Agric. Exp. Sta. College Station. 
Gould, F.W. 1 975b.The grasses ofTexas.Texas A&M University Press. College Static 

JRAHAM, S, 1975. Taxonomy of the Lythraceae in the 

lATCH, S. L, K.N. Gandhi, and L.E. Brown. 1 990. Checklis 

1 655. Texas Agric Exp. Sta., College Station. 

Botanical Gardens. Bronx. 
luNT, D.M., M.H. MacRobfrts, and B.R. MacRobfrts. 19 
Sarg.(Fagaceae) in Texas. Phytologia 79:22-24. 

J, R.J.I 997. Ouercu. 
/ Press. New York, 

]. Privately published I: 

Jones, S.D., J.K. Wipff, and RM. Montgomer 

Y. 1 997.Vascular plants of Texas. A comprehensive 

checklist including synonymy, biblio 

graphy,and index. University of Texas Press. Austin. 


st and atlas with biological attributes for the vas- 

cular flora of the United States, Canada, and Greenland. First edition. In J.T Kartesz and 

C.A.Meacham, Synthesis of the Nor 

h American Flora, Version 1 .0. North Carolina Bo- 

tanical Garden, Chapel Hill. 

Nesom,G.L. 1 993.Three species of Aster (/ 

steraceae:Astereae) disjunct in northern Coahuila, 

Mexico. Phytologia 74:296-304. 

Richardson, A. 1 995. Plants of the Rio Grande Delta. University of Texas Press. Austin. 

RoLUNS, R.C. 1 993.The Cruciferae of con 

inental North America. Stanford Univ. Press. Stan- 


Thomas, R.D. and CM. All EN. 1993. Atlas c 

f the vascular flora of Louisiana. Vol. 1: ferns & fern 

allies, conifers, & monocotyledons. L 

uisiana Department of Wildlife & Fisheries. Baton 


Thomas, R.D. and CM. Aum 1 996. Atlas 

of the vascular flora of Louisiana. Vol. II: dicotyle- 

dons, Acanthaceae - Euphorbiaceae. 

Louisiana Department of Wildlife & Fisheries.Baton 

Thomas, R.D. and CM. Ai i en. 1 998. Atlas of the vascular flora of Louisiana. Vol. Ill: dicotyle- 
dons, Fabaceae-Zygophyllaceae. Louisiana Department of Wildlife & Fisheries. Baton 

Turner, B.L 1 959.The legumes of Texas. University of Texas Press. Austin. 

Turner, B.L.,H. Nichols, O.DoRON,and G.C.Denny. In press. Atlas of theTexas flora Vol.1 Dicots 

and Vol.2 Monocots.Sida Bot.Misc. 
TvETEN,J.L.andG.A.TvETEN.1993.Wildflowers of Houston and southeast Texas. University of 

ERRATUM FOR SIDA 18(4):1227-1245. 1999. 

Allan Nelson 

Department of Biological Sciences, Box T-OlOO 

Negrete et al. wish to correct their paper, A checklistfor the vasci 
Padre Island National Seashore pubhshed m the December 1999 i: 
(volume 18, number 4, pp. 1227-1245) by deleting Sesuvium trianthe 
the checkhst, which will change the total number of species to 4! 


The New York Botanical Garden is pleased to announce that Aaron Liston, cur- 
rently at the Department of Botany & Plant Pathology, Oregon State Univer- 
sity, is the recipient of the Rupert Barneby Award for the year 2002. Dr. Liston 
will be studying the phylogenetic systematics of Astragalus and Trifolium. 
The New York Botanical Garden now invites applications for the Rupert Barneby 
Award for the year 2003. The award of US $1,000.00 is to assist researchers to 
visit The New York Botanical Garden to study the rich collection of 
Leguminosae. Anyone interested in applying for the award should submit their 
curriculum vitae, a detailed letter describing the project for which the award is 
sought, and the names of 2-3 referees. Travel to the N YBG should be planned for 
sometime in the year 2003. The application should be addressed to Dr.James L. 
Luteyn, Institute of Systematic Botany, The New York Botanical Garden, 200th 
Street and Kazimiroff Blvd., Bronx, NY 10458-5126 USA, and received no later 
than December 1, 2002. Announcement of the recipient will be made by 
December 15. 

Anyone interested in making a contribution to THE RUPERT BARNEBY FUND 
IN LEGUME SYSTEMATICS, which supports this award, may send their check, 
payable to The New York Botanical Garden, to Dr Luteyn. 

rco Lambertini (Translated by John Venerella). 2000. A Naturalist's Guide to 
the Tropics (ISBN 0-226-46828 3 pbk) The University of Chicago Press 
5801 S Ell s A e Ch ago IL 60637 1 496 U S A (O de 773 702 0279 773 
702 7956 fax da@p uh ag du^^w^^pe uh ago edu) $25 00 338 
pp 57 olo photo 11 olo plat nu Tie ou b/w halftone 5 Ip 1 1/2 

d h bl 1 

Id g d E gl 

lyd pp d II 

The last two chapters-conservation and traveUng precautions-make this b( 

My recommendation is; Buy this book and read it in depth, book a tropical 
e good field guides for that specific region actually to take with you. And, £ 
m viaggio!"-Roger W. Sanders, Associate CoUectionsManager, Botanical Researc 


David Biek. 2000. Flora of Mount Rainier National Park. (ISBN 0-87071-470-8, 
pbk.) Oregon State University Press. 101 Waldo Hall, Corvallis, OR 97331- 
6407. (Orders: http;//, 800-426-3797, 
541-737-3166, 541-737-3170 fax). $29.95, 520 pp, 64 color photos, line drawings, 
6" X 9". 

bibliogiaphy cite:^ c 

Ls a print too small to be useful. For example, the illustration for P. 
55 section of both leaf and stem. Both of these figures are too s 
se, an illustration of a branch of Tsuga heterophylla is so small i 
s. Perhaps it would have been more useful had the author provide 
ion following the species descriptions that cross-referenced then- 

accessible to those with limited botanical 
T the inside cover for quick reference to its 

imen in the field. 

Without doubt, this book is an important update to the knowledge of the flora of Mount Rainier 
je who enjoy the flora of this area would certainly benefit by purchasing a copy of this book, 
;h should prove a worthy and practical field tool. This is a must-have for amateur and profes- 
al botanists who enjoy the Northwest United States.-Amy TrauthNare, Botanical Research Insti- 
ojTexas, Fort Worth, TX 76102-4060, U.S.A. 

ure. From reading t 

his discussion, it seem 


difficult to classify 

handy features that a 

mal amount of tech 


pledge. The author h 

as provided a map of t 

he Par 

nsfield, the ffora of Steens Mountain, Oregon incl 
:ar Fields to Riddle Mountain east of Diamond, the e 

The book's introduction begins 

short geologic and biogeographic hist( 
tain and an explanation of vegetation 

; iollowedbyabnef descuptionol each species common names 1 
listmg of families a useful glossai y follows, which clearly define 
cross refeiences line drawings that lollow These lUusti ations wou 

. 2001 The Illustrated Flora of Illinois. Grasses: Pani- 
, Second Edition. Illustrations by Miriam W. Meyer and 
Paul W. Nelson. (ISBN 0-8093-2360-5, hbk.) Southern Illinois University 
Press. P.O. Box 3697, Carbondale, IL 62902-3697. (Orders: 618-459-6633, 618- 
453-1221 fax). $50.00, 455 pp, 306 line drawings, 5 1/2" x 8 1/2". 
Robert H. Mohlenbrock. 2001. The Illustrated Flora of Illinois. Flowering Plants: 
Pokeweeds, Four-o'clocks, Carpetweeds, Cacti, Purslanes, Goosefoots, 
Pigweeds, and Pinks. Illustrations by Paul W. Nelson. (ISBN 08093 2380 
X, hbk.) Southern Illinois University Press. PO. Box 3697, Carbondale, IL 
62902-3697. (Orders: 618-459-6633, 618-453-1221 fax). $59.95, 277 pp, 139 
line drawings, 6" x 9". 
Robert H Mohlenbrock 2001 The Illustrated Flora of Illinois. Sedges: Cyperus 
to Scleria, Second Edition. Illustrations by Fredda Burton and Mark W. 
Mohlenbrock. (ISBN 0-8093-2358-3, hbk.) Southern Illinois University Press. 
PO. Box 3697, Carbondale, IL 62902-3697. (Orders: 618-459-6633, 618-453- 
1221 fax), $39.95, 223 pp, 128 line drawings, 5 1/2" x 8 1/2". 

ThcbC thicc books provide one new edition and two revised editions to the flora of Illinois. Each book 
bcgnii w ith a Lounty map of Illinois and a brief introduction about the groups covered within. Taxo- 

e morphology of grasses. The 2 
inois. The key to the genera c 

;s description includes only the most important features for identifi- 
vations of Illinois material. Also included are common name, habi- 
tribution and any pertinent taxonomic information. Following each 

ill genera of Cyperaceae except Carex. The Sedges bears many simi- 
distribution of sedges. Mohlenbrock also discusses the relationship 

1 in the Grossest 
) the Illinois f lor 

luable addition to the knowledge of tl 
Vlidwest United States s 
. Botamtal Research Im 

1 A ParrottA 2002 Healing Plants of Peninsular India. (ISBN 0-85199^501- 
2, hbk ) CABl Publishing, 10 East 40th Street, Suite 3203, New York, NY 
10016,USA (Ordeib 212 481-7018, 212-686-7993 tax, cabi-nao@cabi org) 
$140 00, 944 pp, numerous color photos 7" x 9 1/2" 

I extensive 186 pages of end matter follows the descriptions. A Bibliography is followed by a 

dex comes next, separated into the languages of the region. Also included are a Scientific 
dexandan Index of Medicinal Uses. 


certainly o 


he most e 





ed medicinal plant books 


Hopefully 1 







a standard for other region 



books. This 


ne should 



ken rea 

ders to 

he vast unexplored areas o 


plants. Man 


he species 



m this 


ve not been studied scient 


of great ph 


eutical va 





kes one fully aware of the 





ally dive 




y those 

containing useful and pote 


y healing pi 

Cole Wea 




ustin College, Sherman, TX 


I Research I 







HEN Foster and Christopher hobbs. 2002. Peterson Field Guide to Western 
Medicinal Plants and Herbs. (ISBN 0-395-83806-1, Flexi, ISBN 0-395-83807- 
X, hbk.). Houghton Mifflin Company, 222 Berkeley Street, Boston, MA 02116- 
3764, U.S. A. (Orders: 617-351-3243,, ww w.houghton $22.00 (Flexi), $30.00 (hbk). 448 pp, 530 + color pho- 
tos, 4 1/2" X 7 1/4". 

it surprised to find thick glossy pages, each usually presenting at least one color photograph. This 
ook places photographs immediately adjacent to the individual species descriptions, as other Peter- 
in guides have done. This is a very nice feature, as it can often be frustrating flipping back and forth 

lings are also grouped by various key flower characteristics. This system requires one to thumb 
irough practically every page until they find the section they are looking for A table of contents at 

ling or fruiting dates and 
>{ phnt descriptions oiga 

hyDtpl ojBwhi^y Austin Colk^c Sherman TX' 
tWorth TX76102 USA wweathaby@aushnc 

Books Received/Notices 

Timber Press' Gardener's Guide to Growing 

.haelJefferson-Brown and Harris Howland. 2002. Gardener's Guide to Growing 

Lilies. (ISBN 0-88192-537-3, pbk .; 0-88192-315-X, hbk.). Timber Press, Inc., 
133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Or- 
ders:, 800-327-5680, 503-227-2878, 503-227-3070 
fax). $29.95 (hbk, 1995), $19.95 (pbk, 2002), 160 pp., 70 color photos, 40 b/w 
Ime drawings, 7" x 9 1/2". 

,na Grenfell. 2002. The Gardener's guide to Growing Daylilies. (ISBN 0-88192- 
536-5, pbk.; 0-88192-46I-X, hbk.). Timber Press, Inc., 133 S.W. Second 
Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimber, 800-327-5680, 503-227-2878, 503-227-3070 fax). $29.95 (hbk, 
1998), $19.95 (pbk, 2002), 160 pp., 74 color photos, 5 Ime drawings, 7" x 9 1/ 

RTlN Page. 2002. The Gardener's Guide to Growing Peonies. (ISBN 0-88192- 
535-7, pbk.; 0-88192-408-3, hbk,). Timber Press, Inc., 133 S.W. Second 
Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimber, 800-327-5680, 503-227-2878, 503-227-3070 fax). $29.95 (hbk, 
1997), $19.95 (pbk, 2002), 160 pp., 76 color photos, 18 b/w line drawings, 7" 
X 9 1/2". 

aham Rice and Elizabeth Strangman. 2001. The Gardener's Guide to Growing 

Hellebores. (ISBN 0-88192-517-9, pbk.; 0-88192-266-8, hbk.). Timber Press, 
Inc., 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. 
(Orders:, 800-327-5680, 503-227-2878, 503-227-3070 
fax). $29.95 (hbk, 1993), $19.95 (pbk, 2001), 160 pp., 74 color photos, 20 line 
drawings, 7" x 9 1/2". 

-F Stebbings. 2001. The Gardener's Guide to Growing Irises. (ISBN 0-88192- 
519-5, pbk.; 0-88192-388-5, hbk.). Timber Press, Inc., 133 S.W. Second 
Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimber, 800-327-5680, 503-227-2878, 503-227-3070 fax). $29.95 (hbk, 
1997), $19.95 (pbk, 2001), 160 pp., 74 color photos, 3 line drawings, 7" x 9 1/2". 

JA Grenfell. 2001. The Gardener's Guide to Growing Hostas. (ISBN 0-88192- 
518-7, pbk.; 0-88192-355-9, hbk.). Timber Press, Inc., 133 S.W Second Avenue, 
Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:, 
800-327-5680, 503-227-2878, 503-227-3070 fax). $29.95 (hbk, 1996), $19.95 
(pbk, 2001), 160 pp., 74 color photos, 5 line drawings, 7" x 9 1/2". 

Ecology/Conservation/Landscape Ecology 

Robert H. Robichaux and David 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, Ste, 103, 
Tucson, AZ 85719, U.S.A. (Orders: 520-621-1441, 520-621-8899 fax). $50.00 
(hbk), 259 pp., numerous b/w photos, 6" x 9". 

trees, columnar cacf, Mayo people, crop diversity, amphibians and repuTes," birdT " ''''" ^ '''' " 

Barbara Tellman (ed.). 2002, Invasive Exotic Species in the Sonoran Region . (ISBN 

0-8165-2178-6, hbk.). The University of Arizona Press, 355 S. Euclid, Ste. 103, 

Tucson, AZ 85719, U.S.A. (Orders: 520-621-1441, 520-621-8899 fax). $75.00 

(hbk), 460 pp., 37 illustrations, 6 1/8" x 9 1/4". 

Rodney W. Bovey Woody Plants and Woody Plant Management: Ecology, Safety, 

and Environmental Impact. (ISBN 0-8247-0438-, hbk,). Marcel Dekker, Inc., 
270 Madison Avenue, New York, NY 10016, U.S.A. (Orders: 212-696-9000, 
212-685-4540 fax, www.deekercom). Marcel Dekker AG, Hutgasse 4, Postfach 
812, CH-4001 Basel, Switzerland (Orders: 41-61-261-8482, 41-61-261-8896 
fax) $195.00, 564 pp., 6" x 9". 

Ethnobotanical/Economic/Medicinal/Useful Plants/Toxic 

LA R. Emkry and rebeccaJ. McLain (eds,). 2001. Non-timber Forest Products: 
Medicinal Herbs, Fungi, Edible Fruits and Nuts, and Other Natural Prod- 
ucts from the Forest. (ISBN 1-56022-089-9, pbk.). Food Products Press, 10 
Alice Street, Binghamton, N Y 13904-1580 U.S.A. (Orders: 607-722-5857, 607- 
722-1424 fax, Price not listed, 176 pp., 6" 
X 8 1/4". 

AYAN P Manandhar. 2002. Plants and People of Nepal. (ISBN 0-88192-527- 
6). Timber Press, Inc., 133 S. W Second Avenue, Suite 450, Portland, OR 97204- 
3527, U.S.A. (Orders:, 800-327-5680, 503-227-2878, 
503-227-3070 fax). $69.95, 636 pp., 48 color photos, 3 tables, 834 line drawings, 
2 maps, 8 1/2" X 11". 

A'ords: Plants. Nepalese people, flora, ethnobotany, descriptions, common names. 
This is quite a book with fantastic color photos of plants and Nepalese people. There are no 


3THY ScHWlEDER, Thomas MoRAlN, and Lynn NlELSEN. 2002. Iowa Past to Present: 
The people and the Prairie. Third edition. (ISBN 0-8138-2998-4, hbk.). Iowa 
State Press, 2121 State Avenue, Ames, lA 50014, U.S.A. (Orders: 1-800-862- 
6657, 1-515-292-3348 fax; $32.99, 312 p 
b/w photos and other illustrations, 7" x 10". 

^RT McCoOK 2002 States of Nature. Science, Agriculture, and Environment 

in the Spanish Caribbean. (ISBN 0-292-75256-3, hbk.; 0-292-75257-1, pbk.). 
University of Texas Press, PO. Box 7819, Austin, TX 78713-7819, U.S.A. (Or- 
ders: 800-252-3206 fax; 512-471-4032). $50.00 (hbk), $22.95 (pbk), 201 pp., 
6 photos, 1 map, 6" x 9". 

I of Gerald Holton). 2002. Ivory Bridges: Con- 
necting Science and Society. (ISBN 0-262-19471-6, hbk.). The MIT Press, 5 
Cambridge Center, Cambridge, MA 02142-1493, U.S.A. (Orders: 800-405- 
1619,, $30.00 (hbk), 227 pp., 
6" X 9". 


and Kenneth J. Sytsma. 1997. Molecular Evolution and Adap- 

L (ISBN 1-521-57329-7, hbk.). Cambridge University Press, The 
Edinburgh Building, Cambridge CB2 2RU, UK; 40 West 20^^ s^^gg^^ ^^^ 
York, NY 10011-4211, U.S.A. $39.95, 621 pp., b/w photos, figures, 7" x 10". 

R.S. Singh. 2000. Diseases of Fruit Crops. (ISBN 1-57808-159-9, pbk.; 1-57808-149- 
1, hbk.). Science PubUshers, Inc., PO. Box 699, Enfield, NH 03748, U.S.A. (Orders:;, or $49.50 (pbk), 310 
pp., 31 b/w photos, 6" X 9 1/2". 

)RGt; G. KiiAci-iATcMiRiANS, Alan McHugen, Ralph Scorza, Wai-Kit Nip, and Y.H. 
Ulii (cds.). 2001. Transgenic Plants and Crops. (ISBN 0-8247-0545-9, hbk.). 
Marcel Dckkcr, Inc., 270 Madison Avenue, New York, NY 10016, U.S.A. (Or- 
ders: 212-696-9000, 212-685-4540 fax, Marcel Dekker 
AG, Hutgasse 4, Postfach 812, CH-4001 Basel, Switzedand (Orders: 41-61- 
261-8482, 41-61-261-8896 fax) $225.00, 876 pp., b/w photos, 7" x 10". 

^ORDs: Transgenic plants, crops, transgenic techniques, agriculture, molecular biology, food plant 
^chnology, genetics, social, political, legal aspects of genetic plant manipulation. 

V Waisel, Amram Eshel, and Uzi Kafkafi (eds.). 2002. Plant Roots. The Hid- 
den Half. Third Edition. (ISBN 0-8247-0631-5, hbk.). Marcel Dekker, Inc., 
270 Madison Avenue, New York, NY 10016, U.S.A. (Orders: 212-696-9000, 
212-685-4540 fax, wwwdeekercom). Marcel Dekker AG, Hutgasse 4, Postfach 
812, CH-4001 Basel, Switzerland (Orders: 41-61-261-8482, 41-61-261-8896 
fax) $250.00, 1120 pp., color and b/w photos, line drawings, 8 1/2" x 11". 

RE O. Barel, Marc Poaye, and Howard I. Maibach (eds). Handbook of Cosmetic 
Science and Technology. (ISBN 0-8247-02924, hbk.). Marcel Dekker, Inc., 
270 Madison Avenue, New York, NY 10016, U.S.A. (Orders: 212-696-9000, 
212-685-4540 fax, wwwdeekercom). Marcel Dekker AG, Hutgasse 4, Postfach 
812, CH-4001 Basel, Switzerland (Orders: 41-61-261-8482, 41-61-261-8896 
fax) $235.00, 886 pp., b/w photos, and other figures, 7" x 10". 

Voriin: Cosmetics, cosmetic mgredients, finished products, target organs, delivery systems, cur- 

■get Organs for Cosmetic Products; Part 3) Safety Considerations; Part 4) Vehicles of Cosmetic 
icts; Part 5) Cosmetic Ingredients; Part 6) Cosmetic Products; Part 7) Legislation and Regula- 
Worldwide; Part 8) Testing Cosmetic Products; and Part 9) Cosmetic Claims. Contributions 
some 92 expert scientists from academic dermatology and dermato-cosmetics, the cosmetics 

lAMMAD Pessarakli (ed). Handbook of Plant and Crop Physiology. (ISBN 0- 
8247-0546-7, hbk.). Marcel Dekker, Inc., 270 Madison Avenue, New York, 
NY 10016, U.S.A. (Orders: 212-696-9000, 212-685-4540 fax, 
Marcel Dekker AG, Hutgasse 4, Postfach 812, CH-4001 Basel, Switzerland 
(Orders: 41-61-261-8482, 41-61-261-8896 fax) $225.00, 973 pp., b/w pho- 
tos, graphs, and other figures, 7" x 10". 

th regulators. 

5 physiology. Part 1) Plants/ 

This is a comprehe 

nsive, up-to-d; 

ate refere 


on plani 

Crops Growth Response; 


;ntal Fad 


and Clii 

Plant/Crop Growth and 


d Stages; 


t 3) Cell 

Crop Physiology; Part 4) 

Plant/Crop PI 



i Physio 

tion Processes; Part 5) Pk 

mt Growth Re 




hibitors) and Plant Gem 

;s; Part 6) Ph> 



Drought, and Other Envi 

ronmental Stn 



ons Par 




iK Pan 


i-RiD Blunt (Introduction by William T. Stearn). 2001. Linnaeus: The Compleat 
Naturalist. (ISBN 0-691-09636-8, pbk.). Princeton University Press, 41 Wi 1 liam 
Street, Pinceton, NJ 08540, U.S.A. (Orders: California Princeton Fulfillment 
Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A., 609-883-1759, 
609-883-7413 fax, $35.00, 264 pp., color photos, 
7 3/4" X 9 3/4". 

Acta Botanica Hungarica. (ISSN 0236-6495) Akademiai Kiado, Budapest, H -1519 
Budapest, PO Box 245, HUNGARY (Subscriptions 36-1-464-8221, email 
kiss s@akkrt hu) $192 00 4 issues/vear 6 1/2" x 9 1/2" 

tigation of 4,000- 
,zi_ (E Turkey) 8) 



teach plant taxonomy, f, 
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Botanical Research 

Table of Conthnts 




U V 1 . 








,naphalieaf) in North America 

and Hawaii 


New COMBINATION IN Salix(Salicaceae) 


Taxonomy OF THE MOST 




MiHAi Costea and Franc 

Ois J.Tardif— 525 

Edward E.Terrell and Richard P 

L.Orzell and Edwin L. Bridges— 559 

Botanical History 




\ (Malvaceae) 

George M. Diggs, Jr. and Barney L Lipscomb — 647 

The vascular flora of the Lacuna de Alegria, a national monu; 


Demise Williams, Rudys W. Herrera, and JosE L Linares — 675 

The flora of Oh^ibbeha County, Mississippi 
Andreas Leidoif, Sidney McDAr 

iael H. MacRoberts, and Larry E. Brown— 781 
/If Biosphere Reserve, Mexico: A checklist 

L GarcIa-ArEvalo — 797 

SON— 813 

lAMES H. Peck— 833 
I Dale Thomas— 837 


\, 558, 570, 604, 620, 766, 796, 808, 812, 818, 836, 838, 844,8 

Dahlia neglecta D.E, Saar, sp. nov."-593 




NH.S.Howcroft W.Takeuchi 

Papua New Guinea Forest Service Harvard University Herbaria and Arnold Arboretun 

P.O.Box 406 c/o Papua New Guinea Forest Researcli Institute 

Rabaul, East New Britain 614 Papua New Guinea Forest Service 

PAPUA NEW GUINEA P.O.Box 3 14, Lae, Morobe Province 4 1 1 

hid gatherings from tl 

7>r;U^>K^mge(cfcUT. l994^lC3V-tf'^-x3>"f>^-:^->3:^JHCj; 
Dendrobium archlpelagense t D. vexillarius J.J. Sm. van hansmeyerense 2O 

(DWim<Dm(Dmm\zmi.xtimtmm^m\ m-Dt-xD^^^i^mzmuxtm^t 


Originally a part of the German Territory of New Guinea, tlie Bismarck Archi- 
pelago consists primarily of New Britain, New Ireland, and Manus. These islands 
are now separate provinces of the sovereign state of Papua New Guinea (PNG). 
The Archipelago is collectively designated as the Islands Region' by national 
administrators, and also includes several minor islets scattered outboard of New 
Ireland, as for exam pie, the St. Matthias group, Lihir, and Tabar (Fig. 1). 

The Bismarck Archipelago represents one of PNG's most poorly docu- 
mented f loristic regions (Takeuchi & Golman 2001). It is unfortunately the most 
i ntcnsively logged territory in the country: 50% of the archipelagic land area is 
currently under timber concessional development (ibid.). Because of the pau- 
city of biodiversity information, the forested tracts in southern New Ireland are 
ranked among PNG's 16 terrestrial unknowns and are also included in the country 
portfolio of high-priority sites for conservation action (Johns 1993: 25; Sekhran 

[n order to assess the status of its environments, from January 14 to Febru- 
16, 1994, a 13-member team conducted an elevation-sequenced survey in 
Weitin Valley and Hans Meyer Range of southern New Ireland. Although 
ntly pubhshed (Beehler & Alonso 
:parate checklist (Howcrof 1 1994). 
: 1994 survey are given further 
.s within the Archipelago have 

the survey's collective findings have beei 
2001), the orchid vouchers were treated : 
In the following discussion, the orchids from 
consideration. Occurrence notes from other a 
also been added when appropriate. 

Dendrobium vexillarius J.J. Sm. var hansmeyerense Howcrof t & Takeuchi, 
nOV (Fig. 2). Type: PAPUA NEW GUINEA. NEW IRELAND Province: Hans Meyer R; 
Angil Mountain, expedition camp 4 in pristine subcloud forest, 4^5.2' S. 15256.8' E, 180C 

Distribution and ecology-Known thus far only from the type locality m mossy 
montane forest at 1800 m. 

Etymology-Jhe new variety is named after the type locality 
Dendrobium vexillarius was previously represented by six varieties sepa- 
rated primarily by the color of the perianth (Reeve & Woods 1989: 250). A promi- 
nent abaxial crest on the lateral sepals distinguishes var hansmeyerense from 
other infraspecific taxa. 

The new variety was collected earlier during the 1975 Kew expedition to 
New Ireland (i.e., Sands et al 1985, 2349, 2374, 2399). Reeve and Woods (1990) 
had tentatively assigned these specimens to variety uncinatum. However that 
variety is not prominently crested on the abaxial side of the lateral sepals as 
with var hansmeyerense. Keels are sometimes present among the varieties of D. 

Fig. 2. Dendrobium vexillarius }.i.Sm. var. hansmeyerense Howcroft & Takeuchi. A. Habit. B. Flower, side view. C 
from below. D. Flower, from above. E. Labellum, midsection to apex. F-G. Column apex with and (G) without a 
Anther, front view. I. Anther with pollen, oblique rear (or anterior) view. J. Ovary in cross-section. Scale bars: A, 
E, 2 cm; F-6, 3mm; H-l (enlarged 30x); J, 3mm. Drawn from the type by N.H.S. Howcroft. 

with a prominent abaxi; 
Reeve & Woods, with a 

al keel IS D.brevi 
n otherwise ver 

y differently shaped ovary in cross-sec- 

Dendrobiiini aicliipcla. 

.„sH,.. ,„r 

V Tikcuchi, sp nov (Fig. 3) T>pr PAPUA 

Tufted cpiph\K Rhizomes shoi rsmih.imil.s u ill c base fusifoim m 

the low e I pail di^talK mou sluuiu sun. mukh 1..11 \\ ith pcisistcnt nodal 
sheaths shorter than the uiiei nodes leak t()50b\ lO-iOcm Lcavcsdwtiehous 
lanceolate ovate, 7 5-Q 5 bv 2 0-3 cm as\ mmcti icalK emaigmarc adaxialh 

florescence laccmosc, one or sevcial pci pscudobulb m^crted at the apex or 
shghtly below, erect, rachis to 40 cm by 2-3 mm; peduncle to 9 cm long, pedun- 
cular and floral bracts tubular; pedicel glabrous, ca. 2.2 cm long. Flowers 5-10 
per raceme, usually wide-opening, 4.0-5.5 by 3.5 cm, pale green to yellow-green. 
Dorsal sepal recurved, twisted through one turn, oblong-lanceolate, 2.0-2.6 by 
0.6-0.7 cm, acuminate. Lateral sepals recurved, oblong-lanceolate, 2.5-3.3 by 0.9 
cm at the base, apex acute or acuminate. Mentum narrowly conical, 0.9-1.0 cm 
long, apex straight or slightly decurved. Petals linear, acute, 3.0-4.5 by 0.25-0.4 
cm, twisted through one or two turns, spreading. Labellum porrect, 3-lobed, 2.0- 
2.3 by 1,0-1.5 cm, similar in color to sepals and petals but with some purple 
venation; lateral lobes oblong-elliptic, 1.15 by 1.5 cm, over-lapping base of the 
midlobe,apex rounded; midlobc ovate, 1.0 by 1.0 cm, acute to apiculate, not re- 
curved; callus of 5 keels, the lateral ones not extending past the base of the 
midlobe, the median keel extending to the apex of the midlobe, slightly raised, 
not lamellate or crested. Column 0.5 cm long, apex laterally bilobed, foot at 10° 
to column. Ovary glabrous, 0.9 cm long. 

Distribution and eco log}/.— Widely distributed throughout the Bismarck 
Archipelago, frequently on trees near the coast but also from inland areas. The 
type locality is lowland rainforest at 240 m. 

The new species seems to flower mainly during the first half of each year 
In addition to the t\'pc specimen (I lowering m Jan j. cultivated plants have pro- 
duced flowers continuously from Mav ro Jul. Scecfs from the previous year's set, 
are released suuultaneously with the productit>n o\ new flowers. 

Etymology-The epithet reflects the distributional restriction to the Bis- 
marck Archipelago. 

Within section Spatulata Lindl., the novelty's closest relative appears to be 

r 1 


Q A \\ 

/ \ 
/ / 

V ]■ 1 \ / 


,r^ / 

4 n,n, \ 


V V 


V / 


10 mm 


,iew. Scale bari:J-E,10mm;F,4mm;G-K 

Tin apex with stigma,frombelow.G. Anther 
2 mm. Drawn from the type by N.H.S.Howc 

lateral view 


Dcmhohium antcnnatum Lmdl. In his treatment of this section, Cribb (1986; 
618) recognized several species groups, among them the ante nnatum group with 
9 species. The new species was formerly placed in D. strepsiceros].]. 5m. by Cribb 
(ibid.: 637), O'Byrne (1994: 236), and Lavarack et al. (2000: 262). 

Smith's illustration of D. strepsiceros (1913: Fig. 4, Bl, B2) shows that its la- 
bellum differs from the corresponding characters in the new species. In D. 
strepsiceros the labellum midlobe is larger than the lateral lobes, and the apices 
of the latter do not overlap the base of the midlobe as it does in live specimens 
of D. archipelagense (Fig. 3). This distinction can be lost in the preparation of 
dried specimens so the relationship between the lobes is best seen with spirit 
material. Apart from the difference m shape and extension of the lateral lobes, 
D. archipelagense has a shorter labellum compared to D. strepsiceros and is more 

Compared to its closest congeners, the lateral lobes of the new species are 
proportionately larger in relation to the length of the labellum. In D. 
archipelagense, the lateral lobe is 70-80% of the labellum length, in D. 
antennatum 55-70%, and m D. strepsiceros 50-58% (Fig. 4). 

There are five ridges on the callus of the labellum in all of the preceding 
species. The mid-callus ridge for D. archipelagense extends towards the apex of 
the midlobe, but is not raised to a triangular lamella as described for D. 
strepsiceros. This character may have been overlooked by other authors. 

The new species has long petals which make the flower appear larger than 
typical D. antennatum. As many as ten flowers can be produced by one inflo- 
rescence. The color of the perianth is initially light green but turns yellow-green 
with age. In most individuals, the ridges of the labellum are bordered with vio- 
let, and there are also violet reticulations up to the median, and sometimes the 
distal portion of the midlobe. The perianth is persistent on the fruit, a charac- 
teristic shared by other species in the antennatum group. 

No significant color differences have been seen between specimens from 
West New Britain, East New Britain, and New Ireland. However in terms of 
flower size, the type collection represents one of the largest and most attractive 
forms seen by the senior author. 

Arachnis beccarii Reichb. f . van imthurnii (Rolfe) Tan, Selbyana 1:1-15. 1975; 1:365- 
373. 1976. (Fig. 5). 

uchi&J.Wiakabu 9968 (LAE). 

genus Arachnis is represented by two species 

eccarii occurs in the Bismarck Archipelago. 


iiik ^m 


O '<^ ^t 

Guadalcanal, Malaita, and Santa Isabel (Lewis & Cribb 1991). It was also seen 
by the senior author on the Toriu River side of the Kanak Range in East New 
Britain. The New Ireland specimen is a distributional record for the island. 

Ahhough var. imthurnii is reported as having an entirely white perianth 
(ibid.), the New Ireland provenance is marked by large brown spots. Cultivated 
plants in the National Botanical Gardens at Lae have also produced flowers with 
color markings different from mainland and archipelagic provenances. The dis- 
tinctions can be used to justify eventu; 

PicMousl) known onl) hom Indonesian Papua (van Royen 1983). The collec- 
tion liom New heland repiescnts a significant eastward extension of the spe- 
cies range. Coryhas epiphytitus is still unrecorded from the PNG mainland. 
Dendrobium gnomus Ames, J. Arnold Arbor. 14:106-107. 1933. (Fig. 7). 

Specimen examined: PAPUA NEW GUINEA. New Ireland Province: Hans Meyer Range, ridge adja- 
cent to the Weitm River, expedition camp 3, mossy montane forest, 4'27.205' S, 15256.489' E, 1175 m, 
27 Jan 1994 (IT spirit), W Takcuchi &J. Wiakahu 9597 (LAE). 

This species was initially mistaken for Dendrobium erosum (Bl.) Lindl. because 
of the spathulate shape of the labellum and its finely erose margins (Fig. 7F). 
However the leaf apex is unequally bilobed as with D.gnomus (Fig. 7 A), whereas 
in D. erosum the apex is acute. The cucullate labellum apex on the New Ireland 
specimen is probably an artefact of drying. The shape of the sepals and petals, 
and the mentum being appressed to the ovary and pedicel rather than diverg- 
ing, are also consistent with the redetermination (Figs. 7B-E). 

The species had not been previously recorded for New Ireland. 

Specimen examined: PAPUA NEW GUINEA. New Ireland Province: Hans Meyer Ri 
tain, expedition camp 4, pristine subcloud forest dominated by bryophytic and fei 

Previously known with certainty only from mainland New Gu 

Ireland specimen represents a distributional record. 

Phaius amboinensis Bl., Mus. Bot. Lugd.-Bat. 2T80. 1856. (Fig. 9). 



1^ '<'^V: ; 

5. Corybas epiphyticus (J.J. Sm.) Schltr. A. Flower and leaf, side view. B. Dorsal sepal. C. Labellum, frontal vi 
ed and showing two lateral spurs. Scale bars: A-D, 1 cm; E-F, 5 mm. Drawn from W. Takeuchi&J. Wiakabu 9- 

L Leaf. B. Flower, side view.C. Dorsal sepal. D. Petal. E. Lateral sepal. F. I 
1, oblique side view. H. Column, ovary,and spur. I. Anther from front. J. Anther from side. Scale bars: A 
n; l-J, 2 mm. Drawn from IV. Takeuchi&J. Wiakabu 9597 by N.H.S. Howcroft. 


A widespread orchid found on many islands in the Bismarck Archipelago, but 
not previously recorded from New Ireland. The species distribution includes 
Vanuatu, Fiji, Samoa, and Tahiti in the east. Also occurring in Java, the southern 
Philippines (Sulu Archipelago), Sulawesi, and the Moluccas. 

Phreatia loriae Schltr., Feddes Repert. Spec. Nov. Regni Veg. 3:318. 1907. 

rivers, expedition camp 2, lowland rainforest, 450.210' S, 152^6.242' E, 240 m, 21 Jan 1994 (fl, spirit), 

W. Takeuchi &j. Wiakahu 9885 A (LAE). 

The species was previously known only from the southern PNG mainland. 

:. &J.J. Sm., Acta Fauna Fl. Univ Bucur. ser. 2, 

The labellum of this specimen compares well with the illustration by Ha 
(1977). The new occurrence represents a significant disjunction from the pre^ 
ously known stations in Vanuatu and New Caledonia. 


ain, expedition c 

amp 4. prist 

me su 


)ud tore 










a member of: 

'rachyostek, within which 
3 species have been described for New Guinea. Of these, only B. pachyglossum 
has been recorded from the Bismarck Archipelago. The species is also known 
from the Solomon Islands (Lewis & Cribb 1991). 

Possibly synonymous with the later name C homochroma (J.J. Sm.) Schltr. 
Calanthe triplicata (Willemet) Ames, Philip. J. Sci. Bot. 2:325. 1907. (Fig. 13). 

rivers, expedition camp 2. lowland rainforest, 4°30.210' S, 15256.242' E, 240 m. Jan 1994 (fl, spirit), 

Takeuchi &j. Wiakahu 9976 (A, K, l, LAE). 

Widespread in New Guinea, the Bismarck Archipelago, and Southeast Asia. 

face of the leaves is not verrucose, as in most mainland provenances. 
Schltr. ssp. masarangense, Repert. Spec. Nov Regni 

(Fig. 15) 

Veg. 10:78. 1911. (Fig. 

26 Jan 1994 (fl, spirit), W. Takeuchi &J. Wiakabu 9534 (LAE). 

The white-flowered ssp. masarangense, with a light yellow-tipped labellum, is 
widespread and common on high islands. Lewis and Cribb (1991: 188) list its 
distribution as Sulawesi to New Guinea, New Britain, Bougainville, Guadalcanal, 
Vanuatu, Fiji, and New Caledonia. The senior author has examined this sub- 
species insitu from the Lelet plateau of New Ireland. However much of the habi- 
tat there has been recently destroyed by agricultural development and the or- 
chid has now disappeared from that locality. 

Dendrobium rhodostictum F. Muell. & KraenzL, Oest. Bot. Zeit. 44:300. 1894. 
(Fig. 16) 

cent to the Weitin River, expedition camp 3, mossy montane forest, 4^7.205' S, 15256.489' E, 1175 m, 
25 Jan 1994 (fl, spirit), W. Takeuchi &]. Wiakabu 9514 (LAE); junction of Niagara and Weitin rivers, 
expedition camp 2, lowland rainforest, 450.210' S, 15256.242' E, 240 m.Jan 1994 (fl), W. Takeuchi & 

Previously recorded from New Ireland. The pseudobulbs of these specimens 
are less clavate than other collections from New Britain, Bougainville, and the 
mainland, but the perianth segments and callus agree perfectly in shape and 

Goodyera rubicunda (Bl.) Lindl., Bot. Reg. 25: 61, misc. 92. 1839. (Fig. 17). Neottia 

rivers, expedition camp 2, lowland rainforest, 4°30.210' S, 15256.242' E, 240-300 m, 13 Feb 1994 (fl, 

spirit), W. Takeuchi &j. Wiakabu 9808 (A, K, L, LAE). 

In the broad sense, Goodyera rubicunda is distributed throughout Malesia, from 

peninsular Malaysia to Samoa. 

Spathoglottis plicata Bl, Bijdr Fl. Ned. Ind. 5: 401. 1825. (Fig. 18). 

H>' 'V V'r 

Fig. ^6.Dendroblum rbodostictum F Muell & Kraenzl A. Flowering habit B. D 

W Takeuchi &]. Wiakabu 9972 (LAE). 

The New Ireland specimen undoubtedly belongs to the 5. plicata complex. How- 
ever the ligulate lateral lobes on the labellum, with virtually no dilation of the 
apices, indicate that this is not subspecies plicata. The lack of a distinctly arched 
column is suggestive of cleistogamy. 

Spathoglottis taxa are distinguished primarily by the labellum, especially 
by the shape and angle of the lateral lobes, shape/size of the callus and its 
vestiture, and by claw length and width. Plant habit and morphology is of value 
in defining sections and subspecies (Howcroft 1986). Research by the senior 
author indicates that six subspecies of 5. plicata are present in the New Guinea 
region, of which four have been recorded from the Bismarck Archipelago. A 
formal revision of Malesian Spathoglottis will appear in a future pubhcation by 

Flowers of Dendrohium antennatum were provided by Wolfgang Bandisch, Gen- 
eral Manager of the National Capital Botanical Gardens (NCBG), and by Judith 
Raka, NCBG Administration Manageress and Scientific Officer. Specimens rep- 
resenting New Ireland provenances were received from Oscar Melepia, of 
Gaulim in East New Britain, and from Steven Kami of Port Moresby 

The 1994 New Ireland expedition was funded by Conservation Interna- 
tional and the U.S. Agency for International Development (through the 
Biodiversity Support Program). Bruce Beehler (Conservation International) and 
the Papua New Guinea Department of Environment and Conservation were 
the principal planners and organizers for the expedition. Hitofumi Abe (Eco- 
system Research Group, University of Western Australia) provided the Japa- 
nese translation. Guy Nesom (Botanical Research Institute of Texas) wrote the 
Latin diagnosis for Dendrohium archipelagense, and JohnJ. Pipoly III (Fairchild 
Tropical Garden) wrote the Latin diagnosis for D. vexillarius var hansmeyerense. 
Reviewers Paul Omerod and Andre Schuiteman made many helpful comments 
on the manuscript. 

ifR, B. and L.E.ALONSo(eds). 2001,Sc 

)uthern New Ireland, Papua Nev 

odiversity assessment. RAP Bulletin of 

Biological Assessment 21, Conserv 

Jtional, Washington, DC. 

,, P.J. 1 983. A revision of Dendrohium se 

'Ct. Latouria. Kew Bull. 38:229-306. 

, P.J. 1 986. A revision of Dendrohium se 

"Ct, Spatulota. Kew Bull. 41 :61 5-69: 

, P.J. and C.Z. Tang. 1982. Spafhog/off/s 

in Australia and the Pacific Islanc 

fiG^&.Spathoglottisplkata Bl. A. Mature bud. B. Dorsal sepal. C. Lateral sepal. D. Petal. E. Column with labellum,side 
view.FAabellum,fromabove.G-H.Bilobedcallus.Scale bars A-F,1cm,G-H,5 mm Drawn from IV Meuf/i/SXlVMofcu 

Halle, N. 1 977. Flore de la Nouvelle-Caledonie et Dependances 8; Orchidacees, Museum 
National d'Histoire Naturelle, Paris. 


Guinea Forest Research Institute. 
Johns, R.J. 1 993. Biodiversity and conservation of the native flora of Papua New Guinea. In: 

B. Beehler, ed. Papua New Guinea conservation needs assessment report, vol. 2. PNG 

Dept. of Environment and Conservation, Borol<o. Pp 1 5-222. 
KoRES, P.J. 1 989. A precursory study of Fijian orchids. Allertonia 5:1-222. 
Lavarack, PS., W. Harris, and G. Stocker. 2000. Dendroblum and its relatives. Kangaroo Press, 

Lewis, B.A. and P.J.Cribb. 1 989. Orchids of Vanuatu. Royal Botanic Gardens Kew. Whitestable 

Lithe Printers Ltd. 
LEwis,B.A.and P.J.Cribb. 1 991 .Orchids of the Solomon Islands and Bougainville. Royal Botanic 

O'Byrne, R 1 994. Lowland orchids of Papua New Guinea. SNP Publishers Pty Ltd Singapore. 
REEVE,T.M.and P.J.B. Woods. 1 990.A revision of Dendroblum sect.Oxyglossum (Orchidaceae). 
Notes Royal Bot. Card. Edinb. 46:1 -305. 

Reprinted from: R van Royen.1979.The Alpine flora of New Guinea.Vol.2.0rchidaceae. 
J. Cramer,Germany.Pp 51-812. 

Royen,Rvan. 1 983.The genus Corybas in its eastern areas. Phanerogamarum Monographiae 
Tomus XVI. J. Cramer, Germany. 

ScHLECHTER,R. 1911-14. Die Orchidaceen von Deutsch-Neu-Guinea. Feddes Repert. Spec. 
Nov. Regni Veg. Beih. 1 :1 -1 079. English translation 1 982, D.R Blaxell, ed.The Australian 
Orchid Foundation, Melbourne. 

SEKHRAN,N.and S. Miller (eds). 1995. Papua New Guinea country study on biological diver- 
sity. Colorcraft Ltd, Hong Kong. 

SMiTH,J.J.1913.0rchidaceae.Nova Guinea 12:1-108,t.l4,39. 

from contemporary surveys in Papuasia.Sida 19:445-468. 
Tan, K. 1 975. Taxonomy of Arochnis, Armodorum, Esmeralda and DImorphorchis 



Gilberto Ocampo Acosta 

Centro Regional del Bajio 
Apdo. Postal 386, C.P.61 600 
Pdtzcuaro, Michoacdn, MEXICO mx 

campanulated opercului 

Durante la preparacion del fascicule correspondiente a la familia Portulacaceae 
para la Flora del Bajio y de regiones adyacentes, se encontraron algunos 
ejemplares de herbario del genero Portulaca, los cuales fueron colectados en la 
parte norte del estado de Michoacan (Mexico) y que corresponden a una entidad 
que se puede ubicar en la subseccion Conocarpac D. Legrand (subgenero Portu- 
laca, seccion Catodasis D. Legrand), cuyo unico miembro reportado para Mexico 
es P mcxicana P. Wilson. Estos ejemplares presentan caracteristicas que no 
corresponden a ninguna de las especies conocidas en la actualidad, por lo que 
se propone como: 

rzedowskiana G. Ocampo, sp. nov. (Fig. l). 

■ubra, (3-)4.5-6.5 mm longa, 4-5.5(-6.! 
onga, (3-)3.5-6(-7) mm lata; stamina ] 
3); capsula 3.5-6 mm longa, operculu: 

Planta herbacea perenne, carnosa. Raiz engrosada, suberosa, de 2-6 cm de 
largo por 0.5-3.5 cm de ancho, en raras ocasiones surgen de ella raices 
cilindraceas de hasta 15 cm de largo por 2-4 mm de ancho. Tallos varios 
surgiendo de la raiz, de 4-26 cm de largo, postrados a decumbentes. Hojas 
akernas, aplanadas a subroUizas, carnosas, pelos axilares de 3-5 mm de largo, 
peciolo de l-1.5(-2) mm de largo, lanceoladas, en ocasiones oblongo-lanceoladas 
a oblanceoladas, de 4-20 mm de largo por l-3(-4.5) mm de ancho, apice agudo. 
Flores terminales, sesiles, agregadas en cabezuelas de (2-)3-8 flores, rodeadas 
por un verticilo de (6-)8-10(-13) hojas, pelos de hasta 5 mm de largo; sepalos 
connados en su base, rojizos, ovados, de (3-)4.5-6.5 mm de largo por 4-5.5(-6.5) 
mm de ancho, desiguales, apice agudo, margen escarioso, persistentes hasta la 
madurez de la capsula; petalos de color rosa a morado, connados en su base, 
obovados, de (5.5-)6.5-ll mm de largo por (3-)3.5-6(-7) mm de ancho, obtusos 
a emarginados, la mayoria de las veces con un pequetio mucron; estambres 22- 
44, sus f ilamentos de (2.5-)3-4(-5) mm de largo, connados en su base formando 
un anillo estaminal de ±1 mm de largo, anteras de 0.6-0.8(-0.9) mm de largo; 
estilo de 4.5-6(-8) mm de largo, ramas estigmaticas (3-)4-5(-6), de 1.5-2.5(-3) 
mm de largo. Capsula sesil o sobre un pedicelo de hasta 1.5 mm de largo, la 
capsula de 3.5-6 mm de largo por 3-4.5(-5) mm de diametro, con dehiscencia 
en su tercio inferior o mas cercana a la base, operculo campanulado a semi- 
campanulado; semillas de color negro-azulado cuando maduras, brillantes, 
reniformes, de 0.6-0.7(-0.8) mm de largo, tuberculos redondeados con estelulas 
radiales en su base. 

1986, H, Diaz 2372 (CH APA, lEB). Municipio de Charo: brecha rumbo a L; 
tos 2m (lEB); ±1.5 km sobre el camino a Las Mesas, a un costado de la cort 

(CHAPA, EBUM, lEB, MEXU). Municipio de Morclia: Puerto de Los Copale, 
carretera a Mil Cumbres, 22 Ago 2001, G. Ocampo 1124 (lEB); alrededores 
Sep 1991, E. Garcia y E. Pe rez 3930 (lEB); San Jose Coapa, 4 Oct 1989,J.M. Esc 

Portulaca rzedowskiana se ha encontrado en lugares con matorral subtropical 
perturbado o con pastizal, en terrenos pianos o con poca inclinacion, entre los 
1800 y 2200 m de altitud (5920-7230 ft). Su abundancia local oscila entre escasa 

Fi6. 1 , Portulaca rzedowskiana. A. Aspecto general de la pi 

y francamente abundante, por lo que se puede considerar que carece de 
problemas de supervivencia. Por el momento, unicamente se conoce de la parte 
norte de Michoacan, pero es de esperarse que esta especie se encuentre en zonas 
colindantes de los estados de Jalisco y Guanajuato. 

Las especies de la subseccion Conocarpae D. Legrand, muestran capsulas 
con dehiscencia en su tercio inferior, el operculo es campanulado o subconoideo 
(raramente casi hemiesferico) y sus hojas son rollizas a aplanadas, dispuestas 
en grupos de 6 a 14 en las mf lorescencias (Legrand 1962). P. rzedowskiana tiene 
mayor afinidad con P mexicana P. Wilson, P minensis D. Legrand, P perennis R. 
E. Fr y con P wedermannii Poelln., de las cuales se diferencia por una serie de 
combinacion de caracteres (Cuadro 1), siendo los principales el tamano mas 

Raiz Engrosada.suberosa Engrosac 

Densidad de Regularmente Regularn 

veces oblongo-lanceolada 

oblongo-lanceoladas a 

Tamano 4-20 mm de largo por 4-10 mm de largo 6-12 mm de largo p 

Color Rosa a 



Color Negro-a 

grande que pueden alcanzar sus tallos, asl como las mayores dimensiones de 
sus hojas y de sus capsulas (P rzedowskiana presenta las capsulas mas grandes 
de la subseccion (D. Legrand, op. cit.), las cuales se conocian de hasta 5 mm de 
largo), asi como por el color negro-azulado de sus semillas. En particular, la 
nueva entidad se diferencia de P. mexicana y P. perennis por sus hojas aplanadas, 
por sus petalos de hasta 1.1 cm de largo, por presentar mayor numero de 
estambres, asi como por tener mayores dimensiones en el tamafio del estilo y el 
tamafio de la semilla; de P minensis y P wedermannii, especies de distribucion 
sudamericana, se diferencia principalmente por la abundancia regular de la 
pilosidad axilar y por el menor numero en las ramas del estilo, 

Martin de Sesse y Jose Mariano Mocifio, en su obra Flora Mexicana, registran 
a Portulaca meridiana L.f. (p. 132, 1891-1897) de las cercanias de Puruandiro 
(Michoacan), de donde tambien se ha colectado P. rzedowskiana. En la 
actualidad, se considera que P meridiana L.f. es sinonimo de P quadrifida L., 
especie distribuida en Africa, Asia, Oceania y las Antillas. Por desgracia, de la 
excursion efectuada por los autores mencionados, no se conocen ejemplares en 
el herbario del Real Jardin Botanico de Madrid (MA) (McVaugh, p. 446, 2000) 
ni dibujos en la Coleccion Torner de la expedicion de Sesse y Mocino (Hunt In- 
stitute for Botanical Documentation, Pittsburgh, E.U.A.) asociados con la especie 
aludida. Al revisar la descripcion que Sesse y Mocifio hacen del material 
observado, esta parece coincidir con las caracteristicas de P rzedowskiana. La 
descripcion de P meridiana que se encuentra en la Flora Mexicana se transcribe 

Portulaca foliis lineari-subulatis, carnosis, axillis pilosis; floribus 
polyandris, terminalibus, sessilibus. El. Mex. 

Portulaca foliis ellipticis, carnosis, planis, articulis pilosis, floribus 
sessilibus, terminalibus. Herba dodrantalis, caule tereti, glabro, succulento. Fo- 
lia alterna, lineari-subulata, carnosa, sessilia, glaberrima, pilis fasciculatis, 
utrinque axillaribus ornata. Flores rubro-purpurei, polyandri, terminales, in 
capitulum sessiles, verticillum foliorum suffulti. Capsulae ovatae, circumcisae. 

Habitat prope Oppidium Puruandirum. Floret lulio. 

Portulaca meridiana es una planta anual, con hojas opuestas, aplanadas, 
con flores rodeadas por un verticilo de 4 hojas, con petalos amarillos y con el 
operculo de la capsula de forma campanulado-tubulosa. La disposicion opuesta 
de las hojas, entre otras caracteristicas, la ubican en el subgenero Enantiophylla 
D. Legrand, por lo que la asignacion del nombre de P meridiana que hicieron 
Sesse y Mocino a los ejemplares que observaron cerca de Puruandiro es 
incorrecta. La descripcion en Flora Mexicana concuerda con las caracteristicas 
de la nueva entidad, por lo que muy probablemente se trate de la misma unidad 

Etimologia.-¥J epiteto especifico esta dedicado a la Maestra Graciela 

Deseo expresar mi agradecimiento a Heike Betz (Field Museum) por su 
apreciable ayuda en la consecucion de bibliografia. De igual forma, deseo 
agradecer a Patricia Mayoral, a Emmanuel Perez, a Sergio Zamudio (todos ellos 
del Institute de Ecologia, A.C., Centro Regional del Bajio), asi como a James F. 
Matthews (University of North Carolina at Charlotte) por la revision critica del 

nericanas de Portulaca.Anak 

H, R. 2000. Botanical results of the Sesse & Mociho expedition (1 787-1 803). \ 
; to relevant scientific names of plants. Hunt Institute for Botanical Docume 

Book Notice 

RETT E. Crow and C. Barre Hellqulst. 2000. Aquatic and Wetland Plants of 
Northeastern North America: A Revised and Enlarged Edition of Norman 
C. Fassett's A Manual of Aquatic Plants, Volume 1. Pteridophytes, Gym- 
nosperms, and Angiosperms: Dicotyledons. (ISBN 0-299-16330-X, hbk.). 
$90.00, 480 pp, 338 line illus., 1 map, 8 1/2" x 11". 

RETT E. Crow and C. Barre Hellqulst. 2000. Aquatic and Wetland Plants of 
Northeastern North America: A Revised and Enlarged Edition of Norman 
C. Fassett's A Manual of Aquatic Plants, Volume 2. Angiosperms: Mono- 
cotyledons. (ISBN 0-299-16280-X, hbk.). $90.00, 400 pp, 268 line illus., 1 
map, 8 1/2" x 11". 

The University of Wisconsm Press, 1930 Monroe Street, 3rd Floor, Madi- 
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versity of Wisconsin Press, c/o Chicago Distribution Center, 11030 S. Lan- 
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Greg Starr Jose A.Villarreal Q. 

Tucson, AZ 85745, U.S 







I especi 

^dc maguey. Agave ova 
n es propuesta como nv 
relacion con A. havardi 
la clave pared ymodif 

tifolui Star 
leva. Es ub 
ana TreL y 


la Sierra de Lampaz, 
.Parr^anae desubg. 
;er, Se presenta una i 



:ies, Agave 


[t belongs i 


le Sierra deLampazosm 
Parr^anae of the subgen 

La colecta de material proveniente del norte del estado de Nw 

proporciona informacion de una nueva especie de Agave. 

Agave ovatifolia Starr «Sa: Villarreal, sp nov (Figs. 1 y 2) Tipo MEXICO ^ 

Plantas multianuales; rosetas hemisfericas, algo compactas, 80-120 c 
diametro, 60-90 cm al to; hojas ar regladas en 8-10 hileras, 40-50 hojas por plar 
elipticas, ligeramente concavas a casi planas, 35-45 cm de largo, 20-24 cm 
ancho en la parte media, de color glauco-grisaceo, ocasionalmente con ] 
costiUas dorsales, base ensanchada, apice acummado con borde de color C2 

^ \\ 

espina terminal de cerca de 2 cm de largo, margen recto, con 15-19 pares de 
dientes ligeramente curvados, separados 20-25 mm, mas proximos cerca de la 
base; inflorescencia 3.5-4 m alto, forma eliptica a largamente ovoide, escape 
floral 9-11 cm ancho en la base, 6-8 cm en la primer ramificaclon, raquis algo 
sigmoideo, 13-16 ramillas f lorales, de 30-50 cm de largo, ubicadas en la mitad 
superior del escapo; bracteas eliptico-triangular, esparcidas, las basales 20-25 
cm largo, 8-10 cm ancho en la base, las superiores 12-15 cm largo, 7-9 cm ancho 
en la base, acuminadas con espina terminal, enteras a dentadas, delgadas 
(papiraceas), glaucas, pronto se deshidratan y cambian a color cafe-claro, la 
mayoria erectas; flores en 5-6 umbelas densas, 67-74 mm largo, verde 
amarillentas; pedicelos 4-8 mm largo, ovario cilindrico, 30-35 mm largo, cuello 
del perianto 7 mm largo, tubo del perianto 15-20 mm largo, 15 mm ancho en la 
parte media, tepalos largamente triangulares, 15-17 mm largo, 5-7 mm ancho 


2. Planas de Agave ovatifolia 

en la base, los tres mas externos con evidentes glandulas apicales, filamentos 
50-60 mm largo, ligeramente aplanados msertos en la base de los tepalos, verde 
amarillento, anteras 22-25 mm largo, verdes, estilo 8-9 cm largo; capsulas 
cilindricas 50-60 mm largo, 15-20 mm ancho, color cafe oscuro; semillas 
lacrimiformes, 5-6 mm largo, 3-4 mm ancho, color negro brillante. 

Crece en laderas y mesetas rocosas entre los 1100-1500 m, tanto en bosque 
de encino como matorral de Agave lechuguilla y arbustos espinosos. Su 
distribucion es restringida en el area antes mencionada con la posibilidad de 
que se encuentre tambien en otras sierras aledanas. Las poblaciones son 
esparcidas, donde no se observan hijuelos ni la formacion de colonias. La planta 
es reportada como buena forrajera para el ganado y las poblaciones son 
menguadas por jabalies y otra fauna silvestre. Solo se observaron plantas 
maduras y en floracion en laderas con fuerte pendiente, donde escapan a la 
depredacion. Localmente se le conoce como "noga" y se le diferencia de las 
plantas que carecen de hojas con costillas dorsales, las cuales son denominadas 
"maguey noga." 

La especie se ubica en la seccion Parry anae del trabajo de Gentry (1982) y 
al parecer esta relacionada morfologicamente con A. havardiana Trel. y A. 
parrasana Berger, de las cuales se diferencia por sus hojas proporcionalmente 

mas anchas, de color gris-azuloso y flores con tamano mtermedio, como se 
muestra en la siguiente clave modificada. Aunque las dimensiones de las hoja 
en A. havardiana y A. ovatifolia se sobreponen, la relacion entre el largo y ancho 
es diferente, la de A. havardiana es de 2.0-2.8, mientras que la de A. ovatifolia es 
de 1.5-1.8. La relacion entre largo y ancho de la hoja en A. parrasana es de 2.0- 
2.5, pero con hojas consistentemente mas cortas. Agave havardiana se distribuye 
en el suroeste de Texas y regiones adyacentes de Chihuahua y Coahuila, A 
parrasana en sierras del centre y sur de Coahuila, mientras que A. ovatifolia en 
la Sierra de Lampazos en el norte de Nuevo Leon. 

Agave ovatifolia es ubicado en la seccion Parryanae, y la clave de Gentry 
(1982:526) puede ser modificada como sigue: 

2. Rosettes globose, suckering copiously with long rhizomes forming large, spread- 
ing clones; panicles deep with 20-40 umbels; flowers 60-80 mm long 3 

2. Rosettes not globose, more openly flat-topped, with few or no suckers; panicles 

rather wide and open with 10-20 large umbels; flowers 67-90 mm long 7 

7. Leaves larger, 30-70 x 15-25 cm, long acuminate; teeth reflexed along middle 
and lower leaf margins; tepals about equaling to slightly longer than tube. Big 
Bend Region and adjacent IVlexico 8 

along margin5;tepals about twice as long as tube. Sierras of southern Coahuila. 

gray to light green, spine stouter, 3-5(-10) cm long; flowers 68-88 mm long, 
tepals 18-24 mm long; Big Bend, east-central Chihuahua and northwestern 
Coahuila, Mexico Agave havardiana p. 531 

Al parecer A. ovatifolia es la misma planta que Mrs. Anna B. Nickels colecto y 
distribuyo como A. noah Nickels a finales del siglo antepasado. La especie de 
Mrs. Nickels no fue descrita y tampoco fue apropiadamente publicada, a pesar 
que Trelease (1911) la enlista como smonimo de A. wislizeni Engelm. La 
taxonomia de A. wislizeni ha sido ampliamente discutida por Gentry (1975) y 
Ullrich (1992). El concepto de Trelease (1911) para A. wislizeni acomoda bien 

horticultor texano, colecto esta especie de Sierra de Lampazos, a principios de 
los 1990's. El epiteto especifico hace referencia a las hojas relativamente anchas 
en relacion a su largo. 

A. Encina por 
Miguel A. Can 


RY, H.S. 1 975. The case of Agave scabra and Agave wislizenii. Cact. Succ. J. (Los Angeles) 

^:1 02-1 04. 

RY, H.S. 1 982. Agaves of Continental North America. University of Arizona Press, Tuc- 


ASE,W. 191 1 [191 2]. Revision of the Agaves of the gmup Applanatae. Ann. Rep. Mis- 

)uriBot.Gard. 22:85-97. 

CH, B. 1992. On the history of Agave asperrima and A. scabra (Agavaceae) as well as 

)me taxa of the Porryanoe. Sida 1 5:241-262. 

Book Notice 
Laurence J. Dorr, Basil Stergios, Alan R. Smith, and Nidia L. Cuello A. Chief Edi- 
tor: Laurence J Dorr 2000. Catalogue of the Vascular Plants of Guaracamal 
National Park, Portuguesa and TrujiUo States, Venezuela. (ISSN 00971618, 
pbk). Contributions form the United States National Herbarium, Volume 
20:1-55. Department of Botany, National Museum of Natural History, 
Smithsonian Institution, Washington, DC 20560-0166, U.S.A. Price not given, 
155 pp, 3 figures, 7" x 10". 

applicable Relevant taxonor 

Journal Notice 
Acta Botanica Hunganca (ISSN 0236-6495) Akademiai Kiado Budapest H-1519 
Budapest PO Box 245 HUNGARY (Subsciiptions 36-1-464-8221 email 
kisss@akkithu) $19200 4 issues/yeai 61/2 x91/2 


Perennial herbs 12-17 cm high, the stems seemingly recumbent and arising from 
woody roots which exude a violet-colored dye. Leaves basal at first, but soon 
developing 20 or more overlapping cauline leaves, these gradually reduced up- 
wards and grading into the floral bracts; blades linear-lanceolate to lanceolate, 
essentially nonvenose, except for the midribs, the surfaces appressed-strigose, 
the undersurfaces having finer hairs, the upper surfaces with coarser hairs hav- 
ing attractive flattened basal cells 0.15-0.30 mm across. Inflorescence decid- 
edly scorpioid, having numerous bracteate pale yellow flowers. Sepals 5, linear, 
mostly 7-9 mm long, moderately pubescent without, glabrous within, weakly 
glandular at the throat, markedly nodular-annulate at or near the base; lobes 5, 
ovate, 1.4-1.6 mm long, 1.0-1.2 mm wide. Anthers 5, dorsifixed, ca. 1.7 mm long, 
inserted in the throat but not exceeding the lobes. Styles ca. 11 mm long, gla- 
brous, minutely bifid at apex. Nutlets 2-4, glossy-gray, not at all ornamented, 
ca. 2 mm high, 1.5 mm wide, basally attachment ca. 2 mm across. 

The relationship of this taxon among Mexican species of Lithospermum is 

moot. Assuming the plants to be consistently homostylous with weakly devel- 
oped faucal appendages, it will begrudging! y key to L. calycosum (Macbride) 
l.M.Johnst. in Johnston's (1952) treatment of Lithospermum, to which it is com- 
pared in the above diagnosis. It differs in having much larger, broader, leaves, 
noncrenulate corolla lobes, and larger (ca. 3 mm high vs 2 mm), glossy-gray 

It is a pleasure to name this species for the remarkable Hinton family, who 
first collected the taxon. When first sent to me for identification I recognized it 
as possibly new but lacked fruiting material for descriptive purposes. George 
Hinton (son of the living Jaime Hinton, and grandson of the legendary plant 
collector, George B. Hinton) kindly sent me mature seeds nine years after its 
initial collection, these obtained from garden grown plants collected at the type 
locality. I am grateful to him for such diligence. 

1 am grateful to my wife Gayle Turner for the Latin diagnosis and to Diane M. 
Ferguson and James S. Miller for helpful comments. 


Book Notice 

Stanwyn G. Shetler and Sylvia Stone Orel 2000. Annotated Checklist of the Vas- 
cular Plants of the Washington - Baltimore Area. Part I: Ferns, Fern Al- 
lies, Gymnosperms, and Dicotyledons, (no ISSN or ISBN). Department of 
Botany, National Museum of Natural History, Smithsonian Institution, 
Washington, DC 20560-0166, U.S.A. Price not given, 186 pp [duplicated], 1 
figure, 8 1/2" x 11". 

Stanwyn G. Shetler and Sylvia Stone Orli. 2000. Annotated Checklist of the Vas- 
cular Plants of the Washington - Baltimore Area. Part II: Monocotyledons. 
(no ISSN or ISBN). Department of Botany, National Museum of Natural 
History, Smithsonian Institution, Washington, DC 20560-0166, U.S.A. 95 
pp [duplicated], 8 1/2" x U". 

ginia. "Species were a 





Plant Resources Center 
The University of Texas 

S. platyphylla. 

Schistocarpha is a mostly Mexican genus of ten closely related species (sensu 
Turner 1986 and the present paper). These have been variously interpreted (e.g, 
Rydberg 1927; Robinson 1979; Turner 1996), the most recent appraisal of the 
Mexican taxa, albeit for the state of Chiapas, being that of Strother (1999). With 
the exception of 5. eupatorioides (Fenzl) O. Kuntze and S. pedicellata Klatt, 
Strother placed most of the Mexican species recognized by previous workers 
under the fabric of 5. hicolor Less. Because of Strother's treatment I have looked 
again at the Mexican taxa and have concluded that one can reasonably sink the 
widespread, highly variable 5. longiligula Rydb. into the earher S. platyphylla 
Greenm., but the remainder of Strother's dispositions under a widespread S. hi- 
color is moot. Indeed, having gone over this taxonomic landscape anew I would 
still recognize six species as occurring in Mexico, including 5. liehmannii Klatt, 
S. matudinae H. Rob., and the newly described 5. calzadana, proposed below. 
Schistocarpha calzadana B.L. Turner, sp. nov (Figs. 1, 2). Type: MEXICO. Oaxaca: 

IS of Mexican species of Schistocarpha. 


Perennial, suffruticose herbs 1.5-2.5 m high. Stems moderately to densely pi- 
lose, glabrate with age. Larger leaves 20-30 cm long, 14-16 cm wide; petioles 6- 
8 cm long, winged throughout but much tapered below into a distinct petiole; 
blades broadly subcordate, markedly serrate, their apices rounded or obtuse. 
Primary capitulescences 14-20 cm across, ca. 10 cm high; ultimate peduncles 
mostly 3-9 mm long, pubescent like the stems. Involucres 4-5 mm high and 
about as wide, having imbricate 3-4 seriate bracts, the inner bracts scarious 
with mostly rounded ciliate apices. Receptacle plane or somewhat convex, 
paleate, the pales shorter than the subtended florets. Ray florets ca. 13 in a single 
series; tubes ca. 3 mm long; ligules white, 1.5-2.5(-3.0) mm long, ca. 0.8 mm wide, 
2-4 nervate. Disk florets 30-40; corollas ca. 4 mm long, tubes ca. 2 mm long, 
glabrous or neady so, the lobes 5, ca. 0.5 mm long, hispidulous without. Achenes 
of ray and disk similar, ca. 1.5 mm long, glabrous; pappus of numerous white 
bristles 3-4 mm long. 

rest," 3500 ft, ' 

d poblac 

In the account of Schistocarpha by Robinson (1979), the present taxon will key 
to S. matudae, a species of southern Chiapas. In my treatment of the genus 
(Turner 1986), it will key to or near 5. liehmannii, to which I originally referred 
the collections from Guerrero. With the newly collected Oaxacan material in 
hand, it seems likely that the several sheets relate better to S. hicolor, the latter 
from the Gulf Coast sierras, the former from the Pacific sierras. Schistocarpha 
calzadana differs from its more eastern cohort m characters as noted above. 

It should be noted that Strothers (1999) treatment of Schistocarpha for the 
Asteraceae of Chiapas would reduce nearly all of the Mexican taxa of tf 
nus (other than S. eupatorioides and S. pedicellata) to but a single widespread S. 
hicolor, including the present novelty His comments to justify such redi. 
"I have seen intermediate specimens that link all of the named extremes.' 
exceedingly strained to me, especially since his examination of a wide ra 
collections at LL-TEX resulted in no annotations to that effect. I had no great 
difficulty in placing 'segregate' species names on the holdings at CAS, LL-TEX 
and UC, most of which served as the basis for Strother's remarks. I have " 
ever, accepted the likelihood that S. longiligula and S. platyphlla are sy 
mous, the two being largely sympatric and separated by relatively trivi 
tures (ligule length and floret number). I also freely admit that 5. matudae, S. 
liehmannii, and S. calzadana might be swept under S.platyphyUa,3iS interpreted 
here (the earliest name for the complex then being 5. liehmannii), but such ti- 
dying should take place only after more detailed field studies of the popula- 
tions concerned. Regardless, S. bicoior (with its markedly winged petioles) does 
not extend so far south in Mexico so as to include the state of Chiapas. 

Etymology— The species is named for J.T. Calzada, a Mexican botanist who 
was the second worker to garner the taxon and who participated in all subse- 
quent collections. 

I include below a revised key to the Mexican species of Schistocarpha, which 
includes the present novelty Figures two and three show the distributions of 
these taxa. The revised key and up-to-date distribution maps should prove help- 
ful to future workers interested in the group. 


inged throughout, often 

Fig. 3. Distributions of Mexican species of Sf/j/sfofor/j/io. 

. Capitulescence strict, congested, coi 

peduncles mostly l,5-2.0(-2.5) cm I 

4. Stems glabrate or nearly so at ma 

veins) glabrous, the upper surfaC' 

north-central Oaxaca and adjacer 

4. Plants without the above comb 

5. Ray florets 5-1 2; ligules mostly 

5. Ray florets 12-21; ligules most! 

^ cloud forests of 

_S.platyphylla (Fig.'^ 

S.calzadana (Fig.: 

all of the above-keyed taxa are treated i 

I am grateful to my wife, Gayle Turner, for the Latin diagnosis and to Jose Panero 
for calling my attention to the Oaxacan plants concerned. John Strother re- 
viewed an early draft of the manuscript, improving considerably its terseness 
and readibility. The study is based upon materials from the following herbaria: 
CAS, LL-TEX, UC, as follows: Schistocarpha hicolor CAS (22); LL-TEX (39); UC 
(7). Schistocarpha liehmannii CAS (4); LL-TEX (10); UC (1). Schistocarpha 
matudinae CAS (4); LL-TEX (3); UC (0). Schistocarpha pedicellata CAS (9); LL- 
TEX (15); UC (1). Schistocarpha platyphylla CAS (68); LL-TEX (54); UC (8). 


Robinson, H, 1979. A study of the genus Schistocarpha (Heliantheae: Asteraceae). 

Smithsonian Contr.Bot. 42:1 -20. 
Rydberg, P.A. 1 927. Schistocarpha, in N. Amer. Fl. 34:303-306. 
Strother, J.L 1 999. Schistocarpha. In: Flora of Chiapas, Pt. 5: Heliantheae s.l. , Calif. Acad. Sci., 

' genus Schistocarpha 



William A.Weber 


2 Rocky Mountains of northwestern Montana. It is distinguishec 
ieowehsteri Blake, by glabrous foliage, broader and shorter leaf 1 


neowehsteri Blake, por sus hojas glabras, lamina foliar m; 

Senecio spribillei W.A. Weber, sp. nov. (Fig. 
net Mountains, E face of Snowshoe Peak, ca. 150 1 
on sparsely vegetated alpine rock ledges, with C 
alt., 25 Aug 2001, TobySpribille&M.An'idson . 

longepetiolatis 0.5 dm longis, folia basalibus caulis f lor 
folia caulina 4-5, laminibus late triangulo-ovatis vel s 
truncatis vel ad basi brevissimo-cuneatis, grosse spinulo 

Perennial from a well-developed, often branched caudex, up to 2 dm tall, to- 
tally glabrous; first year shoots 0.5 dm long, consisting of a few basal leaves on 
long petioles; flowering stem (second season) with withered basal leaves, cauline 
leaves with lamina broadly triangular-ovate to subrotund, 3.0-4.0 x 2.5-3.0 cm, 
truncate or very shortly cuneate at the base, coarsely spinulose-dentate, peti- 
oles scarcely winged, 4-5 cm long, purple; cauline leaves 4-5, similar to the basal 
leaves; heads 4-6, nodding, the lower on long, erect peduncles; involucre 8-10 
mm high, the disk 5-7 mm diam, with a single row of phyllaries and a few 
minute basal bracteoles; phyllaries lance-oblong, obtuse or broadly acute, with 
broad hyaline margins, glabrous; ray flowers few, little exceeding the involucre, 
the lamina 7x2 mm, the veins 4-5, simple, the tube 4 mm long; disk flowers 
with limb 4 mm, tube 4 mm long; pappi 3 mm long; cypselae linear, strongly 
ribbed, 3 mm long. 

This new species is most closely allied with S. neowehsteri Blake (S. wehsteri 
Greenman, non Hook.), an endemic of the Olympic Mountains of Washington 
(Table 1). It appears to be a narrow endemic of northwestern Montana. 

The ray flowers in the new species are so short and so narrow that the col- 
lector believed that the plant was rayless. The structural anatomy of the ray 
flowers differs from that of Senecio neowehsteri. In S. sprihillei the nerves of 
the ray flowers are simple, unbranched; in 5. neowehsteri each nerve is forked 
near the base, the branches continuing closely parallel distally. Baaghoe (1977, 
1978) published on the taxonomic application of ligule micro-characters in the 

Senecio neowebsteri 

Foliage initially floccose,tl 

Ligules ot ray tlowers /-I 
hardly longer than the ir 

Asteraceae but made no mention of venation or possible differences involving 
venation. Her work dealt with light and electron microscope observations, 
mostly on epidermal features. The existence of branched and non-branched 
nerves in Senecio, s. lat. deserves attention. 

Distribution.— Known only from the type collection. According to the col- 
lector, the locality is likely a nunatak north of the southernmost limits of the 
Pleistocene ice sheets. 

Etymology— Senecio sprihillei is dedicated to the young taxonomist and 
phytosociologist, Toby Spribille, with whom I have been associated for several 
years. His keen eye and ability to deal with the lichen and bryophyte as well as 
the phanerogamic flora, his knowledge of phytosociology, and his fine collec- 
tions, marks him as an important figure in the recent history of the Rocky Moun- 
tain flora in Montana. 

?, and Tageteae. Bot.Tidss^ 

Book Notices 
LES M. Allen, Dawn Allen Newman, and Harry H. Winters, 2002. Trees, Shrubs, 
and Woody Vines of Louisiana. (ISBN 0-9718625-0-8, pbk). Allen's Native 
Ventures, LLC, 5070 Hwy 399, Pitkin, LA 70656, U.S.A. (Orders: 337-328- 
2252, Price not given, 333 pp, numerous b/w line draw- 
ings, 3 maps, 5 1/2" x 8 1/2". 

throughout the state (FACU) as an ornamental with some 

recent interest in its wood. Th 

records include 14 parishes" (Paulownia tomentosa). Apart 

from the enumeration of whic 

species occur in the state and their reproductive status, perh 

laps the most useful feature of tl 

the line drawing for each, drawn from a range of pre 

viously published sources. Iden 

keys are not provided but 8 'tables' in the introductory mat 

eriallist genera with distinctly 

ters such as strong odor, compound leaves, opposite or wl 

lorled leaves, etc. This compila 

serve good use as a reference in company with an identifi 

cation manual and range map: 

Nesom,Botanical Research Institute of Texas. 509 Pecan Stre. 


1 Graves, Photos by Wyman Meinzer. 2002. Texas Rivers. ISBN 1-885696- 
38-8, hbk.) University of Texas Press, PO. Box 7819, Austm, TX 78713-7819, 
U.S.A. (Orders 800252-3206 512-471-4032, 800-687-6046 fax)) $39.95, 144 
pp.80 color photos, 11" x U". 
as Rivers were only a picture book it would be an outstanding work. The photographs by Wyman 
ler are breathtakingly beautiful. Panoramas and close-ups alike are full of radiant color and 


Guy LNesom 


:al Research Institute of Texas 







ium primarily on the 

ncy for stolon produce 


d, non-mucilaginous 
ca: E. sphaericus (nat, 


ornia and Oregon), E. 

iia and Oregon), and Einvoluc-ra(usG 

apparent waif m Cali- 

ts, the 1 


)ecies in North A 




1 ecological sum 




e en base a su ten 


:onpapUas epidermic 



en Norte 



a California y Oregon), 

Euchiton is treated apart from Gnaphalium primarily on the basis of its ten- 
dency for stolon production and cypselar surfaces with paired, non-mucilagi- 
nous epidermal papillae (Anderberg 1991). The plants are mostly perennial and 
produce heads in terminal, capitate clusters immediately subtended by a whorl 
of leafy bracts. The distinctiveness of the group was emphasized by Drury (1972) 
and Holub (1974); nomenclatural transfers to Euchiton were made by Holub 
(1974) and completed by Anderberg (1991) and Ward and Breitwieser (1998). 
The species number about 22 (Drury 1972; Anderberg 1991), including recent 
additions (Buchanan 1999, Walsh 1999) or 20-30 (Walsh 1993). They are native 
to Australia and New Zealand and probably to New Guinea and eastern Asia; 
some species have weedy tendencies and have become widely naturalized. 

The following synopsis of Euchiton in North America and Hawaii is based 
on surveys of collections at ARIZ, BRIT-SMU, CAS-DS, GH, HSC, MO, NCU, TEX- 
LL, UC-JEPS, and WIS. 
Euchiton Cass, in E Cuvier, Diet. Sci. Nat. ed. 2, 56:214. 1828. Type species: Euchiton 

Plants perennial or (less commonly) annual, herbaceous, stoloniferous in most 
species (taprooted m 1), white-tomentose, eglandular. Leaves basal and cauline, 
sometimes in a rosette, lanceolate to linear, entire, sessile or petiolate, bicolored, 
usually with a close tomentum at least on the abaxial surfaces. Capitulescences: 
heads usually in a terminal cluster subtended by a whori of leafy bracts, some- 
times with axillary clusters below. Capitula narrowly campanulate to short- 
cylindric; phyllaries chartaceous, transparent, mner with larger stereomes. Pis- 
tillate florets fertile, more numerous than the bisexual florets, corollas purple 
or purple-tipped. Bisexual florets fertile, corollas purple or purple-tipped. 
Cypselae oblong, 0.6-1.5 mm, epidermis minutely papillate (the papillae im- 
bricate and paired, nonmyxogenic); pappus of caducous, separate or weakly 
basally connate bristles separating in groups. Ba 

nnual, taprooted; leaves 

not at all clasping or sheathing; capitula in a globose 

uster; bisexual florets! p 

ercapitulum;capitulescence bracts 4-8. Euchiton sphaericus 


us-rooted; leaves subclasping or sheathing; capitula in a 

mispheric cluster; bisex 

ual florets 3-7 per capitulum;capitulescence bracts 2-5. 

Stolons commonly pr 

sent, plants forming colonies; basal leaves present in a 

rosette at flowering; ca 

uline leaves 2-4(-6), linear to oblanceolate, 0.5-3(-4) cm 

(Labill.) Maiden & Betciie, Census New South Wales PI, 204. 1016. Typi;; AUSTRALIA. 
Flowering (late March-)May-October(-November). Grassy hills, margins and 
openings in woods, roadsides, cutover areas; ca. 100-2500 ft (30-750 m); California 
and Oregon, Hawaii. Apparently native to Australia and New Zealand. Recorded 
in Australia as "common and widespread, particularly in cooler parts ..., often 
colonizing bare ground, stream and track margins etc." (Walsh 1993, p. 823) and 
"woodland areas near Canberra and also extending to upper slopes of moun- 
tains; widespread in many parts of Australia and Tasmania" (Burbidge & Gray 
1970). In New Zealand, it occurs in a wide variety of habitats but is "typically 
associated with forest and scrub communities between sea level and 1,000 (- 
2,000) ft" (Drury 1972, p. 144). 

A handwritten note by J.P. Tracy on the JEPS sheet of Tracy HllO notes the 
following: "My record of this plant [Euchiton gymnocephalus, presumably in 
Humboldt County] extends back to 1900 when it was already well estab." Cita- 


L971, Anderson and 



serpentine soil, 15 May 1965, J 


670 (DS); Lord Ellis 

road near I 

ity One, 600 ft, 30 Jul 1932, 

Parks and 


of the Mad River, 

grassy hil" 

: Quad., 

old skid road, shac 

ed rock 

y subsoU, 13 Aug 1 

e Oat C 

reek, logged dougf 

ir forest, 2^ 

below indicate that Tracy collected the species repeatedly in Humboldt 
ty from 1904 through 1946 and collections between 1960 and 1989 have 
nued to document its occurrence in counties of northeastern California 
outheastern Oregon. 

:ar coast, 300 ft, 12 May 1929, Tracy 8588 (CAS, UC). Humboldt Co.: near Azalea Reserve State 
ool woods in partial shade, 100 ft, 15 Jul 1960, Adams s.n. (HSO; along Mad River ca. 2 mi S of 
SO; 8 mi W of Berry Summit along US 299, 

ake, 25 Nov 1926, Ki Idale 2830 (DS); Trinidad, 
1133 (UC); above dirt road from old 101 to the 
n half shade, 22 Jul 1967, Stevens 61 (HSC); 
od forest and mixed evergreen forest, 850 ft, (HSC); Shubrick Peak Quad., Kinsey 
1978, Sutherland (HSC); immediate 

-I, UC); Boynton Prairie road, near Carroll place, 1000 ft, 23 Jun 1918, Tracy 4958 (NCU, UC); near 
ue Lake, in woods, spreading by stolons and forming patches, 1000 ft, 1 Apr 1923, Tracy 6175 Q^FS, 
:); near Boynton Prairie, in recently logged area, among "fireweeds," 2000 ft, 28 Aug 1927, Tracy 
58 (UC); Trinidad, logged-off lands, spreading in mats, 10-200 ft, 13 Aug 1932, Tracy 10349 (ARIZ, 

20 Jul 1935, Tracy 14110 (CAS, JEPS, UC-2 sheets); 4 mi SE of Korbel, near "Angels Ranch," local in 
assy moist places, 1000 ft, 22 Mar 1936, Tracy 14784 iCAS, DS, JEPS, UC-2 sheets); Fickle Hill, 6 mi 
; of Areata, in recently cleared land, 2000 ft, 17 May 1936, Tracy 14811 (UC); 2 mi NE of Orick, on 
id Hills road, local by roadside, 500 ft, 29 May 1936, Tracy 14829 (CAS, DS, GH, JEPS, UC); hillsides 
ar Canyon Creek, 6 mi SE of Blue Lake, in logged-over land, 1200 ft, 1 Aug 1936, Tracy 15058 (UC); 

ag 1936, Tracy 15175 (UC); vicinity of Carlotta, in recently cleared land, ca. 100 ft, 15 Jun 1938, Tracy 
936 (UC); "Riverside Park" near old Strong's Station District School House (burned down), 30 May 
46, Tracy 17548 (UC); Kneeland Prairie, Dan McBride's place (SE end of Kneeland), local, in partial 
ade of open fir woods, 2500 ft, 23 Jun 1946, Tracy 17596 (UC). Mendocino Co.: 15 air mi SW of 
irberville, 60 air mi SSE of Eureka, 1/4 mi E of Bear Harbor on Lost Coast Trail, edge of trail bed, 
difornia bay riparian forest, 120 ft, 8 Jul 1989, Bowcutt 1369 {HSC); 3.2 km N of jet with Rte 1, along 
;al Rd, openings in second growth redwoods, 305 m, 11 Jun 1981, Smith 6632 (CAS, HSC). Siskyou 
).: Forest Service Road 17N04, Doe Flat, T17N, R4E, Sec. 35, [no date]. Van Deventers.n. (HSC). HA- 

just N of Brookings, 10 Jun 1964, Chambers 2241 (DS, NCU); Brookings, 100 ft, 1 Aug 1937, Tracy 15606 
(UC); moist slope 5 mi N of Brookings, 7 Jul 1939, Peck 20445 (CAS, UC). Klamath Co.: Winema Na- 
tional Forest, Cold Springs Road and State Route 140, T36S R6E. Sec. 18, 14 Oct 1978, Sawyer 3320 

Wagner et al. (1997) recorded Euchitongymnocephalus, identified as E.japonicus 
(Thunb.) Holub, as a new state record for Hawaii. In Australia it was identified 
as Gnaphaliumjaponicum Thunb. by Burbidge and Gray (1970), but the syn- 
onymy of G.japonicum with G.gymnocephalum was later rejected without spe- 
cific comment by Cooke (1986) and Walsh (1993). In the present review, plants 

from Japan and China appear to be consistently distinct from related ones of 
Australia and New Zealand, althougfi differences are subtle, and until some- 
one may provide a more detailed study of the complex, the morphological af- 
finity of the North American plants with those of Australia is reinforced by the 
nomenclature. Differences between the two taxa are outlined in the following 

Basal leaves usually narrowly oblanceolate, attenuate to a distinct petiolar portion, 
the cauline abruptly differentiated in size and shape from the basal, smaller and lin- 
ear-lanceolate to linear-oblanceolate; adaxial leaf surfaces quickly glabrescent and 

mm long. Euchiton gymnocephalus 

Basal and cauline leaves linear-lanceolate to linear-oblong, without a distinct petiolar 
portion, the lower cauline slightly reduced in size but similar in shape to the basal; 

cypselae ca. 1.0 mm long. Euchiton japonicus 

Plants f romjapan, China, and Taiwan (specimens examined), and probably from 
Ryukyus and Korea are typical E. japonicus (see citation below). Collections re- 
ferable to E. japonicus or E. gymnocephalus from New Guinea, New Caledonia, 
and Java need to be studied in order to assess their relationship. Reports of E. 
japonicus from the Phihppines evidently refer, at least in part, to Gnaphalium 
ohlanceolatum Elmer, which apparently is more closely related to E. 
involucratus (but probably not conspecific with it) than to E. japonicus. 



1974 (non A. A 

55. 1786. Type: NEW ZEALAND. 

places, often moist or wet; ca. 50-600 m; 
o Australia and New Zealand; collections 
also observed (present study) from New Guinea, New Caledonia, Java, and Tai- 
wan. The species in Australia is recorded as a "weed of gardens and disturbed 
ground" (Burbidge & Gray 1970); it also is common "particularly in swampy 
sites, from near sea-level to the higher alps where usually in Sphagnum bogs" 
(Walsh 1993). 

Both of the North American collections cited below probably represent 
waifs, because this species apparently has not been subsequently recorded in 
floristic summaries from anywhere in the USA. This apparently is the first 
pubhshed report of E. involucratus from North America. 

Collections examined. UNITED STATES. California. Humboldt Co.: S end of Fickle Hill, 6 mi SE of 
Areata, ca. 2000 ft, moist ground, among rushes and sedges, spreading by rootstocks, perennial, 26 

s. 8:58. 1919. 1 

Type: HAWAII. 
Gnaphaliumjaponicum sensu various authors, non Thunberg 1784. 

Flowering (late June, Tracy 15974; July-)August-October. Grassy open places m 
wooded areas, recent clearings and clearcuts, disturbed soil, especially along 
roadsides; ca. 100-2000 ft 130-600 ml; California and Oregon, Hawaii (Molokai, 
Lanai, Maui, and Hawaii, fide Wagner et al. 1999). "Reported from San Joaquin 
County," California (Ferris 1960; voucher not seen in present study). Native to 
Australia and New Zealand; also documented from New Guinea, New 
Caledonia, Java, Philippines, Japan, and Taiwan (Drury 1972). In Australia re- 
corded as "a very common and widespread species, occurring on a wide variety 
of substrates, e.g., mallee and coastal sands, clayey floodplain area, and often 
colonizing disturbed ground" (Walsh 1993). 

Euchiton sphaericus has long been identified in California, apparently be- 
ginning with Howell (1937), as Gnaphaliumjaponicum; it recently was correctly 
identified by Wagner et al. (1997) and by Peter W. Michael (annotations on speci- 
mens at GH). Correct nomenclature was earlier provided by Drury (1972). 
Euchiton sphaericus was established in California at least by 1915, when appar- 
ently first collected there.J.P Tracy collected it repeatedly from that year through 
1949; from one locality (Tracy W52) he noted that it was "scarce now, abun- 
dant 15 years ago." Tracy also observed (fide label of Tracy 5091) that the species 
was "probably introduced in grass seed sown on logged-over country" Relatively 
recent collections (1954, 1977, 1992) confirm its persistence in northwestern 

Euchiton sphaericus was collected in Hawaii, perhaps for the first time, in 
1909 (on Maui, Eaurie 92S-BISH, as noted by Wagner et al. 1999). Rock (1914, p. 
352) noted that in Hawaii E. sphaericus was among several species "imported 
accidentally during the last 10 or 20 years, by the cattle estates with grass seeds." 
Collections examined. UNITED STATES. California. Del Norte Co.: Sutton Creek, 2 mi E of the Van 
Deventer Ranch and 2.5 mi air4ine E of Fort Dick, 350 ft, 15 Oct 1954, VanDevmier 412 (UC). Humboldt 
Co.: Eel River, 16 mi N of Garberville, 29 Aug 1936, How d\ 12888 {GH, LL, WIS); logged redwood flat, 
2 mi E of Carlotta, 15 Aug 1936, Jepson 11.898 QEPS); mouth of Laribee Creek, common as a weed but 
local, 200 ft, 12 Sep 1915, Tracy 4677 (CAS, MO, UC); around Humboldt Bay, Newells Camp on old 

1918, Tracy 5091 (NCU, UC-2 sheets); South Fork of Eel River, 4 mi above the mouth, common road- 

C) North Fork ot Mad River along newly con: 
on there 1000 It 19 Sep 1920 Ttacy5399i\JQ 
925 Inuy 7229 (UC) along 1 ord Ellis Road 1 
8Au£^lQ31 TtaLy9608(UO near Dyerville n 

r Creek recentl) lot^gtc 
>dwoodHoui,e 1500 it 
(UC) Fickle Hill bmi 

tl949 T>aLyl859HUC WIS) Aikens Campground TION R5E Sec 
1977 Sawyer 2990 (HSC) Lake Co road to Bartlett Mountain Aug 
ino Co 10 air mi W ol Leggett along and m dirt roadbed oi Hotel 

ml 30)1111927 Topf int;sn(G 
I 20 Jul 1919 Ptcfe 8924 IGH) 

1 am grateful to the staffs at GH, MO, NCU, and TEX-LL for help during recent 
visits to those institutions. Loans were studied from ARIZ, CAS, DS, HSC, JEPS, 
UC, and WIS. Comments by John Strother contributed to 
clarity of presentation. 

Opera Bot. 104:5-195. 
Buchanan, A.M. 1 999. A new species of Euchiton (Gnapha 

Tasmania, Australia. Pap. Proc. Roy. Soc.Tasmania 133:115-1 16. 
BuRBiDGE,N.T. and M.Gray. 1970. Flora of the Australian Capital Territory. Australian Natio 

Univ. Press, Canberra. 
Cooke, D.A.I 986. Gnap/ia//L/m. In: Jessop, J. P and H.R.Toelken,eds. Flora of South Austra 

Part III, Polemoniaceae-Compositae. South Australian Govt.Printing Division, Adelai 

Drury, D.G. 1972. The cluster and solitary^headed cudweeds native to New Zealand: 

{Gnopholium section Euchiton -Compositae).NewZeal.J. Bot. 10:1 12-179. 
Ferris, R.S.1960.Vol.lV,Bignoniaceae to Compositae. In Abrams, Land R.S.Ferris, Illustrated 

flora of the Pacific States. Stanford Univ. Press, Stanford, California. 
HoLUB, J. 1 974. New names in phanerogannae. Folia Geobot. & Phytotax. 9:261-275. 
Howell, J.T. 1937. Three species of Gnaphalium adventive in California. Leafl. W. Bot. 2: 


RegniVeg. 13:352-361. 

Bishop Mus.Occ. Papers 48:51-65. 

(rev. ed.), Volume 1 . Univ. of Hawai'i Press and Bishop Museum Press, Honolulu, Hawaii. 
Walsh, N.G. 1 993. Euchiton. In: N.G.Walsh and T.J. Entwhistle, eds. Flora of Victoria, Volume 

4. Inkata Press, Melbourne, Australia. Pp. 820-825. 
Walsh, N.G. 1 999. New species in Asteraceae from the subalps of southeastern Australia. 

Muellena 12:223-228. 
Ward, J.M. and I. Breitwiesfr. 1998. New combinations in Euchiton (Compositae - 

Gnaphalieae) from New Zealand. New Zeal. J. Bot. 36:303-304. 

Book Notici- 
Dennis Blagg. 2002. Big Bend Landscapes. (ISBN 1-58544-202-X, hbk.). Texas A&M 
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3982, 800-826-8911, 979-847-8752 fax). $40.00, 160 pp, 50 color paintings, 
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Dorn (1998; following Cronquist 1964) has treated Salix interior Rowlee at infra- 
specific rank within S. exigua Nutt, observing that a broad region of morpho- 
logical intergradation exists between the two where their ranges are in contact 
(see Dorns Fig. 2, p. 200). This observation is confirmed in field studies relating 
to a f loristic inventory of Lake Meredith National Recreation Area in Hutchin- 
son, Moore, and Potter counties of the north-central Texas panhandle (Nesom 
& O'Kennon in progress). In this area many populations exist of 5. interior stnsu 
stricto (colonial shrubs with completely glabrous, linear-lanceolate leaves with 
widely spaced and elongate teeth) as well as S. exigua (similar to S. interior but 
with densely silky-sericeous lower leaf surfaces). Also common are populations 
with foliar vestiture in varying degrees of intermediacy (e.g., Nesom & O'Kennon 
LM359, LM198 - BRIT). Salix nigra Marsh and S. amygdaloides Anderss. also 
commonly occur in the area, apparently without intergrades. 

Argus (1986) treated Salix exigua as including 5. interior without formal 
rank, but he noted (p. 91) that the "eastern phase (5. interior) differs from the 
western phase (S. exigua) in having leaves less densely sericeous, more distinctly 
toothed, and more veiny." In my observation (based on BRIT and VDB collec- 
tions), variability in vestiture is greater in the western phase than in the east- 
ern phase, where the green-glabrous leaves are consistently evident. Sericeous 
undersurfaces are characteristic of the western phase. 

Dorn recognized three varieties within Salix exigua subsp. exigua but 
treated S. interior only at subspecific rank: S. exigua subsp. interior (Rowlee) 
Cronquist. The following combination provides consistent nomenclatural rec- 
ognition of all infraspecific taxa, following ICBN Article 4.1: "The secondary 
ranks of taxa in descending sequence are tribe (tribus) between family and ge- 
nus, section (sectio) and series (series) between genus and species, and variety 
(varietas) and form (forma) below species" (see commentary in Turner & Nesom, 
Sida 19:257-262. 2000). 

Salix exigua var. sericans (Nees) Nesom, comb, nov Basionym: Salix longijolia var. 
sericans Nees in Wied-Neuw,, Reise Nord-America 2:448. 1841. TYPE: not seen. 

This is the oldest available name at varietal rank for plants identified as Salix 
interior. As noted by Dorn (1994, p. 92), the type locality apparently is "in or 
near the present Cass Co., Nebraska" (eastern Nebraska, well within the range 
of typical S. interior and east of the area of its co-occurrence with 5. exigua var. 

exigua, although the epithet imphes that the lea\ 
pletely glabrous. 

Comments by George Diggs and an anonymous re 

Argus, G.W. 1 986, The genus Salix (Salicaceae) i 

Monogr. 9:1-1 70. 
Cronquist, a. 1 964. Salix. Vase. PI. Pacific Northw. 2:37- 
DoRN, R.D. 1994. North American Salix (Salicaceae): 

DoRN, R.D. 1998. A taxonomic study of Solix sectioi 





MihaiCostea^ Fran(;ois J.Tardif 

Department of Plant Agriculture Departmentof Plant Agriculture 

e new characters, the relatio; 
corded from Romania. 

idcs^LdE. hTspidu^cfaTellsodi'scusS. ForThe"f°rrdmI, E.colo, 



Echinochloa is an economically important genus because many of its species 
are noxious weeds. In Europe, E. hispidula, E. oryzoidesand E. oryzicola are most 
often found as weeds in rice fields while E. crus-galli and E. colona although 
present in rice fields, are more often encountered in irrigated fields and amongst 
vegetable crops (Kossenko 1947; Vasconcellos 1954; Pnola 1965; Chirila 1984; 
Tzvelev 1976; Haf linger & Scholz 1980; Carretero 1981; Mosyakin 1996; Ciocarian 
2000). The taxonomy of the genus is controversial due to nomenclatural prob- 

lems and the continuous morphological variation exhibited by the taxa. For 
the practical purpose of plant identification, the species often lack conspicu- 
ous identification characters. Previous treatments and taxonomic studies are 
often contradictory and, consequently, the limits of the taxa are uncertain and 
specimens in herbaria are often misidentified. Other important taxonomic treat- 
ments of the genus (or part of the genus) are those of Stapf (f 899, 1934), Hitch- 
cock (1920), Wiegand (1921), Kossenko (1947), Ohwi (1942, 1962), Martinez 
Crovetto (1942), Vasconcellos (1954), Bor (1960), Pirola (1965), Chinla (1967, 
1984), Yabuno (1962, 1981), All (1968), Gould et al. (1972), Tzvelev (1976), 
Haf linger & Scholz (1980), Clayton (1980), Carretero (1981), Michael (1983) and 
Brussoni (1994). A general characterization of the genus can be found in Wat- 
son and Dallwitz (1992-on wards). Carretero (1981) provided a summary of the 
morphologic characters for European weedy species of the genus Echinochloa. 

The identification of new characters useful in determining taxonomic 
boundaries have shed light on our understanding of difficult groups such as 
grasses. The study of lemma micromorphology can provide characters valu- 
able in elucidating the delimitation of taxa, and these characters may reflect 
systematic relationships. Micromorphological features of glumes and bracts 
(lemma and palea) have been studied in other genera from Poaccae by: Bjorkman 
(1960), Hsu (1965), Baum (1971), Lucas (1979), Clark and Gould (1975), Thomas- 
son (1978a; 1978b, 1980, 1981, 1986), Shaw and Smems (1979), Terrel and Wergm 
(1981), Terrel et al. (1983), Webster and Hatch (1983), Bark worth (1983), Peterson 
(1989), Soderstrom and Zuloaga (1989), Kellogg (1990), Zuloaga and Judziewicz 
(1991), Valdes-Reyna and Hatch (1991), Molina (1993), Naredo et al. (1993), Ball 
et al. (1993, 1999) and Snow (1996, 1998). 

Caryopsis morphology has received a lot of attention since the beginning 
of the century; however, most studies have focused on the general morphology 
and the structure of the embryo. There is comparatively less information avail- 
able on the anatomy of the caryopsis coat in various other grasses (Netolitzky 
1926; Anderson 1927; Avery 1930; Krauss 1933; Hayvard 1938; Akerberg 1943; 
Bradbury et al. 1956; MacLeod & Palmer 1966; Kowal & Rudnicka-Sternowa 
1969; Rost 1973; Jones & Rost 1989; Rost et al. 1990; Ungurean & Costea 1994, 
1997). According to our knowledge, micromorphology and anatomy of the cary- 
opsis m Echinochloa have not been studied so far 

The purpose of this study is to evaluate the taxonomic usefulness of select 
floral characters in Echinochloa, namely, micro- and macrocharacters of the 
lemma and caryopsis, the caryopsis coat, and the size of starch grains. Further- 
more, the nomenclature, the taxa limits and the interspecific relationships in 
Echinochloa are also discussed. 

Samples were collected and identified from populations in Spain and Romania. 

The accessions from Spam were collected and identified by Professor Jose Luis 
Carretero from Polytechnic University Valencia, Spain (Table 1). The samples 
from Romania were collected and identified by the first author (Table 1). Mature 
spikelets were collected from the median region of the panicles as the spikelets 
from the upper parts of the inflorescence tend to be smaller Micromorphologi- 
cal characteristics were assessed on 20 specimens in each population. Voucher 
specimens for all the taxa are preserved in the VALA, BUAG and BUG Herbaria 

Micromorphology of sterile lemmas and caryopses— Ten mature caryopses 
for each specimen were examined. Only the adaxial (ventral) face of sterile lem- 
mas and caryopses was observed. Sterile lemmas and caryopses were mounted 
on aluminium stubs with Avery's spot-o-glue and then coated with 20 nm of 
gold using Bio-Rad Sputter-Coatter SC-500. Samples were examined with 0° 
tilt at 5-15 KV on a Hitachi S-4100 Scanning Electron Microscope. 

Structure of the caryopsis coat.— Mature caryopses with the glumes and 
bracts removed were soaked in warm water for 1 hour. Afterwards, they were 
fixed in FAA (90 ml 95% ethanol, 5 ml formalin and 5 ml glacial acetic acid) for 
48 h. Caryopses were transferred to 70% ethanol, dehydrated in tertiary butyl 
alcohol (TBA), and embedded in Tissuemat. Ten mature caryopses for each speci- 
men were serially sectioned to 5-10 |im, stained with safranin and fast green, 
mounted in Canada balsam, and examined with standard brightfield optics 
and with polarized light. A small number of caryopses belonging to each ac- 
cession were soaked, fixed, embedded and sectioned without removing the 
glumes and bracts. The anatomy drawings were made using a Reichart camera 
lucida. The lengths of the largest fifty starch grains located in the mealy en- 
dosperm from each caryopsis, from each accession were measured as well. 

Depending on the environmental conditions, an Echinochloa spp. plant can 
produce 1500-22000 spikelets distributed on 9-25 panicle bearing culms. The 
spikelets have 2 flowers; the lower floret is sterile consisting only of lemma and 
palea. Sometimes the lower floret is staminate (especially in E. colona). The ster- 
ile lemma may be awned; but this character is relatively insignificant. The spike- 
lets of E. colona are unawned, those of E.oryzoides are always awned, and those 
of the other species may be awned or not. An important character, emphasised 
by all authors, is the size of the spikelets. The lower glume is about 1/2 the length 
of the spikelets in £. colona; 1/2-3/4 in E. crus-galli, E.oryzoides and E. hispidula, 
and 1/2-3/5 in E. oryzicola. The lemma (in the species examined) is becoming 
indurated and difficult to remove. Micromorphological characters of the lemma 
for Echinochloa include: short cells (silica cells), long cells, bicellular microhairs, 
papillae and microhairs. Stomata may occasionally occur in all species. Cork 
cells were not observed on the Echinochloa sterile lemmas. Silica bodies are 







Echinochloa hi 


Spain C 






Echinochloa or 





la (BUAG) 

a-Department of Botany, Poiyteclnnic 

b-Department of Botany, University of Agronomical Sciences Bucharest, Romania 

c-Botanical Garden, University of Bucharest, Romania. 

restricted to the intercostal regions. They can be cross-shaped or i 
shaped. Within each category, 2 types can be further recognised. 
Cross-shaped type (ratio length:width = 1:1). 

CI. Sihca bodies with the two endings deeply divided (Fig. le). 

C2. Endings only slightly bilobed (Fig. 2b). 

D2. The two endings emarginated or bilobed (Fig. Id). 
Even if one or two types of silica bodies predominate in one species, one sterile 
lemma may often contain sporadically the other types as well. 

Long cells are easily recognizable by their length and their sinuous mar- 
gins. Long cells are associated with papillae at their distal ends, especially near 
the apex of lemmas. Bicellular hairs, 30-60 jim long, are present in all species 
and belong to the panicoid type (Tateoka et al. 1959; Amarasinghe & Watson 
1988; Watson & Dallwitz 1992-onwards). All the trichomes observed in 
Echinochloa (bicellular microhairs excepted) are more or less macroscopic. If 
shorter, they are only an earlier stage in the development of macrohairs. They 
tend to increase in frequency and length toward the distal parts of the lemmas. 
The marginal veins usually have the longest hairs. They are somewhat shorter 
or even missing from the median and lateral veins. Macrohairs are present in 
the intercostal regions but they are usually shorter compared to those of the 
marginal veins. In E. oryzicola, sterile lemmas often appear glabrous in the in- 
tercostal regions. The macrohairs are rigid, pointed and oriented toward the 
apex of the lemma. 

Morphology of caryopsis (Fig. 3, a-e). Caryopsis is the dry, monospermous, inde- 
hiscent fruit of grasses in which the layers of pericarp are fused with the seed 
coat. The adnation between the pericarp and the seed coat starts in the placento- 
chalazal region and extends toward the rest of the caryopsis (Izaguire de Artucio 
& Laguardia 1987). 

The size of caryopsis varies proportionally with the size of the spikelet and 
therefore is also an important differential character. The caryopsis is round, ellip- 
soid to ovate, with the ventral (adaxial) face rounded and the dorsal face (abaxial) 
more or less flat. The ventral face shows the axis of the embryo. The embryo is 
large, about 0.5 times the caryopsis length in E colona; almost as long as the 
caryopsis in E. oryzicola and 0.6-0.7 times the caryopsis length in the other 
examined species. The scutellar region is visible around the embryo axis. The 
dorsal face features the basal pointed coleorhiza, which is also encircled by a 
rounded scutellar region. The hilum may or may not project on the outhne of 
the caryopsis. When plants shed their spikelets, they land with the convex face 
down, m a position that brings the embryo m the most favourable position for 
germination. The glumes and sterile lemma then absorb the water necessary 
for germination. When they land in water, the spikelets float until imbibition 
is completed after which they sink and fall to the bottom of the water (Costea, 
unpublished). The mature caryopsis coat is shiny and whitish, yellowish or 
brown. The scutellar region prolonged around the embryo (on the convex, ven- 
tral face), is smooth or wrinkled, and may have black spots (E. oryzicola). 

Micromorphology of caryopsis (Fig. 2, c-f) is not similar in the Echinochloa 
species examined. The epicarp cells are elongated ranging from 60-120 |im in 
length and 10-20 ^im in width. The periclinal walls are flat, except for E. 
oryzicola which has concave periclinal walls (Fig. 2f). The anticlinal walls are 
protuberant and undulated with the amplitude, the width at the base and the 
shape of the undulations having diagnostic value. For example, in E. crus-galli 
(Fig. 2d) the anticlinal walls are ^-undulated (undulations rounded, wider to- 
ward the apex and narrower at the base), in E. orzyoides and £. hispidula they 
are S-undulated, (Fig. 2c and e) and in E.oryzicola they are Z-undulated (Fig. 2f). 

Anatomy of the caryopsis coat (Fig. 4, a-c) m the examined species is simi- 
lar. The differences observed are minute and quantitative. The caryopsis coat 
consists of adnate layers of pericarp, seed coat and nucellus that surround the 
endosperm and embryonic axis. If the lemma is not removed, cross-sections 
reveal its connivence with the caryopsis. The structure of the lemma resembles 
the structure of the leaf. A homogenous mesophyll consisting of 2-4 cell layers 
can be observed between the 2 epidermis. During the early stages of develop- 
ment the mesophyll cells contain chloroplasts. These cells are larger than the 
epidermis cells, with thickened, sclerified cell walls at maturity (Fig 4, 1-c). 


'/, b. f. hispidula, c. £. oryzkola, d. E. oryzoides.e.E. colona. Scale bar 


FiG.4. Anatomy of caryop; 

,sg-starch grains. Scale bar 10 

The veins in young lemmas resemble the tertiary veins of the leaves. The cary- 
opsis coat is thin (7-15 )im thick), with a simple structure. It consists of only 2-3 
cell layers. The one-layered epicarp is the most representative component of the 
pericarp. The epicarp cells have thickened walls. The mesocarp is also single- 
layered with smaller and often crushed cells. At caryopsis maturity the endocarp 
is usually no longer visible. In younger caryopses it consists of long thin-walled 
cells ("tube-cells"), parallel to the epicarp cells. The seed coat is fused with the 
pericarp and it is not structured, visible only as brown line. The nucellus per- 
sists as a thin remnant. A single layered alleurone is continuous around the entire 
endosperm. The alleurone layer can be locally 2-layered in E. oryzoides (Fig. 
4c). The central endosperm consists of large, irregularly shaped cells that con- 
tain simple starch grains. The central endosperm does not have a homogenous 

appearance. Around the embryo the endosperm is "mealy" and soft while in 
the rest of the caryopsis is hard and glassy In most of the species exammed, the 
glassy endosperm predominates. However, in E.oryzoides the mealy endosperm 
is equaly or better developed than the glassy endosperm. The mealy endosperm 
contains rounded, isolated starch grains; only rarely they are aggregated (but 
not compound) in small groups of 2-4 granules. The hard endosperm has po- 
lygonal (rarely round) starch grains, closely packed together. This last type cor- 
responds to the Panicum type described by Tateoka (1962). The average size of 
the starch grains is significant and it varies between 5.5 |im and 10.5 |im de- 
pending on the species. The embryo has a scutellar tail, without an epiblast. 

Nomenclature.— Most of the nomenclatural problems of E oryzicola are related 
to the binomial Panicum phy/lopogon on which Echinochloa phyHopogon (Stapf) 
Stapf ex Koss. is based. Stapf 's diagnosis of Panicum phyllopogon addresses only 
the vegetative features of the plants, particularly the presence of hair tufts in 
the collar region of the leaves. For a long time this feature was erroneously con- 
sidered to be an exclusive characteristic of only one species— £. phyllopogon, 
thus generating extensive nomenclatural confusion. The individuals of 
Echinochloaoryzicola—a more recent name— are constantly showing this veg- 
etative characteristic. Consequently E.oryzicola has almost generally been con- 
sidered a synonym of E. phyllopogon (Pirola 1965; Morariu 1972; Kerguelen 1975, 
1993, 2002; Czerepanov 1981; Michael 1983; Chirila 1984; Ciocarlan 2000). 
Kossenko (1940) further contributed to this confusion when he subordinated 
E.oryzicola as a subspecies of E. phyllopogon (1940). Other authors erroneously 
considered E. oryzicola to be a synonym of E. hispidula (Ohwi 1962; Gould et 
al. 1972) or even of E oryzoides (Clayton 1980). 

Stapf 's collections of Panicum phyllopogon at Kew comprise both vegeta- 
tive and fertile specimens. Tufts of hairs are obvious in most of the vegetative 
specimens and in several plants bearing inflorescences. Carretero (1981) noted 
that tufts of hair might also occur in the leaf collar of E. hispidula (1981). After 
examined Stapf 's collection, Carretero (1981) synonymized E phyllopogon with 
E hispidula. Michael initially regarded Stapf 's collection (1975 in herb, 1983) as 
a mixture of E phyllopogon (the plants showing the vegetative characteristics) 
and E oryzoides. Later, after examining populations from Spain, Italy and 
France he observed that individuals of E. oryzoides may also have leaves with 
hair tufts in the collar region and accordingly he synonymized E. phyllopogon 
with E. oryzoides. The author selected as a lectotype for Panicum phyllopogon 
the one sheet in which the inflorescence and tufts of hairs are evident on the 
same specimen (1985, note in the Kew Herbarium on the specimen selected as a 
lectotye, collected by Jacometti from "Novarra, in rice fields"). We support 

Michael's view in that E. phyllopogon may be a synonym of £. oryzoides and that 
Stapf 's collection is a mixture of E.oryzicola and E.oryzoides ( = E phyllopogon). 

Many recent studies on the biology, ecology, herbicide resistance and physi- 
ology have been also using the binomial '"Echinochloa phyllopogon" referring 
probably to E. oryzicola (Fox & Kennedy 1994; Fox et al. 1995; Mujer et al. 1995; 
Gibson et al. 1999; Fischer et al. 2000; Fischer et al. 2000). 

Identification key-The following key is modified from Carretero (1981) 
and Michael (1983). 

In order to improve chances of correctly identifying the species, a range of 
individuals from the same population and a range of spikelets and caryopses 
belonging to the same plant should be collected and examined. Spikelets length 
measurements do not include the awns. Caryopsis measurements refer to dry 
caryopses. Starch grains should be observed in the mealy endosperm (around 

1. Spikelets 2-3 mm long.regularly arranged on the racemes.Caryopses whitish, 0.7- 

1.2 mm long with the embryo 0.4-0.5 of the caryopsis length E.colona 

Coleoptyle red-f 

)urplish. Leaves often with a tuft of t 

jrown hairs in the collar 

region. Lower gl 

ume 1/2-3/5 the length of the spi 

kelets, with the 3 veins 

visible only at the base. Sterile lemma often glabroi 

js and shiny, rarely with 

stiff macrohairs. 

up to 0.8 mm. Caryopses brownish, : 

?-2.4 mm long, with the 

embryo 0.9 of th 

e caryopsis length, and the glassy endosperm equal or more 

the mealv endosperm 

E. oryzicola 

Coleoptyle g reel 

1. Leaves only rarely with a tuft of hai 

TS in the collar region of 

the leaves. Lowe 

r glume 1/2-1/3 the length of the sp 

3ikelets, with the 3 veins 

visible along the 

vided with dens 

e, stiff hairs up to 1 mm long. Caryof 

Dsis light-yellow 2.2-2.8 

mm long, with t 

he embryo 0.6-0.7 of the caryopsi: 

5 length and the mealy 

endosperm mor 

e developed than the glassy endosf 



4. Weeds of rice 

fields, occasionally with a tuft of hai 

rs in the collar region of 


> panicle is not pyramidal with branches often whorled.Spike- 

lets ovate-elli 

ptical, 3.3-3.8 mm long. Caryopsis 2 

-2.2 mm long with the 


■transverselv wrinkled 

4. Weeds not ot 

3ligate of rice, but sometimes occui 

-ring in rice fields espe- 


nargins of the ponds. Leaves never v 

.ith hairs in the collar of 

the leaves. Panicle pyramidal with the branches n. 

ever obviously whorled. 

Spikelets ovat 

:e, 2.8-3.4 mm.Caryopsis ovate, 1. 4- 



, Echinochloa colona (L.) Link, Hort. Berol. 2:209. 1833. type "JAMAICA:" 
Browne (LINN). Panicum colonum L, Syst. Nat., ed. 10, 2;870. 1759. Echinochloa crus- 
Beauv. subsp. colona (L.) Honda, Bot. Mag. (Tokyo), 37:22.1923. 

There is still disagreement over how to write the specific epithet, as various 
authors have used either \olonum" or \olona" to designate the species The spe- 
cific epithet is most likely derived from the medieval Latin adjective "colonus- 
a-um" in which case ''Echinochloa colona" is the correct spelling. Another hy- 
pothesis (Carretero 1981) is that the specific epithet results from the contraction 
of the word "colonorum," the plural genitive of ''colonus-i" although we believe 
this to be less probable. 

Echinochloa colona is the most easily recognised species. It is annual, often 
rooting at the lower nodes. The leaves may have purplish transversal bands. The 
inflorescence is usually erect, with short branches (the lower ones are shorter 
to equalling 3 cm long). The spikelets are regularly arranged on the racemes. 
Often the lower floret of the spikelets is staminate. The lower lemma is awn- 
less, not exceeding 2 mm long. The spikelets are ovate, 2-3 x 1.2-1.8 mm. The 
lower glume is 3-veined, about 1/2 as long as the spikelet. The upper glume is 5- 
veined. The sterile lemma is 7-veined. The median vein may or may not have 
macrohairs. Longest macrohairs, 1 mm long. Lateral veins are only distally con- 
spicuous. The silica bodies are of type D2. The stigmas are dark-red. 

The caryopsis (Fig. 3e) is round to almost so, whitish and translucent, (0.7-) 
0.9-1.2 X 0.7-1.1 mm. The embryo is 0.4-0.5 of the caryopsis length. The epicarp 
cells are 40-65 x 8-12 ^m. The periclinal walls are flat. The anticlinal walls are 
moderately thick (2.1-2.7 |im), undulated; the amplitude of the undulations is 
5-10 |im, and their width is 6-12 |im. The undulations are Q-shaped or S-shaped. 
Starch grains, 5.6 (± 0.24) |im long. The chromosome number is 2n = 6x = 54 
(Carretero 1981; Yabuno 1985; Koul & Gohil 1991; Devesa et al. 1991). 

This species is a widespread weed m rice fields in tropical and subtropical 
areas of the globe. In Europe it has been recorded in the warmest countries 
(Spain, Portugal, Italy, France, Turkey Greece), where it grows as a ruderal or 
segetal weed in irrigated crops (and sometimes also in rice). It seems to be ex- 
panding towards typical temperate countries where it usually grows as a rud- 
eral. As an example, we are recording the occurrence of this species for the first 
time in Romania where we have encountered it since 1996 growing as a ruderal 
in the Railroad Station in Bucharest. 

2. Echinochloa oryzicola (Vasing.) Vasing. in Komarov, Fl. U.R.S.S. 2:33. 1934. Type: 
"Oriente Extremo inter segetes Oryzae satime L." Panicum oryzicola Vasing., Bull. Appl. Bot. 
Pl.-Breed. (i^enmgrad) 25(4);125. 1931. Echinochloa phyllopogon (Stapf) Koss. subsp. oryzicola 
(Vasing.) Koss., Not. Syst. Herb. Acad. U.R.S.S. 8(12):210. 1940. Echinochloa crus-galli (L) Beauv. 

= Echinochloa phyllopogon auct., non Stapf 
Annuals; the European populations with a tuft of brownish hairs in the collar 

region of the leaves. The hairs are obvious from the two or four leaf stage. Yabuno 
(1962, 1981) studying Asian populations found that different forms - notably 
his C and F forms - do not always exhibit the tuft of hairs in the collar region of 
the leaves. The inflorescence is normally erect and spreading but occasionally 
can be horizontal or pendent and may be green to red coloured. Spikelets are 
ovate-elliptical, awned (awn not exceeding 2 cm in length) or not, measuring 
3.9-4.8(-5) X 2.2-2.4 mm. The spikelets persist in the panicle longer than in E 
oryzoides. The lower glume is about 1/2-3/5 the length of the spikelet, with 3 
veins only partially visible. The upper glume is 5-7 veined. The sterile lemma 
IS 4-5 veined. The median veins may or may not have short hairs (0.5-0.6 mm); 
the laterals veins are conspicuous only toward the apex. The longest macrohairs 
are up to 0.8 |Lim long. The sterile lemma can be: a) glabrous and shiny, convex— 
with the lateral veins visible only along their distal part. In the intercostal re- 
gions the macrohairs are rare, short, 0.1-0.4 mm, with a swollen base (Fig. 1, c 
and d) (C-form of Yabuno 1962, 1981). This is the most common type encoun- 
tered in Europe, b) more or less flat and coarse, with dense long macrohairs, up 
to 0.8 mm in the intercostal regions (F-form of Yabuno 1962, 1981). The silica 
bodies may be type C2 (Fig. le) or D2 (Fig. Id). The stigmas are red. The cary- 
opsis is ellipsoidal to almost round measuring 2-2.4 x 1.8-2.1 mm (Fig. 3c). The 
embryo is 0.75-0.9 of the caryopsis length. Often the stigmas are persistent. The 
hilum is prominently visible on the outline of the caryopsis. The colour is 
brownish-red, brownish-green or brownish-grey. The scutellar zone is irregu- 
larly wrinkled often with black spots. The epicarp cells are 100-120 x 15 |im. 
The periclinal walls are concave (Fig. 4a). The anticlinal walls are moderately 
thick (2.6-3.2 iim) and undulated; the amplitude of the undulations is 8-10 jam, 
and their width is 6-7 |im (Fig. 2f). The undulations are Z-shaped (acute and 
narrow toward the apex) (Fig. 2f). The glassy endosperm predominates or it is 
as developed as the mealy endosperm. Starch grains are 7.14 (± 0.51) ^m in 
length. The chromosome number is 2n = 4x = 36 (Carretero 1981; Yabuno 1985, 

This species originated in China and SE Asia and is now a weed of rice in 
Europe, Asia, North and South America. The biology and ecology of a culti- 
vated form of E. oryzicola has been recently described by Hirosue et al. (2000). 

3. Echinochloa ( 

li (L.) Beauv, Ess. Agrost.:53, 161, tab. 11, fig. 2. 1812. ' 



um sheet r 

lumber 8C 

,18 (LINN) 


1920; Ba 

um 196; 




ituation oc 

:curs with I 


80.20) by L 

innaeus. A 


= Pamcumcriis-corvi L., Sp. PI, ed. 2, 1:84. 1762. 
Annuals with erect to spreading or decumbent stems, up to 120 cm. The panicle 
is often pyramidal with purplish nuances, erect, but sometimes nodding. The 
inflorescence branches are longer than 3 cm, patent or erect-patent, never ob- 
viously whorled. The lower branches are frequently twice branched. The spike- 
lets are irregularly arranged on the racemes and the lower floret is rarely stami- 
nate. Spikelets are ovate to broadly-ovate, 2.8-3.4 x 1.6-1.8 mm. The lemma of 
the lower floret is awnless or awned, the awn not exceeding 5 cm in length. The 
lower glume is 3-veined and about 1/2-1/3 the length of the spikelet. The up- 
per glume is 5-7 veined. The sterile lemma is 7-veined. The lateral veins are 
visible along their entire length or only toward the apex. The longest macrohairs 
are about 1 mm long. The silica bodies belong to types C2 or D2 (Fig. 2b). The 
stigmas are white or red. 

The caryopsis is ovate, 1.4-1.9 x 1.3-1.6 mm, brownish-red or brownish-grey 
in colour. The embryo is 0.6-0.7 of the caryopsis length (Fig. 3a). The epicarp 
cells are 60-80 x 10-15 |im. The periclmal walls are flat. The anticlmal walls 
are thick (2.6-3.2 |im) and undulated; the amplitude of the undulations is 8-10 
|im, and their width is 6-7 |j,m. The undulations are Q-shaped (Fig. 2d). The 
glassy endosperm predominates. The starch grains are 5.9 |im (± 0.37) in length. 
The chromosome number is 2n = 6x = 54 (Carretero 1981). 

This species is probably the most widespread of the genus, being a com- 
mon and noxious weed all over the world, especially in irrigated crops. It is less 
limited to warm chmate compared to E. colona and E. hispidula. In rice it usu- 
ally grows on the pond margins, but it may also penetrate the interior of the 
rice fields. 

4. Echinochloa hispidula CRctz)Neesc\Royle 111 Bot Hunal 11416 420 1840 

gani(L)Beau\ bubbp tftcfa (PollaciJCiierri &Giacomini Nomencl Fl Ital 120 1950 

Annuals which may occasionally have tufts of hairs in the collar region of the 
leaves. The panicle is rarely pyramidal, erect to pendent, green or with purplish 
nuances. The branches are whorled and more or less erect, except for the lower- 
most ones. Spikelets are ovate-elliptical of 3.3-3.6(-3.8) x 1.7-1.8 (-2) mm. The 
lower glume is 3-veined, about 1/2-1/3 of the spikelet length, and the upper 
glume is 5-7 veined. The sterile lemma is 7-veined resembling E crus-galli. The 
longest macrohairs are about 0.8 mm long (Fig. lb). The silica bodies are very 

variable, depending on the population - CI (h802) (Fig. lb), CI and Dl (h803, 
h805), CI and D2 (h805). The stigmas are white but may occasionally be red. 

The caryopsis is larger that of E. crus-galli, ovoid to oblong, 2-2.2 x L5-L8 
mm and brownish-red or brownish-grey in colour (Fig. 3b). The embryo is 0.6- 
0.7 the caryopsis length, and the scutellar zone is transversally wrinkled (Fig. 
3b). The hilum is projecting on the outline of the caryopsis. The epicarp cells 
are 65-90 x 10-15 |im. The periclinal walls are flat. The anticlinal walls are thick 
(2.6-3.2 |im) and S-undulated; the amplitude of the undulations is 8-10 )im and 
their width is 6-6.5 |im (Fig. 2e). The glassy endosperm predominates. The starch 
grains are in average 6.3 (± 0.47) |im in length. The chromosome numbers re- 
ported are: 2n = 6x = 54 (Devesa et al. 1991) or 2n = 36 (Carretero 1981; Feng & 
Zhang 1993). 

This taxon probably originated m the SE Asia. It is widespread in tropical 
and temperate areas of Asia, Africa, America and Australia. In Europe it grows 
only in the warmest regions (Spain, Italy, Portugal and France) as a weed of rice 

3. Echinochloa oryzoides (Ardumo) Fritsch, Verb. Zool.-Bot. Ges. Wien 41: 742. 
1891. type: "Semina hujus Panici inventa a me fuere inter Oryzam" Pamcnm oryzoides 

= Echinochloa crus-gaWi (L.) Beauv. subsp. oryzoides Bolos & Masclans, Collect. Bot. 4:420. 1955. 
= Panicum hostii Bleb., Fl. Taur. Cauc. 3:57. 1819. Echinochloa crus-galli (L.) Beauv. subsp. hostii 

= Panicum phyllopogon Stapf, Hook, Ic. Plant., ser. 4, plate 2698. 1901. Echinochloa phyllopogon 
(Stapf ) Stapf. ex Koss., Not. Syst. Herb. Acad. U.R.S.S. 8 (12):208. 1940. 

= Echinochloa macrocarpa Vasing. in Komarov, FL U.R.S.S. 2:739. 1934. Echinochloa crus-galli L 
Beauv, van macrocarpa (Vasing.) Morariu in Savulescu, Fl. Rom. 12:86. 1972. 

= Echinochloa coarctata Koss., Not. Syst. Herb. Acad. U.R.S.S. 9(1):28. 1941. 

= Echinochloa commutata Schultes in Roemer & Schultes, Syst. Veg., ed, 15, Mant. 2:267 1824. 

Annuals with erect to spreading or decumbent stems, up to 130 cm tall. The 
seedlings have a reddish coleoptyle. Tufts of hairs in the collar region may be 
infrequently present. The inflorescence is green, pendent and at maturity hang- 
ing almost horizontally, resembling rice in habit. The branches of the panicle 
are often adpressed to the main rachis. The spikelets are rather persistent, usu- 
ally awned (awn up to 5 cm long), broadly-ovate to ovate and (3.6-)4.1-5 x 2.2- 
2.6 jim. The lower glume is about 1/2-1/3 the length of the spikelet, with 3 veins 
visible across their entire length. The upper glume has 5 veins also entirely vis- 
ible. The sterile lemma is 7-veined, with the lateral veins usually visible along 
their entire course. The longest macrohairs are about 1 mm long (Fig. la). The 
silica bodies belong to the type Dl (Fig. la). The stigmas are red. 

The caryopsis is ovate to almost round, 2.2-2.8 x 1.9-2.3 mm (Fig. 3d) and 
yellowish. The embryo is 0.6-0.7 the caryopsis length, with the scutellar region 

zone smooth (Fig. 3d). The epicarp cells are 100-120 x 10-15 ^m. The periclinal 
walls are flat. The anticlinal walls are moderately thick (2-2.6 |im) and weakly 
S-undulated (Fig. 2c). The amplitude of the undulations is 1-3 |im, and their 
width is 8-11 |j.m. The mealy endosperm predominates. The starch grains are 
9.5 ^im (± 0.58) in length. The chromosome numbers reported are: 2n = 36 
(Carretero 1981) and 2n = 6x = 54 (Yabuno 1984; Feng & Zhang 1993). 

Asia, Europe, North America, South America and Australia. It is probably one 
of the most widespread species after E. cru^-galU and E. colona. 


The genus Echinochloa does not have qualitative characters which clearly de- 
marcate its species. A good example is the presence of hair tufts in the collar 
region of the leaves which has been considered an exclusive feature of E. 
phyllopogon. This assumption has subsequently caused a widespread nomen- 
clatural and taxonomic confusion because as we have previously indicated, three 
species may show this feature (E. oryzicola commonly; E. hispidula and E. 
oryzoides rarely). Even the most significant qualitative character states overlap 
between species. When such characters are noted they are always accompa- 
nied by the words "often," "usually," "normally," etc. because exceptions are pos- 
sible. The occurrence of a feature should be understood as a predilection and 

Lemmatal micromorphology—The long cells, microhairs and papillae are 
the same in all species examined. The pattern of papillae in Echinochloa spe- 
cies is similar to the pattern described in the fertile lemmas of some Panicum 
species (Clark & Gould 1975). The macrohairs are more or less similar in the 
examined taxa. Shorter hairs that could be described as prickles (Metcalfe 1960; 
Ellis 1979) were observed in all species. However, because these shorter hairs 
represent a developmental stage, only the macrohairs are recognised in this 
study as a distinct category (see Snow 1996, 1998 for considerations on homol- 
ogy and ontogeny of hairs in Poaceae). Variations in length, distribution and 
density of macrohairs are minute and there is considerable overlapping among 
species. The only exception is the short, rare macrohairs, with swollen bases 
from the intercostal regions of sterile lemmas in E.oryzicola (in the "form C of 
Yabuno 1962, 1981). These hairs are constant m their morphology and can be 
considered intermediate to prickles (Metcalfe 1960; Ellis 1979). Silica bodies have 
previously been considered structures of taxonomic significance in Poaceae 
(Metcalfe 1960; Ellis 1979; Palmer & Tucker 1981; Evoli & Pirola 1971). In 
Echinochloa, silica bodies vary in shape even within a small area of the same 
lemma. However, in most species there is a tendency toward one or two types. 
Echinochloa hispidula was the most variable of all species examined in this 

study. The silica bodies varied from population to population, with all the dif- 
ferent types observed. The previous epidermis studies conducted on leaves 
(Sanchez 1968; Pirola & Evoli 1971; Carretero 1981; Jin et al. 1986) are rather con- 
tradictory but they all showed an even greater variation of this character Even 
if the results obtained in this study revealed some differences between species 
(E. hispidula excepted), the shape characteristics of the silica bodies should be 
regarded with caution. More populations need to be examined before a final 
conclusion is reached. The same cautionary approach towards the taxonomic 
utility of the silica body shape is suggested by other studies conducted on 
Zizania (Terell & Wergin 1981), Oryza (Whang & Kim 1994) and Leptochloa 
(Snow 1996). Ball et al. (1999) showed that silica phytoliths examined individu- 
ally are taxonomically irrelevant. The authors instead used detailed 
morphometries, computer-assisted imagery and statistical analysis to develop 
a classification key of several Triticum and Hordeum species. The presence or 
absence of silica bodies in lemmas can prove useful at higher ranks (genera 
and above) since their presence is probably symplesiomorphic (Snow 1996). 

Caryopsis—Tht simple morphology of the caryopsis provides good char- 
acters for species identification. The size of caryopsis is important and corre- 
lated with the size of the spikelets. The other characters overlap between spe- 
cies but may be useful diagnostics is some cases, eg: caryopsis whitish, 
translucent (E. colona), yellowish (E. oryzoides); embryo 0.4-0.5 of the caryopsis 
length (E. colona) or 0.75-0.9 (E.oryzico la). Surprisingly, surface patterns of the 
caryopsis proved to be significant in the differentiation between almost all 
Echinochloa species. However, more populations should be examined before any 
definite conclusions can be reached. 

The anatomy of the caryopsis is of minimal systematic value. The overall 
organization of the caryopsis coat resembles the structures described in Setaria 
(Rost 1973) or Melica (Rost and Izaguire de Artucio 1990). The structural sim- 
plicity of the pericarp can be a functional result of the indurated lemma that 
protects the caryopsis. In those grasses where no coalescence between the cary- 
opsis and lemma occurs, or when the lemma is thin, the pericarp is more com- 
plex structurally (as for example in many Triticoideae— Netolitzky 1926; Avery 
1930; Krauss 1930; Bradbury et al. 1956; Ungurean & Costea 1994). The anatomy 
of the sterile lemmas in Echinochloa species could be more important as a taxo- 
nomic character than the structure of the caryopsis coat. The structural fea- 
tures and surface pattern of the lemma indicate its ontogenetic connection with 

The average size of the starch grains examined in a large number of en- 
dosperm cells and caryopses is constant and significant among the species ex- 
amined and therefore can be considered a diagnostic feature. This character is 
as significant in the taxonomy of the genus as other quantitatively important 
characters such as the size of spikelets, and size of caryopses. 

Relationships between species 

The individuality of Echinochloa species as treated in the present study is ap- 
parently reinforced by studies of their biology and ecology (Holm et al. 1977, 
1997; Szilvassy 1976; Chirila 1967; Carretero 1981; Yabuno 1983; Norris 1996; 
Honek &r Martinova 1996). The appropriate understanding of E oryzicola was 
delayed by nomenclature problems since this binomial has been almost gener- 
ally considered a synonym of E. phyllopogon (Pirola 1965; Morariu 1972; 
Kerguelen 1975, 1993, 2002; Czerepanov 1981; Michael 1983; Chirila 1984; 
Ciocarlan 2000). Apart from this nomenclatural confusion E. oryzicola is the 
second species easiest to differentiate (after E colona) even using classic mor- 
phologic characters. It has the second largest (after E oryzoides) spikelets, cary- 
opses and starch grains; the micromorphology of the sterile lemma and cary- 
opsis are quite peculiar as well. 

Yet apart from E. colona and E. oryzicola, which are usually easily 
recognisable, the other 3 taxa comprise a difficult complex. Echinochloa 
hispidula is extremely variable and shares close affinities with both E oryzoides 
and E crus-galli. Several current treatments tend to view E hispidula as a sub- 
species or as a variety of E crus-galli (Michael 1983; Mateo Sanz 1990; Devesa 
1991; Janzein 1993; Mateo Sanz & Crespo Villalba 1995; Kerguelen 1993, 2002; 
Asins et al. 1999). However, Gonzales-Andres et al. (1996) studying the izoenzyme 
variation of these species, reached another conclusion: only 3 species-E colona, 
E. crus-galli and E. oryzicola— were clearly defined. Echinochloa hispidula and 
E oryzoides clustered together, and the variability between the populations of 
E hispidula was higher than the variability between both taxa. Therefore, the 
authors suggested that the most appropriate classification would be E hispidula 
as a subspecies of £. oryzoides. Asins et al. (1999) studying the morphologic and 
isozyme patterns of variation of the same species, found only E. colona and E 
oryzicola to be distinct from both morphologic and molecular point of view. 
Based on their morphology, the populations of E. crus-galli, E. hispidula and E 
oryzoides congruently clustered together within each species. However, based 
on the isozyme variation, there was a considerable overlapping between spe- 
cies suggesting a high degree of genetic variation. Consequently, Asins et al. 
(1999) advocated that E. hispidula and E oryzoides as infraspecific taxa of E. 
crus-galli would be the most appropriate classification. We consider that in 
order to reach a final conclusion more populations worldwide should be 
analysed using combined molecular (such as RAPD, AFLP, ISSR and DNA fin- 
gerprints) and morphologic methods, and the data resolved m a cladistic ap- 
proach involving all characters. The enormous variation observed makes the 
boundaries between these taxa rather uncertain and in order to avoid an arbi- 
trary classification, we maintained each taxa at specific level. In Echinochloa 
the amplitude of morphologic variation can serve as a basis for taxa differen- 

I owe many thanks to Jose Luis Carretero for the useful discussions and the 
material from Spain. I am also very grateful to Julio Iranzo who kindly made 
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Edward E.Terrell Richard P.Wunderlin 

Research Collaborator Institute for Systematic Botany 

tment of Systematic Biology-Botany Department of Biology 

Hedyotideae. These data support the taxonomic recognition of V 

Seed morphology in Houstonia L. and related genera in the tribe Hedyotideae 
Cham. & Schltdl. ex DC. (Rubiaceae) has contributed valuable information, es- 
pecially when combined with data from chromosome number and pollen struc- 
ture (Terrell et al. i986; Terrell 1996). More recent taxonomic studies have con- 
tinued to emphasize the importance of seed data (Terrell 2001a, 2001b, 2001c). 

This study investigates the morphology of the seeds and fruits in selected 
genera in the tribe Spermacoceae Bercht. & J. Presl. We contrast this informa- 
tion with previously acquired seed data for the Hedyotideae. The two tribes in- 
clude some superficially similar genera and species (e.g. Oldenlandia L. and 
Spermacoce L.), but structure and development of their fruits and seeds are fun- 
damentally different and deserve further study. The Hedyotideae have few to 
many small seeds per locule, nearly always in subglobose biloculate capsules. 
In contrast, the fruits of Spermacoceae have one large seed per locule and are 
generally more structurally complex and vary considerably among the genera. 
Bremer and Manen (2000) expanded the tribe Spermacoceae to include the 
Hedyotideae; however, our data from seed and fruit morphology in five genera 
do not appear to support this inclusion. 

Verdcourt (1958) proposed a classification of the Rubiaceae in which he 
recognized three subfamilies. The Rubioideae, the largest subfamily contained 
17 tribes including the Hedyotideae and Spermacoceae. Bremekamp (1966) 

modified Verdcourts system and recognized eight subfamilies in the family and 
19 tribes in the Rubioideae (including the Hedyotideae and Spermacoceae). 
Verdcourt (1976), still recognizing his original three subfamilies, reduced the 
number of tribes in the Rubioideae to 12 and maintamed Hedyotideae and 
Spermacoceae. Robbrecht (1988, 1993) recognized four subfamilies and 19 tribes 
in subfamily Rubioideae. The latter included Hedyotideae and Spermacoceae. 

Scanning electron microscopy was employed in this study of seeds and fruits. 
The investigation was carried out at the Biology Department, University of 
South Florida, using a JEOL JSM-35 microscope operating at 15 kilovolts. Ten 
species in five genera were examined from herbarium material at the University 
of South Florida (USF) (Table 1). The species examined were those most readily 
available and representative of species native to tropical America, especially of 
the Caribbean basin. Tables 1 and 2 provide nomenclatural author's names. 

antropical tribe that may consist of as many as 19 gen- 
era and 450 species. Its generic delimitations are a matter of dispute. Eleven 
currently recognized genera native to the New World are summarized in Table 
2. Morphological characters of the seeds and fruits of five genera of Spermacoceae 
native to the Americas are described as follows: 

Diodia: Ovary 2(3-4)-locular; ovules solitary in each locule, attached to 
the middle of the septum. Fruit with 2(3-4) indehiscent mericarps, these fleshy 
or dry capsules. Seed enclosed within the hardened mericarp (accessible only 
by dissection), oblong, somewhat compressed, the dorsal surface convex, the 
ventral surface with a slightly sunken, central, longitudinal groove (see com- 
ments in Discussion), the testa finely reticulate (Fig. lA-D). 

Ernodea: Ovary 2-locular; ovules solitary in each locule, attached to the 
middle of the septum. Fruit fleshy, oblong, becoming hardened in drying, not 
easily separated into indehiscent mericarps (accessible only by dissection). 
Seeds oblong, somewhat compressed, the dorsal surface convex, the ventral sur- 
face with a slightly sunken, central, longitudinal groove, the testa finely reticu- 
late (Fig. 3C,D) 

Mitracarpus: Ovary 2(3-4)-locular, ovules solitary m each locule, attached 
near the middle of the septum. Fruit a thin-walled circumscissile capsule, the 
distal portion falling away with the calyx limb, the septum usually persistent. 
Seed oblong to globose, the dorsal surface convex, the ventral face divided into 
4 distinct areas by an x-shaped groove, the testa finely reticulate (Fig. 3A,B). 

Richardia: Ovary (2-)3-4(-6)-locular, ovules solitary m each locule, at- 
tached near the middle of the septum. Fruit dehiscing into dry mericarps, the 
ventral mericarp face with either a medial keel or narrow groove. Seed filling 


Source/Voucher Information 

Florida. Collier Co.: Lakela 27450 (USF) 

Florida. L 

is Gomes Florida. Hillsborough Co.: Gregory 7 (USF) 

a (Cham.S Florida. Escambia Co:.Wilhelm & Burkhalter 


Diodia L. 5 spp. North America, Mexico, Centra 

South America; Bacigalupo & C 

Emmeorhiza Pohl ex En( 


Tus Zucc. ca. 30 spp. Mexico, Central America, and South A 

■a L. 1 5 spp.; Mexico, Central America, South Amer 

nia E.L.Cabral & Brazil;Cabral & Bacigalupo 2001. 

the mericarp tightly (sect. Richardia) or sHghtly smaller than the mericarp (sect. 
Asterophyton (KSchum.) W.H.Lewis & R.L.Oliv.), oblong or broadly ovate, the 
dorsal surface convex, the ventral surface with a slightly sunken, central, lon- 
gitudinal groove, the testa finely reticulate (Fig. 3E,F;4A-F)). 

Spermacoce: Ovary usually 2-locular, ovule solitary in each locule. Fruit 

dehiscing into 2 mericarps, both opening to release the seed (sect. Borrcria ( 
Mey.) Verde.) or one mericarp opening to release the seed while the other : 
mains closed by a fragile, easily removable septum (sect. Spermacoce). Seeds i 
ellipsoidal, ovoid, or oblong, the dorsal surface convex, the ventral surface wi 
a slightly sunken, longitudinal groove extending to the ends, this sometirr 
with small elaiosomes, the testa reticulate (Fig. 1E,F; 2A-F). 


Some workers continue to treat Spermacoce in the 
a section of Spermacoce based on fruit deh 

ense, while others 
ler as a separate genus or as 
(capsule with both carpels 
:ds in Borrcria or a capsule with 1 carpel opening to 
laining closed). Based on our study of seed morphol- 
ppear to warrant recognition as a genus. Recently, 

(E-F). Scale bars are 

Ga lianthe was resurrected as a segregate genus from Borrcria (Cabral 1991) and 
Scandentia was described (Cabral 6a: Bacigalupo 2001). Both Galianthe and 
Scandentia are distinguished from Borrcria primarily in habit with more or 
less lax, thyrsoid, apical inflorescences in contrast to the congested, apical and/ 
or axially glomeruliform inflorescences of Borrcria s. str Emmeorhiza, a mo- 
notypic South American genus closely related to Borrcria, Galianthe, and 
Scandentia has an umbelliform inflorescence. The seeds of Spermacoce 
assurgens, 5. prostrata, and 5. vcrticillata, which would be placed in Borrcria 
on the basis of fruit dehiscence, do not differ significantly from that of S. 
tetraquetra traditionally placed in Spcrmacoce (Fig. 1E,F; 2A-F). Further work 
on the Spcrmacoce complex is needed and is underway by various other work- 
ers (e.g., E.L. Cabral, N.M. Bacigalupo, and S. Dessein). 

Diodxa also presents similar problems in circumscription. Hemidiodia, a 
monotypic genus with 2 indehiscent mericarps and long considered closely 
related to Diodia, was transferred to Borrcria subg. Dasycephala (DC.) 

Bacigalupo & E.L.Cabral by Bacigalupo and Cabral (1996). Another study by 
Bacigalupo and Cabral (1999) defined Diodia as comprised of only five Ameri- 
can species; the other species previously referred to the genus are transferred to 
Galianthe or Borreria. These workers exclude Diodia teres and related species 
from Diodia s. str. The placement of these species is still under study by 
Bacigalupo and Cabral. The seed morphology is markedly different between 
the two southeastern U.S. species, Diodia virginiana and Diodia teres, exam- 

ined in this study (Fig. lA-D). The seed of D. vi rginiana is oblong with an equally 
rounded apex and base. The testa has more or less isodiametric cells. In con- 
trast, that of D. teres is oblong with an apical projection and a truncate base. On 
the ventral surface, the base and sides are somewhat enrolled onto the longitu- 
dinal groove. The cells of the testa surface are elongate (2-3 times as long as 
wide). Further study by other workers may prove D. teres and related species to 
be distinct at the generic or subgeneric level. 

The seeds of the four Richardia species examined are fairly uniform, differ 
only slightly in size and shape, and the genus appears to be a natural assemblage. 

Ernodea seeds and fruits are most similar to those of Diodia virginiana. 

The seeds of Mitracarpus are unique among those examined in this study. 
The distinct x-shaped groove and the circumscissile dehiscent capsule, both 
features unique in the tribe, suggest that it may be a distinct subtribe. 

The Hedyotideae is much simpler in seed morphology than is the 
Spermacoceae, but has much more variation in seed shape and size. The seeds 
are typically in subglobose capsules, each bearing few to many seeds borne on 
complexly structured placentas. The variation shown by the seeds of the vari- 
ous taxa is very great and each genus, subgenus, or section often has its own 
particular seed shape. The following are some examples of seed diversity found 
in the Hedyotideae: (1) Houstonia L. has crateriform seeds, each with a ventral 
subglobose cavity (subg. Houstonia) or with a hilar ridge in a ventral shallow 
depression (subg. Chamisme Raf. sect. Amphiotis (DC.) Terrell) or with a hilar 
ridge in a ventral boat- or cup-shaped depression and with other complex struc- 
ture (subg. Chamisme sect. Ericotis (Terrell) Terrell) (Terrell et al. 1986, Terrell 
1996); (2) Oldenlandia L. typically has very small trigonous seeds, 50-100 or 
more per capsule (Terrell 1996); (3) the Hedyotisfruticosa L. group has dorsiven- 
trally compressed seeds with a short to long raised hilar ridge (Terrell 1996); 
and (4) Stenotis Terrell and Stenaria (Raf.) Terrell have eUipsoid seeds with a 
central punctiform hilum (Terrell 2001a, 2001b). 

The Spermacoceae, in contrast, have one seed per locule, and the locules 
develop into varied mericarps. The complexity occurs in the mericarps as well 
as m the seeds; the seeds examined here do not seem to exhibit much variation, 
although the Mitracarpus seed is an exception. In the Spermacoceae, some 
mericarps are hardened and indehiscent (e.g., Diodia), others are fleshy 
(Ernodea), and some open widely (e.g., Spermacoce). Each genus often has its 
characteristic kind of mericarp, and its structure is important in classification 
(e.g., Richardia, Fig. 4A,B). The mericarps may be more important in classifica- 
tion than the seeds (e.g., our figures show the similarity of seeds among the vari- 
ous genera). The presence of such varied mericarps contrasts with the mor- 
phology of the Hedyotideae, which have no such seed covering, just the bare 
seeds attached to a multi-branched placenta. 

An additional seed feature not found in the Hedyotideae is the ventral 

(adaxial) longitudinal groove which resembles a raphe. This structure was des- 
ignated as a strophiole by Kirkbride (1979), Bacigalupo and Cabral (1996), and 
other South American botanists. The structure and terminology of this appar- 
ent raphe needs further study It occurs on all seeds of Spermacoceae that we 
have examined. The Hedyotideae differ in having a so-called hilum, a puncti- 
form scar or a more elongated ridge-top scar on the seed. 

Although our evidence from five selected genera represents a limited sur- 
vey of the possibly 19 genera of Spermacoceae, it is sufficient to lead us to seri- 
ously question the union of the Spermacoceae and Hedyotideae. 

We thank Betty Loraamm, Electron Microscope Manager, and Clinton J. Dawes, 
Biology Department, University of South Florida for providing SEM data and 

advice on SEM mattersjoseph Kirkbnde and John Wiersema contributed valu- 
able reviews. Piero Delprete provided early taxonomic advice. Paul Peterson and 
Pedro Acevedo contributed the Spanish translation, 


Anderson, W.R., 1972. A monograph of the genus Crusea (Rubiaceae). Mem. New York Bot. 

Card. 22(4):1 -128. 
Bacigalupo, N.M. and E.L. Cabral. 1996. Infrageneric classification of Borreria (Rubiaceae- 

Spermacoceae) on the basis of American species. Opera Bot. Belg. 7:297-308. 
Bacigalupo, N.M. and E.L. Cabral. 1999. Revision de las especies Americanas del genero 

Diodia (Rubiaceae, Spermacoceae). Darwiniana 37:1 53-1 65. 
Bremekamp, C.E.B. 1 966. Remarks on the position, the delimitation and the subdivision of 

Bremer, B. and J.-F. Manen. 2000. Phylogeny and classification of the subfamily Rubioideae 

(Rubiaceae). PI. Syst. Evol. 225:43-72. 
Cabral, E.L. 1 991 . Rehabilitacion del genero Qalianthe (Rubiaceae). Bol. Soc. Argent. Bot. 

Cabral, E.L. and N.M. Bacigalupo. 200] . Scandentia, nuevo genero de Rubiaceae- 

Spermacoceae. Darwiniana 39:29-41 . 
KiRKBRiDE, J.H. 1 979. Revision of the genus Psyllocarpus (Rubiaceae). Smithsonian Contrib. 

Lewis, W.H. and R.L. Oliver. 1 974. Revision of Richardia (Rubiaceae). Brittonia 26:271 -301 . 
Nlgron-Ortiz, V, and R.J. Hickey. 1 997.The genus Ernodea (Rubiaceae) in the Caribbean Ba- 

sin:2. Morphological analyses and systematics.Syst. Bot. 21:445-458. 
RoBBRECHT,E.1988.Tropical woody Rubiaceae. Appendix 4:235-250. NationaiePlantentuin 

RoBBRECHT, E. 1 993. Supplement to the 1 988 outline of the classification of the Rubiaceae. 
Index to genera. Opera Bot. Belg. 6: 1 73-1 96. 

Terrell, E.E. 1 996. Revision of Houstonio (Rubiaceae-Hedyotideae).Syst. Bot. Monogr.48:l -1 1 8. 

Terrell, E.E. 2001 a. Taxonomy of Stenaria (Rubiaceae-Hedyotideae), a new genus includ- 
ing Hedyotis nigncans.Sida 19:591-614. 

Terrell, E.E.2001b.5tenot/5 (Rubiaceae), a new segregate genus from Baja California, Mexico. 
Sida 19:899-911. 

Terrell, E.E. 2001c. Taxonomic review of Houstonia acerosa and /-/. pa/mer/, with notes on 
Hedyotis and Oldenlandia (Rubiaceae). Sida 19:913-922. 

Terrell, E.E., W.H. Lewis, H. Robinson, and J.W. Nowicke. 1 986. Phylogenetic implications of di- 
verse seed types, chromosome numbers, and pollen morphology in Houstonia 
(Rubiaceae). Amer. J. Bot. 73:1 03-1 1 5. 

Vrrdcourt, B. 1 958. Remarks on the classification of the Rubiaceae. Bull. Jard. Bot. Etat 28:209- 

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Issimus var. subtropicanus are 


lology for 

e landscape 

Carphephorus odoratissimus znd C. 
ired. Carphephorus odoratissimus var. 
1 status is quantified. It occurs in a fire- 
, where it is a conspicuous, perennial, 
sular Florida. 


de Flor 

:te y compara la morfologia, disi 







Carphephorus sensu lat. (Asteraceae-Eupatorieae-Liatrinae) comprises seven 
species (Correa & Wilbur 1969), and one recently recognized variety ( Wunderlm 
& Hansen 2001) of Carphephorus odoratissimus, all of which are endemic to the 
southeastern United States. Six of these taxa occur in Florida, of which five [C. 
carnosus (Small) C.W.James, C. corymhosus (Nutt.) Torr & A. Gray, C paniculatus 
(J.F. Gmelm) H. Hebert, C odoratissimus (J.F. Gmelm) H. Hebert, and C odoratissimus 
var. subtropicanus (DeLaney, N. Bissett & Weidenhamer) Wunderlin & B.F 
Hansen] are known from peninsular Florida (DeLaney et al. 1999; Wunderlin 
1982; Wunderlin et al. 1996; Wunderlin 1998; Wunderlin & Hansen 2001). 

The circumscription of Carphephorus has been much debated (Hebert 1968; 
Correa & Wilbur 1969; Cronquist 1980; Cox 1998). Some authors (Hebert 1968; 
Correa & Wilbur 1969) include Trilisa and Litrisa within Carphephorus, while 
others recognize them as distinct genera (Cassini 1823, 1828; Robinson 1913; 
Gaiser 1954; James 1958; King & Robinson 1987; Bremer 1994). Evidence pre- 
sented by both Hebert (1968) and Correa & Wilbur (1969) support the inclu- 
sion of Trilisa within Carphephorus. Most recent floristic works have adopted 
this broad definition of Carphephorus (Cronquist 1980; Godfrey & Wooten 1981; 
Long & Lakela 1971; Wunderhn 1982; Wunderlm et al. 1996; Wunderlm 1998). 

In the most recent taxonomic study of Carphephorus sensu lato (Correa & 
Wilbur 1969), Irilisa and LUrisa are included withm Carphephorus. Since 
Correa and Wilbur (1969), no new taxa in Carphephorus were recognized until 
DeLaney et al. (1999), described C. suhtropicanus DeLaney, N. Bissett, & 
Weidenhamer Following DeLaney et al. (1999), Wunderlin and Hansen (2001) 
treated C. suhtropicanus at varietal rank within C odoratissimus. 

Based upon field observations in the fall of 1995, there seemed to be two 
entities of C odoratissimus within peninsular Florida. Carphephorus specimens 
from f latwoods and prairie habitats of south-central peninsular Florida seemed 
somewhat morphologically different from C odoratissimus, as known to us from 
previous fieldwork in southern Alabama, southern Mississippi, southern Geor- 
gia, and the Florida panhandle. Subsequent field investigations and critical taxo- 
nomic study of Carphephorus conducted from 1995-1997 led us to conclude 
that the Carphephorus entity in south-central peninsular Florida was worthy 
of recognition. Since that time, our on-going (1998-2001) field investigations 
and taxonomic studies of Florida Carphephorus, particularly in the region of 
range overlap for C. odoratissimus and C. suhtropicanus in central Florida, lead 
us to conclude that C suhtropicanus should be recognized as a variety of C 
odoratissimus. In this paper we present ecological, geographical, morphologi- 
cal, and phenological evidence to corroborate Wunderlin & Hansen's (2001) 
recognition of C. suhtropicanus at varietal rank. All our data on morphological 
characters are derived from field sampling of randomly chosen individuals in 
each of ten populations of these varieties scattered in central Florida. Both en- 
tities have mainly separate geographic ranges but intergrade where they are 
contiguous or overlapping; both exhibit slightly divergent but overlapping flow- 
ering phenology; and despite quantitative differences in mean values of veg- 
etative morphology and numbers of flowers per head, they lack differences in 
achene and floral morphology; all these features are indicative of varietal rather 
than species level recognition. Phenotypically, C. odoratissimus var. 
suhtropicanus is distinguished from C. odoratissimus var. odoratissimus by its 
lack of a coumarin (vanilla-like odor), shorter and narrower basal leaves, 
strongly clasping and entire stem leaves, broader and more diffuse inflorescence, 
and more flowers per head. An illustration of Carphephorus odoratissimus var. 
suhtropicanus is provided in Figure 1. 

Carphephorus odoratissim 
most notably in leaf cha 
(1933:1337) noted that "extreme forms" of Trilisa odoratissima (= C 
odoratissimus) might represent two entities, "one with a strong coumarin odoi 
broad, clasping, coarsely toothed upper leaf -blades and slightly viscid involu 
cres Ivar. odoratissimus], the other with only a faint coumarin odor, narrow entin 

lnflorescence,C. Single head. 

upper leaf blades and very viscid involucres. Small never published the latter 
as a distinct taxon, and subsequent floristic v^/orkers have treated C. 
odoratissimus as a single entity. A revision of the genus (Correa & Wilbur 1969) 
also failed to account for the two as separate entities. According to Correa and 
Wilbur (1969), the type of C odoratissimus is "presumably from the outer Coastal 
Plain of South Carolina ... which was the site of most of the species mentioned 
in Thomas Walter's Flora Caroliniana." Although this type has not been seen 
by the authors, the collection location is outside the range of C odoratissimus 
Y^r. subtropicanus. 

There is considerable character overlap between the two C odoratissimus 
varieties (Table 1). The presence of numerous quantitative differences in inflores- 
cence and leaf morphology are the most divergent character states. However, 
the lack of major differences in achene and floral morphology indicates differ- 
entiation at the varietal level. One useful character is the average number of 
inflorescences per clump in well-developed plants. Most plants of C odoratissimus 
var. suhtropicanus have one or two flowering stems per clump (mean=1.6, 
std.=0.8), whereas in central Florida var. odoratissimus averages over twice as 
many inflorescences per clump (mean=4.1, std.=1.9). Several characters of the 
basal and stem leaves have different mean character states. Although there is 
wide variation in leaf size and shape between plants within a population in 
either entity and between lower and upper leaves on individuals, comparison 
of leaves in the same relative position on plants of the two varieties reveals con- 
sistent differences. The basal leaves of C. odoratissimus var. suhtropicanus are 
both shorter (mean=ll cm) and narrower (mean=3.0 cm) than those of var. 
odoratissimus (mean length=19.8 cm; mean width=5.6 cm). The stem leaves of 
var. odoratissimus are broadly elliptic, and although the base of the leaf clasps 
the stem, the apex of the leaf is often divergent or flared away from the stem. 
The margins of the stem leaves in var. odoratissimus are often coarsely shal- 
lowly toothed. In contrast, the stem leaves of var. suhtropicanus are narrowly 
elliptic, clasp the stem for their entire length, and typically have entire margins. 

Other characters that differentiate the two varieties involve the size and 
branching pattern of the inflorescence. The inflorescence of var. suhtropicanus 
is much broader than that of var. odoratissimus, and on well-developed mature 
plants it is almost always as broad as tall or broader than tall (height:width 
ratio mostly from 0.5:1 to 1:1). This difference is mostly attributable to the angle 
of branching of the primary inflorescence branches from the main axis. The 
inflorescence branches of var. suhtropicanus diverge from the main axis at an 
angle of 30°to 45°and are often arcuate, resulting m a rather open and diffuse 
inflorescence. In contrast, the main inflorescence branches of var. odoratissimus 
diverge from the main axis at an angle of 10° to 20° resulting in a narrower, 
taller, inflorescence that is typically taller than broad (height:width ratio from 
1.5:1 to 3:1). This open, diffuse inflorescence, more diffuse than any other species 

Achene length 

of Carphephorus, is a reliable field character but is often difficult to discern m 
dried herbarium specimens or plants with an immature inflorescence. 

Differences in inflorescence, floral, and achene characters between the two 
varieties are minor There is considerable variation even within an individual 
in the number of florets per head, size of heads, and size of floral parts. 
Carphephorus odoratissimus var suhtropicanus tends to have more florets per 
head (10-13) than is typical for var odoratissimus (7-10). However, this could be 
attributable to the fact that most var suhtropicanus inflorescences consist of 
only 50-75 heads, in contrast to the hundreds of heads present in some inflo- 
rescences of var odoratissimus] therefore a larger proportion of heads fully de- 
velop all of their potential florets. We can not find any consistent differences in 
achene length (1.9-2.5 mm) (-2.8 mm) for C odoratissimus var suhtropicanus in 
comparison to var odoratissimus (2.0-2.5 mm) or other floral characters be- 
tween the two varieties. DeLaney et al. (1999) also noted overlap in achene length 
and number of florets per capitulum. 

The following key can be used to distinguish the two varieties: 

Carphephorus odoratissimus var. suhtropicanus is endemic to central and south- 
ern peninsular Florida, where it has been collected from 21 counties (Fig. 2), an 
area encompassing approximately 50,000 sq. km (20,000 sq. mi). It is a con- 
spicuous, autumn-flowering forb of burned pine savanna-f latwoods/dry prai- 
rie landscape throughout south-central Florida and is quite frequent in High- 
lands, Polk, and Okeechobee counties. In the western part of the peninsula, its 
range extends somewhat further north as compared to the eastern part of its 
peninsular range. In south Florida, C odoratissi mus var. suhtropicanus is rather 
infrequent in Collier County, having been collected most frequently in scrubby 
flatwoods near Immokalee and historically from scrubby flatwoods at Marco 
Island, the only offshore island location presently known. In southeastern 
Florida it has been collected from flatwoods and prairies in St. Lucie, Martin, 
and northern Palm Beach counties. It is not known from the Everglades region 
of south Florida, presumably due to the absence of acidic pine flatwoods and 
prairies. There is an historical collection from a sandy low pineland m north- 
ern Miami-Dade County, presumably from sandy flatwoods that once occurred 
on the Atlantic Coastal Ridge. 

Carphephorus odoratissimus var. odoratissimus, in contrast, is found 
throughout most of the southeastern United States coastal plain, from south- 
eastern North Carolina south and west to southeastern Louisiana (Correa & 
Wilburl969). It is quite common in northern Florida and m southern Georgia 
and Alabama, where it is typically found in upland longleaf pine (Pinus 
palustris Mill.) dominated woodlands and savannas. Throughout its range var. 
odoratissimus tends to be associated with sandy surface soils that are better 
drained than those for var. suhtropicanus. Carphephorus odoratissimus var. 
odoratissimus does not occur in South Florida slash pme (Pinus elliottii Engelm. 
var. densa Little & K.W. Dorman) dominated flatwoods. However, ranges of both 
varieties overlap in central Florida. Here Carphephorus odoratissimus var. 
odoratissimus seems to reach its southern limit in ecotonal habitats on the lower 
sideslopes of sandy ridges, both associated with the Central Ridges in extreme 
northern Polk and Osceola counties and with coastal sand ridges in Brevard, 
Volusia, Citrus, and Hillsborough counties. 

Carphephorusodoratissimus var. sukropicanus is characteristic of the pine- 
savanna flatwoods/dry prairie landscape of the Okeechobee, Osceola, and 
Desoto Plains (as defined by Cooke 1939; Schmidt 1997) in south-central penin- 
sular Florida. Pine savanna-f latwoods and dry prairie occupy nearly level, in- 
ter-drainage f latlands on acidic, low nutrient, poorly drained sandy to sandy 
clay alfisol or spodosol soils (Abrahamson & Hartnett 1990; Bridges 1997; Orzell 
& Bridges 1997). Pine savanna-flatwoods are fire-maintained, open-canopied 
pine stands dominated in south-central Florida either by Pinus palustris or P. 

fiin county is shown when species is restricted to a single or few locations in a county. For Collier County 

stars, one for mainland and one for the only known island population oi 

county when species is widely distributed throughout county. Distribution ol 

'5 extends beyond figure north of Florida to North Carolina and west to Louisiana. 

dliottii var. densa, with Aristida heyrichiana Trin. & Rupr., Serenoa repens (W. 
Bartram) Small, and low-growing Quercus minima (Sarg.) Small typically domi- 
nant in the ground cover as well as numerous regionally endemic species that 
vary f loristically between geographic regions (Orzell & Bridges 1997). Dry prai- 
ries are similar in ground cover composition to pine-savanna f latwoods but are 
naturally treeless. Historically, dry prairies had annual or biennial naturally 
occurrmg fires (Harper 1921, 1927; Orzell & Bridges 1999) and are the highest 
fire-frequented community type in central Florida (Orzell & Bridges 1999). 
Associated species that are endemic or near-endemic to peninsular Florida in- 
clude Asimina reticulata Shuttlew. ex Chapm., Polygala setacea Michx., Liatris 
tenuijolia Nutt. var quadriflora Chapm., Rhexia nuttallii C.W.James, Bejaria 
racemosa Vent., Andropogon hrachystachyus Chapm., Andropogon ternarius 
Michx. var cahanisii (Hack.) Fern. & Griscom, Gymnopogon chapmanianus 
Hitchc, Phoehanthus grandiflorus (Torr. & A. Gray) S.F Blake, and Polygala 
rugelii Shuttlew. ex Chapm. 

Carphephorusodoratissimus var subtropicanus can be sympatric with any 

of the four other Carphephorus species occurring m central Florida. It most of- 
ten occurs with C corym hosus at drier sites, C panicu latus at mesic sites, and in 
very rare cases with C carnosus at wet-mesic sites. Carphephorus odoratissimus 
var. subtropicanus is sympatric with var. odoratissimus at its northern (Polk 
and Osceola counties) and northwestern (Citrus and Hillsborough counties) 
range limits. Plants intermediate between the two are occasionally found in 
ecotonal and disturbed sites in central Florida where the boundaries of the two 
overlap. Where the two varieties are sympatric at a site in Osceola County, var. 
subtropicanus grows in dry-mesic pooriy drained pine flatwoods, whereas var. 
odoratissimus is found on a somewhat poorly drained slight sandy rise between 
the pine flatwoods and a wetland depression. 

At a disturbed site in Orange County specimens intermediate in characteris- 
tics (leaf morphology, inflorescence, and coumarin odor) between var. 
odoratissimus and var. subtropicanus were observed with both flowering in late 
September 1999. The co-habitation and overlapping flowering at the site of both 
varieties indicates that ecological separation of the two varieties is incomplete. 
In 1997 at an Osceola County site, C. odoratissimus var. odoratissimus flow- 
ered from early September till late October, and C. odoratissimus var. 
subtropicanus flowered from late October into November. Elsewhere in central 
Florida, var. odoratissimus generally flowers from early to mid September into 
late-October and even earlier in north-central Florida (August thru mid-Sep- 
tember), whereas var. subtropicanus flowers primarily from early to mid Octo- 
ber thru early to mid November, with flowering having been noted in Septem- 
ber at several locations. Plants flowering in early or late September were also 
noted by DeLaney et al. (1999). Furthermore, atypical flowering dates for var. 
subtropicanus (usually the result of burning followed by available soil mois- 
ture) have been observed. Flowering plants were collected on 2 April 1998 fol- 
lowing a 19 August 1997 burn in Osceola County. 

Citations here are abbreviated, however full label data is 
from the authors, including detailed habitat and locati 
Orzell & Bridges collections. 


ig 240ctl945 Brass J56J3(ARCH) dry mtbiL snndy treelebi,tlatwoodb/dr> prairie AvonPaik 
)rceRinge 12 Nov 1997 Orzdl& Budges 25261 {ARCH BRIT FLAS FSU MO NY TEX USF) 
orough Co Flatwoods Park ca 7 mi E of Lutz 2 Oct 1994 Wunderhn W558 (USF) Lee Co Six 
:ypiess Preserve 17 Oct 1997 Bradley 791 (FTG) Manatee Co ca 1 mi N of Myakka River State 
L7 Sep 1978 Dodson 4798 (USF) Martin Co W of canal S of Salerno jct off US 1 29 Sepl962 
I 25383 (FI AS USF) Okeechobee Co US 68 15 mi N of Okeechobee 19 Oct 1969 McCart llNl 
. USF) Osceola Co CI 12 air mi S ol Kissimmee 11 Oct 1997 Orzell & Bridges 25237 (FLAS 

^SjCitrusCo Inverness 10 Nov 1987 MawJunne) 
\^) Dixie Co NoiOldtown 18 Aug 1937 Arnold 
L8 Aug 1963 Oeagei 269 (FLAS) Escambia Co E 
aglerCo Palm Coist 4 Oct 1978 Tabbsn (USF) 
■.cmkda &Ruhaidson6719(VS¥) Nof Andalusia 
Co E of Trenton 10 May 1940 Martin DeVall & 

.1 1 inetdtLfcon 28 Sep 1981 Hansen 8906 iU^F) 
4 U USF ) Lake Co Oc d 1 National Foiest 3 Sep 
1 Ninon i\ Fiitbt 29 Oct 1983 Parkei 1790 (USF) 

,nCo SiherRuer State Park 10Sepl990 Buckner 

ISepl979 Hansen &RKhard-^on 6240 iUSY) 10 mi S of Oakland 7Augl958 Krai7745(USF) 
ilaCo W of Intercession Qt) 14 Aug 1981 Hni nf-^RoJ^mson 8478 (USF) Polk Co EoiHaines 
' Sep 1968 Conard s n (FLAS) i lU W il 1 O i 1 io4 ( maid s n (FLAS) Putnam Co Welaka 
, 1939 DeVai/sn (FLAS) Santa Rosa Co \ III i uk Pt 28Nox 1980 Wi;helm8280(USF) 
Holley26Sepl967 Snnlh WmI I ^S) Seminole ( o Oxicdo 7Sepl947 Sc ha llert 6762 (USF) 

West & Arnold s n (FLAS) Sumter Co W ildwc 
r Co US 27 W of Lafayette/Taylor Co line 12 

12 Aug 1956 Knobloch 1429 (FLAS) Union Co S of Raiford 22 May 1942 West & Arnold s n 
i) Volusia Co Near Ormond 5 Aug 1943 Butts s n (USF) Walton Co S of DeFuniak Springs 
1967 Smith 2006 (FLAS) Washington Co N of Cfnpley 29 Jul 1954 Ford 3682 (FLAS) Ala 

Covington Co F of Wmg 20 Oct 1969 Krai 38120 (USF) Escambia Co Riverview 27 Nov 
Wi/hem 8180 (USF) GEORGU Thomas Co N of Boston 3 Oct 1967 Mo (ton 2668 (USF) Mcln 
:o NEol Sipelo Island 17 Sep 1956 Diuican 20544 (USF) LOUISIANA Washington Par Nof 
ley 16 Sep 1983 TaylorcS Dutton 5618 (USE) MISSISSIPPI Harrison Co P'^ss Christian 22 Oct 
Dtmaree 36238 (USF) Jackson Co Hurley 27 Sep 1953 Dtmajtt 34398(USF) Pearl River Co I 

5epl966 Bradley/ 6^ Seari 3551 (USF) 

We thank Paul Ebersbach, Chief of the Environmental Flight at Avon Park Air 
Force Range (APAFR), for his support of scientific research on the military in- 
stallation; Rebecca Yahr for the illustration; Scott Penfield (APAFR) for provid- 
ing funding from the Rangelands Program at APAFR for the illustration; and J. 
Douglas Ripley formerly of the Environmental Planning Division of the U.S. 
Air Force in Washington DC, who secured funding for both the illustration 
and manuscript preparation. Fieldwork was facilitated by Sam Vanfiook, who 
conducted prescribed burns of sites for C odoratissimus var. suhtropicanus at 
APAFR, and Chris-Ann Kosel formerly of the TNC Disney Wilderness Preserve, 
who provided access to the preserve. We especially thank Guy Nesom of BRIT, 
Billie Turner of TEX, and Richard Wunderlm of USE for providing stimulating 
views and critical edits to earlier drafts of the manuscript. 


Abrahamson, W.G, and D.C. Hartneit. 1 990. Pine flatwoods and dry prairies. R.L Meyers and 

J.J. Ewel. eds. In: Ecosystems of Florida. University of Central Florida Press, Orlando. 

Bremer, K. 1 994. Asteraceae, cladistics, and classification. Timber Press: Portland, Oregon. 
Bridges, E.L 1997. Vegetation analysis of selected dry prairie/treeless flatwoods sites for 

GIS vegetation mapping on Avon Park AirForce Range,Florida. Unpublished reportto 

Avon Park Air Force Range, Florida. 65 p. + Appendices l-IV. 
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FCuvier, Diet. Sci. Nat. 26:223-235. 
Cassini, 1828,77/7/50. In RCuvier, Diet. Sci. Nat. 55:310-31 1. 
Cooke, CW. 1 939. Scenery of Florida, interpreted by a geologist. Florida G 

No. 17. 
Correa, M.D. and R.L.Wilbur. 1969. A revision of the genus Carphephorus ( 

Eupatorieae).J.EIisha Mitchell Sci. Soc. 85:79-91. 
Cox, RB. 1998. An overview of the subtribe Liatrinae (Eupatorieae: Compos 

tion Southeastern Biologist Bull. 45:121. 
Cronquist,A. 1980. Vascular flora of the southeastern United States. Volume 

University of North Carolina Press, Chapel Hill 
DeLaney, K.R., N. Bissett, and J.D. Weideniiamer. 1999. A new species of Carphephorus 

Gaiser, LO. 1 954. Studies in the Kuhniinae (Eupatorieae). J. Arnold Arbor. 35:87-1 33. 

States: Dicotyledons. University of Georgia Press, Athens. 
Harper, R.M.I 921. Geography of central Florida. Florida Geol.Surv. 1 3"' Annual Report. Pp. 

Harper, R.M. 1 927. Natural resources of south Florida. Florida Geo!. Surv. 1 8^^ Annual Re- 
port. Pp. 27-206, 

Hebert, H.J.C. 1968, Generic considerations concerning Carphephorus and Trillsa 
(Compositae).Rhodora 70:474-485. 

James, C.W. 1958, Generic considerations concerning Carphephorus, Trilisa, and Litnsa 
(Compositae). Rhodora 60:1 1 7-1 22, 

King, R. and H. Robinson. 1987, The genera of the Eupatorieae (Asteraceae), Monogr. Syst. 
Bot, Missouri Bot.Gard. 22:277-280. 

Long, R.W.and 0. Lakela. 1 971 .A flora of tropical Florida. University of Miami, Coral Gables. 

Orzell, S.L and E.L Bridges. 1997, Regional floristic diversity in peninsular Florida pine 
flatwoods and savannas. Abstr. of the Southeastern Flatwoods Ecosystem. First Ann. 
ConfSoc.EcoLRest.CoastalPlainChapter.May 16-17, 1997. Gainesville, FL 

Orzell, S.L. and E.L. Bridges. 1999. Dry prairie. U.S. Fish and Wildlife Service, eds. in: South 
Florida multi-species recovery plan, a species plan, an ecosystem approach. South- 
eastern Region, Atlanta. Pp. 3-279 thru 3-346. 

Robinson, B.L 1913. A key to the genera of the Compositae - Eupatorieae. Proc. Amer. 
Acad. Arts. 49:429-437. 

Schmidt, W. 1997. Geomorphology and physiography of Florida. A.F. Randazzo and D.S. 
Jones, eds. ln:The geology of Florida. University Press of Florida, Gainesville. Pp. 1 -1 2. 

SMALL,J.K.1933.Manualofthe southeastern Flora.Published by author,NewYork. 

Wunderlin, R.P 1982. Guide to the vascular plants of central Florida. University Press of 

WuNDERUN,R.PandB.F.Hansen.2001. Seven new combinations in the Florida flora.Novon 

Wunderlin, R.P., B.F. Hansen, and E.L, Bridges. 1 996. , 

partment of State, CD-ROM version of public. 

h ttp://www.plan tatlas. 
Wunderlin, R.P. 1 998. A guide to the vascular plants of Florida. University Presses of Florida, 

David G. Frodin. 2001 Guide to Standard Floras of the World. Second Edition. 

(ISBN 0-521-9077-0, hbk). Cambridge University Press, Cambridge, U.K. 
$240.00. 1100 pp, 7 1/2" X 10". 

in's book is arranged by major geographic area, i.e., North i 

ng the tidbits in the Guide are these about Easter Island. Th( 
:d in 1914, with others in 1920, 1922, 1951, 1958, and 1991. The 

of the tree from the island was by the Thor Heyerdahl 1955-1956 expedition. The island's sol 
More tidbits? Which U.S. state has the most plants? Leading the pack are California, \ 
The compiler of all this wealth, David Frodin, is a botanist at Kew Gardens in Eng 

botanical explorers who have worked over the centuries to discover and make known th 




Paula Power 

San Marcos National Fish Hatchery and Technology Center 
500 E.McCarty Lane 


Its, submersed leaves, and reproductive culms with associated leaves and inflorescences, then dried 
i weighed. At the end of the study, mean total biomass of plants grown in the fast site was 28.42 
^ vs 4.24 gdw for plants grown in the slow site. Resource allocation patterns differed among sites 
i two distinct phenotypes were apparent. One phenotype, associated with relatively higher cur- 
,t velocities, had higher net productivity a well-developed root system, and allocated proportion- 
/ more biomass to non-reproductive organs (49.2% gdw root biomass in fast site vs 24.8% gdw 
It biomass in slow site). A second phenotype, associated with relatively slower flowing water, had 
/er net productivity and allocated proportionally more biomass to reproductive organs (22.1% 
V reproductive culm in fast site vs. 65.0% gdw reproductive culm in slow site). Because of lower 
ductivity and the potential for loss from herbivory to a significant proportion of the plant in 

rante 7 meses. Las velocidades normales variaron entre 0.0-0.010 m/s (el sitio lento), 0.038-0.142 m/ 
s (en el sitio moderado), y 0.250-0.369 m/s (en el sitio rapido). Las plantas se cosecharon de cada 

reproductores con las hojas e inf lorescencias asociadas, luego se secaron y pesaron. A final del estudio, 
la biomasa total media de plantas del sitio rapido fue 28.42 gdw contra 4.24 gdw para plantas del 
sitio lento. Los modelos de recursos difirieron significativamente entre los diferentes sitios y fueron 
aparentes dos fenotipos claros eran. Un fenotipo asociado con velocidades actuales relativamente 

). Un segundo fenotipo, 

una productividad neta mas baja y 

Trade-offs between reproductive allocation and vegetative allocation with 
changes in environmental conditions have been reported by Harper and Ogden 
(1970); Hickman (1975); van Baalen, et al. (1990); Dunn and Sharitz (1991); 
Madsen (1991) and Neill (1993). Environmental factors associated with alloca- 
tion trade-offs include substrate, nutrient availability, and water depth 
(Idestam-Almquist & Kautsky 1995; Blanch et al. 1999; Lorenzen et al. 2001; 
Vretare et al. 2001). Little information is available on current velocity and its 
affect on allocation patterns in macrophytes although numerous studies have 
identified current velocity as an important factor influencing macrophyte dis- 
tribution, photosynthesis and growth (Nilssen 1987; Chambers et al. 1991; 
Madsen & Sondergaard 1983; Power 1996a). The work reported here was under- 
taken to document resource allocation patterns in the endangered Zizania 
texana Hitchc. over seven months in three different habitat types differing in 
current velocity and pH. 

Zizania texana is an endangered macrophyte endemic to the San Marcos 
River, Hays County Texas (U.S. Fish & Wildlife Service 1995). It commonly oc- 
curs midchannel in swiftly flowing water. Terrell et al. (1978) and Poole and 
Bowles (1999) provide thorough descriptions of the habitat for Z texana. Threats 
to the species include reduced spring flow from the source aquifer due to 
overpumping of ground water for human use, competition and herbivory by 
nonnative plant and animal species, and absence of sexual reproduction m the 
wild, along with other human impacts including alteration of historic hydro- 
logic patterns from dams located along the river and within the watershed, and 
recreational use (U.S. Fish & Wildlife Service 1995). 

Under flowing water conditions, Z. texana produces long ribbon-like, sub- 
mersed leaves and emergent culms, each with a terminal, wind pollinated in- 
florescence; however, sexual reproduction is nearly absent in the wild (U.S. Fish 
& Wildlife Service 1995). Historical accounts suggest this was not always the 
case. Photographs depicting fully developed inflorescences and verbal accounts 
suggest recruitment by seed occurred (Silveus 1933). More recent descriptions 
of floral structures and floral development by Emery and Guy (1979), and Power 
(1996b; 1997) indicate that sexual reproduction is most likely limited by envi- 
ronmental factors rather than cytological factors. It is not well understood why 
fully developed, emergent inflorescences are rare in the wild, although drifting 
mats of floating vegetation and herbivory play a role (Power 1996b, c). The spe- 

cies produces asexual clones (tillers) which form at the nodes of reproductive 
culms and it is assumed that tiller production is the primary mechanism for 
recruitment of new individuals in the wild (U.S. Fish & Wildlife Service 1995). 
Zizania texana has two distinct phenotypes under wild and cultured con- 
ditions. When grown in the wild, Z. texana is primarily submersed and tends 
to be a long lived perennial plant. Under cultivated conditions, Z. texana is pri- 
marily emergent, reproductive, and short lived (Terrell et al. 1978 and pers. 
observ). Information on plant response to environmental conditions, gains im- 
portance given the endangered status of the species and the need for restora- 
tion of the habitat as outlined in the Recovery Plan for the species (U.S. Fish & 
Wildlife Service 1995). Adequate restoration protocols cannot be developed with- 
out a thorough understanding of the response of Z. texana to a variety of con- 
ditions found in its historic range. 

This research project was carried out in Spring Lake on the Southwest Texas 
State University (SWT) campus. Spring Lake is an impoundment formed by a 
dam and spillway originally constructed across the San Marcos River in 1849. 
The dam and spillway are approximately 750 m downstream from the San 
Marcos Springs. 

Study plants were obtained by germinating captive grown seed in the lab. 
Seedlings were then transplanted to 15 cm plastic pots, lined with small plastic 
bags and filled with sediments collected from one location in the study site in 
Spring Lake. Pots were placed in an outdoor cement raceway on the SWT cam- 
pus and seedlings were allowed to grow for about 6 weeks. Water was supplied 
by an artesian well from the source aquifer for Spring Lake and the San Marcos 
River In March 1995, potted plants were transplanted into three sites in Spring 
Lake and one site in the outdoor raceway on the SWT campus. This was a nested 
design with plants nested in plots and plots nested in sites. There were three 
replicate plots at each site. Each replicate plot had 36 potted plants for a total of 
432 plants. All plants were protected from herbivores with 1 m^, floating 
exclosures constructed of polyvinyl chloride (PVC) pipe and 2.5 cm wire mesh. 
Initially, four plants were harvested to obtain baseline biomass values for the 
newly transplanted individuals. The study design called for harvesting four 

plants from each replica 


me separate occasion 

s. However, during the 

study period some plant 

s were lost 

to herbivory and son 

le plants were washed 

away by the current. As ; 

a result, pla 

nts were harvested f r 

om each replicate plot 

on six dates, May 3, Jun 

el, June 3C 

),July 27, September 

1, and October 16. At 

each harvest, plants weri 

2 selected at 

random. Sediment v 

.as washed from plant 

roots, then plants were i 

divided mt^ 

vegetative parts, re 

:productive parts, and 

roots! Vegetative parts w, 

sre defined 

as submersed leaves a 

md reproductive parts 

were defined as reproduc 

live culms ^ 

vvith associated leaves, inflorescences, seeds, 

and tillers. Plants were then dried at 65°C for at least 48 hours and weighed to 
the nearest 0.01 g. Data are reported as means plus-minus standard error. 

On nine occasions between May 1995 and October 1995 pH, water depth, 
and current velocity were recorded in each replicate plot, in each site. Current 
velocity was measured with a Marsh McBirney Model 201 portable water cur- 
rent meter and calculated as the average velocity at 20%, 60%, and 80% depth. 
The four sites differed in mean current velocity and were identified as follows: 
slow (raceway), no flow, moderate and fast. 

Plants in the no flow site in Spring Lake were lost to herbivory, probably by 
crawfish, prior to the second harvest and this site was dropped from the study 

Water depth, current velocity, and pFI at the remaining sites were recorded 
at the study sites eight times between May and October (Table 1). In the slow 
site, mean water depth was 0.71 m (±0.03) mean current velocity was 0.001 m/ 
5 (±0.004), and pH ranged from 7.50 to 7.68. In the moderate site, mean water 
depth was 0.88 m (±0.06), mean current velocity was 0.090 m/s (±0.048), and 
pH ranged from 7.23-7.26. In the fast site, mean water depth was 0.85 m (±0.07), 
mean current velocity was 0.290 m/s (±0.076) and pH ranged from 7.16-7.28. 

Plants in all study sites increased in size during the study period (Fig. 1). At 
each harvest, fast site plants had the greatest root, submersed leaf, and total net 
biomass, furthermore, data from the September harvest showed mean net total 
biomass of fast site plants was an order of magnitude greater than mean net 
total biomass of slow site plants (34.05 g vs. 3.66 g). 

Number of submersed leaves increased in all sites during the study period, 
however fast site plants produced six times as many submersed leaves as slow 
site plants. At the end of the study period, mean number of submersed leaves in 
fast site plants was 18.0 (± 1.414) while mean number of submersed leaves in 
slow site plants was 3.0 (± 0.211). 

Reproductive culms were present in every harvest between 1 June 1995 and 
16 October 1995. Culm number was greatest during the September harvest in 
plants grown in the fast site with 2.9 (± 0.380) culms/plant. For plants grown 
in the slow site, number of culms was greatest in the October harvests with 1.6 
(± 0.159) culms/plant. (Fig. 2). 

Biomass allocation to plant roots and reproductive culms varied through 
time and among sites (Fig. 3). The proportion of biomass allocated to roots in- 
creased from 27% in March at the beginning of the study to 49% in October in 
plants grown in the fast site. In contrast, the proportion of biomass allocated to 
roots decreased from 27% at the beginning for the study to 25% at the end of the 
study in the slow site. 

Plants in all sites produced similar numbers of culms, however biomass 
allocation varied greatly among sites. In October, culm biomass constituted over 

half of total plant biomass in slow site plants (65%; culm number = 1.6+0.159), 
m moderate site plants, 48% of total net biomass was culm biomass (culm num- 
ber = 1.7+0.129 and fast site plants allocated only 22% of total net biomass to 
reproductive culms (culm number = 2.25+1.591). 

The proportion of biomass allocated to submersed leaves was similar 
among sites even though the number of leaves varied among sites. In October, 
submersed leaf biomass was 29% of total biomass in the fast site (leaf number 
= 18+1.414), 15% of total biomass in the moderate site (leaf number = 5.3+1.862), 
and 28% of total biomass in the slow site (leaf number = 3+0.211). 

In this study, plants exhibited markedly different growth patterns among study 
sites. Net total biomass accumulated over the study period was an order of 
magnitude greater in plants grown in water flowing between 0.146-0.442 m/s 
compared with plants grown in water flowing between 0-0.01 m/s. Net biom- 
ass accumulation in individual plant organs (roots, submersed leaves, and re- 
productive culms) also was greater in fast flowing water compared with slow 
flowing water Submersed leaf biomass was 18 times greater in fast site plants 
compared with slow site plants and 4.5 times greater compared with moderate 
site plants. Zizania texana exhibited a similar response to flowing water in 
other studies (Power 1996a, b). 

Flowing water has been shown to influence macrophyte photosynthetic 
rates (Westlake 1967; Smith & Walker 1980; Madsen & Sondergaard 1983), dis- 
tribution (Fonseca & Kenworthy 1987; Nilssen 1987); and growth (Chambers 
et al. I99I). Plants occur in a range of current velocities and there is consider- 
able variability in optimum flow rates for macrophytes. Chambers et al. (1991) 
found an inverse relationship between biomass and current velocity between 
0.01-1.0 m/s. Nilssen (1987) found species richness reached a peak at about 0.3 

Fig. 1 . Growth by potted Zizania te 

7 months from two sites in Spring Lake (fast and moderate sites) and 
w site). After 7 months, plants grown in the fast site had the greatest 
;s compared with the other two sites. Points on graph represent the m 

I ^ S S " s s 

ersed leaves and reproductive culms from potted Zizania texana harveste 
<e and one at Southwest Texas State University. Plants grovt/n in fast flo 
s and reproductive culms compared with plants grown in slow flowing w 

ir.Vertical bars rep- 


s s I s s 

O Culms 
M Leaves 

3. Proportional allocation of plant nr 

m/s along a current gradient from 0.04-1.23 m/s, with some species growing 
in current velocities greater than 1 m/s. Many species occur in slower current 
velocities, while fewer species are specialized to withstand the forces of veloci- 
ties over 0.3 m/s. Apparently, Z texana with smooth, ribbon-like leaves is one 
of few species able to withstand velocities over 0.3 m/s. Poole and Bowles (1999) 
found wild Z. texana stands primarily in current velocities > 0.46 m/s and the 
current velocity tolerance range for Z texana exceeds 1.0 m/s; from this study 
however, it was not possible to identify a maximum or an optimum current 
velocity for growth. 

Zizania texana exhibits phenotypic variation in response to current ve- 
locity. One phenotype, associated with relatively higher current velocities, has 
higher net productivity, a well-developed root system, and allocates propor- 
tionally more biomass to nonreproductive organs. A second phenotype, associ- 
ated with relatively slower flowing water, has lower net productivity and allo- 
cates proportionally more biomass to reproductive organs. Plants face a tradeoff 
between the need for a well-developed root system to anchor plants in poten- 
tially unstable sediments, the need for submersed leaves with which to photo- 
synthesize, and the need for emergent stems for reproduction. In this study, the 
proportional allocation to nonreproductive organs (roots and submersed leaves) 
decreased with decreasing current velocity as the proportional allocation to 
reproductive organs increased. Apparently there is a trade-off in favor of sub- 
mersed organs in faster flowing water. 

Other factors directly and indirectly influenced by current velocity may 
play a role in net biomass accumulation in plant organs. They include herbivory 
deposition of debris and sediments on leaves interfering with metabolic pro- 
cesses, colonization of leaves with epiphytes, and the plant's inability to utilize 
HCO3' and its dependence on CO2 as an inorganic carbon source (unpublished 
data). Ribbon-like submersed leaves of Z texana are adapted to withstand the 
forces of flowing water and can reduce carbon limitation by exploiting flow- 
ing water habitat where boundary layer surrounding leaves and diffusion dis- 
tances for CO2 are reduced, and leaves are continually bathed with carbon rich 
water In contrast, in slower flowing water, photosynthesis by submersed leaves 
of Z. texana is carbon limited and few submersed leaves are produced. Emer- 
gent reproductive culms with associated emergent leaves most likely are not 
carbon limited because culms obtain CO2 from the atmosphere where CO2 is 
more readily available owing to the higher diffusion rate and current velocity 
in air relative to water (Madsen & Sand-Jensen, 1991; Denny 1993). 

Increased proportional allocation to emergent reproductive organs in rela- 
tively slower flowing water concurs with observations of captive grown Z. 
texana when grown in current velocity =0.015 m/s in which plants allocate a 
greater proportion of biomass to reproductive organs and typically set seed and 
senesce after one growing season. This is similar to Z. palustris and Z aquatica, 

annual species which commonly occur in shallow water along the margins of 
lakes and streams (Ferren & Good 1977; Weir and Dale I960). The importance 
of having leaves and flowers above the surface of the water may be due to CO2 
limitation in submersed leaves. In relatively slower flowing water, gas exchange 
and photosynthesis may be insufficient to support vegetative organs and re- 
sources shift to emergent organs where CO2 is plentiful. 

Herbivory is a factor contributing to sexual reproductive failure in Z. texana 
(U.S. Fish & Wildlife Service 1995; Power 1996c). Plants growing in microhabi- 
tats with relatively slow flowing water and potentially over 60% of biomass 
allocated to reproductive parts, are especially vulnerable to herbivory by wa- 
terfowl. Microhabitats with conditions which would trigger a low productiv- 
ity/high reproductive phenotype in the wild include back eddies and protected 
stream edges, emergent macrophy te beds, impoundments upstream from dams, 
and potentially, reduced springf lows due to drought and overpumping of the 
source aquifer (the Edwards Aquifer) for human use. These microhabitats would 
not be I 

This research was supported by a Section 6 grant from U.S. Fish and Wildlife 
Service and Texas Parks and Wildlife, 1 would like to thank Kathryn Kennedy 
Center for Plant Conservation, Robert Doyle, Baylor University for their sup- 
port and Francis Rose, Southwest Texas State University for permission to use 
outdoor raceways and Spring Lake. 


id K.F.WALKtR. 

1999.Growth and resource, 

allocation in respons 

rgent sedge I 

Solboschoenusmedianus. Ac 


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Power, R 1997. Moisture, seeds, and reproductive failure in Texas wildrice (Z/zan/atexono). 
Southw. Naturalist 42:435-440. 

SiLVEU5,W.A. 1933. Texas grasses, classification and description of grasses. The Clegg Co., 
San Antonio, Texas. 

Smith, FA and N.A.Walker. 1 980. Phtosynthesis by aquatic plants: effects of unstirred layers 

New Phytol. 86:245-259. 
Terrell, E.E.W.PH. Emery, and H.E.Beaty. 1 978.0bservations on Zizania texana (Texaswildrice), 

an endangered species. Bull. Torrey Bot.Club 105:50-57. 
U.S. Fish and Wildlife Service. 1995. San Marcos/Comal (revised) recovery plan. Albuquer- 

VAN Baalen, J., W.H.O. Ernst, J. van Andel, D.W. Janssen, and H.J.M. Nelissen. 1 990. Reproductive 

soil fertility. Acta Bot. Neerl. 39:1 83-1 96. 
Vretare, V.,S.E.B.Weisner,J. A. STRAND,andW.GRANeLi. 2001. Phenotypic variation in Phragmites 
australis as a functional response to water depth. Aquatic Bot. 69:1 27-1 45. 

VEiR,C.E.and H.M. Dai L. 1 960. A developmental study of wild rice,Z/zon/a ac 

J. Bot. 38:71 9-739. 
Vestlake, D.F., 1 967. Some effects of low-velocity currents on the metabc 

macrophytes.J.Exp. Bot. 18:187-205. 
;ar,J.H., 1984.Biostatistical analysis. Prentice-HallJnc.,Englewood Cliffs, f 


Royal Botanical Gardens 

P.O. Box 399 

)ilton, Ontario, CANADA L8N 3, 

Stachys cordata was described from Ohio by Riddell in 1836. The identity of the 
species so named was never in question, and the use of this name prevailed dur- 
ing the following 85 years. Riddell's type specimen is extant at US and micro- 
fiches are widely accessible. 

In 1921, House published the name Stachys riddellii in direct substitution 
for S. cordata Riddell, although he inappropriately used the designation "n. sp." 
Because this was merely a new name for the same species, no new description 
was provided and no new type was cited. House considered the name 5. cordata 
Riddell to be illegitimate, in that it was a homonym of S. cordata Gilib. Whether 
the new plant names in Gilibert's Flora Lituanica Inchoata should be consid- 
ered validly published under the provisions of the International Code of Bo- 
tanical Nomenclature was formerly a matter of doubt. This matter is now settled. 
Flora Lituanica Inchoata and Gilibert's other publications are listed in the Codi 
among the "opera utique oppressa," and new names at the rank of species an 
decreed not to have been validly published in those works. 

Immediately upon its being published, the name Stachys cordata Gilib. wa< 
widely recognized as being taxonomically synonymous with 5. syhatica L, th( 
name of a species native to Europe. Consequently, it was not inadvertently vali- 
dated by subsequently being accepted by any other author. Bentham, in his com 
prehensive treatment of the genus in 1848, did not note the use of this name b) 
any other authors m the interim, nor did he even cite it in synonymy Since the 
provisions of the International Code of Botanical Nomenclature pertaining tc 
homonyms apply only to validly published names, the use of the combinatior 

by Gilibert no longer constitutes a reason for rejecting the name S. cordata 

Following Houses publication of the name S. riddellii, that name came into 
general use for the species, although as late as 1933 the name S. cordata was 
used in Small's Manual of the Southeastern Flora. In 1979, Nelson and Fairey 
noted that the long-misapplied name 5. nuttallii Shuttlew. ex Benth. (1848), 
which antedated 5. riddellii House, was correctly applicable to the same spe- 
cies. The species has usually been called 5. nuttallii since their paper was pub- 
lished. They rejected the name 5. cordata Riddell only for the reason given by 
House. In his monograph on Stachys in the southeastern United States, Nelson 
(1981) included S. cordata Riddell m the synonymy of 5. nuttallii but did not 
address the question of the legitimacy of the earlier name. 

The only post-1981 flora in which the name 5. cordata Riddell was accepted 
is that by Gleason and Cronquist (1991). The names S. riddellii and S. nuttallii 
were cited in synonymy, without comment. In other standard references, both 
printed and electronic and including those published since 1991, the name S. 
nuttallii has continued to prevail. 

I conclude that there is no basis for the rejection of the name Stachys cordata 
Riddell, and that it is the correct name for the species that has been called 5. 
riddellii House and 5. nuttallii Shuttlew ex Benth. 


Benthan.G. 1848, Labia 

jtae. In: 

Candolle, A.P. de, ed. Prodromus systematis nati 


ctor Mc 

jsson. 12:27-603. 



revision of American Stachys. Repert. Spec. Nov. 


Gleason, H.A. and A. C 


T. 1991. Manual of vascular plants of northeasts 

States and adjacen 


ja, ed. 2. Bronx:The New York Botanical Garden. 

House, H.D.I 921. Nea 

?ssary ( 

ihanges of certain plant names. New York State 

Nelson, J.B.I 98 l.Stochys (Labiatae) in southeastern United States. Sida 9:104-123. 
Nelson, J. B. and J.E. Fairfy III. 1 979. Misapplication of the name Stachys nuttallii (Lamiaceae) 

to a new southeastern species. Brittonia 31:491-494. 
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Smali, J.K. 1933. Manual of the southeastern flora. Chapel Hill: The University of North 



M. Patrick Griffith Stephanie C. Bartel 

Rancho Santa Ana Botanic Garden Department of Biology 

Claremont, CA 9171 1, U.S.A. AlpineJX 79832, U.S.A. 

xted in the Davis Mountains ot Jeff Davis County, Texas. Chloroplast DNA sequences were obtained 

ion species of tlie Davis Mountains. Based on this evidence, the present known distribution of C 
nzonica in Texas remains restricted to the Chisos Mountains of southern Brewster County. 

lemos investigado un reciente registro de un individuo disyunto de Cupressus arizonica 
Zupressaceae) colectado en las montafias de Davis del condado de Jeff Davis, Texas. Se obtuvieron 
:cuencias de ADN cloroplastidial del nuevo registro y los supuestos taxa relacionados. El analisis 
logenetico muestra que el especimen de C arizonica de las montanas de Davis consiste en tejidos 
egetativos d(t]umperus deppeana, una especie comun de las montaiias de Davis. En base a esta 

Cupressus arizonica, as broadly circumscribed (Wolf & Wagener 1948; 
Eckenwalder 1993) includes a number of variable, isolated populations in south- 
western North America. The recent report of a single individual of Cupressus 
arizonica at Bridge Gap Spring in the Davis Mountains of Jeff Davis County, 
Texas (Karges & Zech 2001) suggests a more continuous distribution in suit- 
able habitats from California, through the southwestern United States, into 
Trans-Pecos Texas, and south into Mexico. Previously, C arizonica was known 
from northern Mexico (Standley 1920; Wolf and Wagener 1948; Little 1978 
Correll 6a: Johnston 1979; Rehfeldt 1997) southern Brewster County, Texas (Pow- 
ell 1988; 1998), and isolated localities in Arizona, CaHfornia, and New Mexico 
(Sudworth 1927; Vines 1960; Little 1971; Minnich & Everett 2001). The Davis 
Mountains specimen led Karges and Zech (2001) to postulate a greater Pleis- 
tocene range for C arizonica, represented by an occurrence in this additional 
Madrean sky island habitat. 

The specimen upon which this phytogeographic postulate is based (Karges 
& Hedges I'^SO, SRSC) may not represent Cupressus arizonica, however This 

sheet consists of several short (< 12 cm), separate, yellowed terminal branches, 
and three small (< 8 mm), detached woody scales. Another voucher of the Bridge 
Gap putative Cupressus arizonica (Karges s.n., SRSO consists of a single branch 
with green leaves, but no visible reproductive structures. Common members of 
the Cupressaceae that occur in the Davis Mountains Are Juniperus deppeana 
var. deppeana and J. pinchottii (Powell 1998). Also present in the Davis Moun- 
tams, J. deppeana var sperryi is an extremely rare endemic known from three 
individuals (Adams 1973; 1993; Watson & Eckenwalder 1993; Powell 1998). The 
Bridge Gap specimens could represent one of the above taxa, as concluded by 
B.L. Turner in an annotation (17 Jan 2001) of Karges and Hedges 2^80 d^sjuniperus 
deppeana. The distal vegetative growth of Cupressus arizonica and the above 
taxa of Juniperus are nearly indistinguishable. This may reflect on the sister 
relationship between these genera (Brunsfeld et al. 1994; Gadek et al. 2000). 

The putative occurrence of Cupressus arizonica in the Davis Mountains is 
of biogeographic importance, and we attempt to address the veracity of the re- 
cent report in Jeff Davis Co., Texas, through an alternative means. Cupressaceous 
taxa that are difficult to identify by conventional means have been accurately 
identified through chemical analysis (Gough 62: Welch 1978; Adams 1993; Hsiang 
& Huang 2000). In order to verify the determination of the two Davis Moun- 
tains putative Cupressus arizonica specimens, we have gathered and compared 
chloroplast DNA sequence data from these specimens and specimens of puta- 

The Davis Mountains specimens were compared to 4 mdividuals representing 
Cupressus arizonica, 2 other Cupressus species, 11 individuals oi juniperus, and 
1 specimen each of the outgroups Calocedrus, Chamaecyperis, and Thuja (Table 
1). This sampling was based on the generic relationships within Cupressaceae 
suggested by recent studies (Brunsfeld et al. 1994; Gadek et al. 2000). Specimens 
used for DNA sequencing were either collected during fieldwork in Mexico and 
the United States in 2001, previously vouchered herbarium specimens, or 
vouchered live plantings growing at Rancho Santa Ana Botanic Garden 

For all specimens collected m 2001, DNA was extracted using 2X CTAB, 
followed by precipitation in cold isopropanol (Friar et al. 1996). One gram of 
terminal shoot tissue (including leaves) was used for these extractions. For speci- 
mens collected before 2001, a modified small prep for dried leaf tissue was used 
for extractions (Doyle & Doyle 1987). Amplification of cpDNA templates of the 
trnl intron and trnL-trnF intergenic spacer follows the methods outlined by 
Porter et al. (2000). Purified template amplifications were sequenced directly 
with four primers, trnlc, f rnLd, trnle, and trnLi (Taberlet et al. 1991), using "big 

ORNiA. San Diego Co,: 

RSABG 11 746 {HS^) 

Karges and Hedges 2480 

..Jeff Davis Co.: 

Kargess.n. {Sf<SQ 

:o. D.F.: 


Griffith 287 (RSA) 

ORNiA. Shasta Co.: 

RSABG 11 434 {RSA) 

Brewster Co 

Bartel 575 {SRSQ 

Brewster Co 

Powell 5146 ISRSQ 

.) Puebia 

Griffith 324 [RSfi.) 

Jeff Davis Co 

Powell 51 86 {SRSQ 

Jeff Davis Co 

Kollel 193 {SRSQ 


Bartel 574 {SRSQ 


Griffith 253 (RSA) 

() Jalisco 

Griffith 279 (RSA) 


Gr/ffith 304 (RSA) 

Jeff Davis Co 

Keough 205 [RSA) 

Griffith 352 (RSA) 

3KNIA Humboldt Co.: 

Thuja plicata Donn ex D. Don 

dye" chemistry from Applied Biosystems Incorporated, according to the 
manufacturer's specifications. All sequences were gathered using an Applied 
Biosystems Incorporated 3100 automated DNA sequencer 

Chromatograms derived from sequencing were assembled into contigs, and 
edited using Sequencher v4.1 (Gene Codes Corporation, Inc.). Consensus se- 
quences were manually aligned usmg Se-Al v2.0a72 (Rambaut 1996). The 
aligned DNA data matrix is available from the first author upon request. The 
phylogenetic relationships among these specimens were estimated using Fitch 
parsimony, in PAUP* v4.068 (Swofford 1998). Estimations of confidence in the 
clades were obtained through bootstrap analysis (Felsenstein 1985) with 10,000 
pseudoreplicates, and through jackknifmg (Farris et al, 1996), also with 10,000 
pseudoreplicates (63% deletion) as implemented in PAUP*. 

The specimens of Cwpressus arizonica form a well-supported (94% bootstrap, 
80% jackknife) monophyletic group with three other Cupressus specimens, and 








^^ 51 

s(Karges and Hedges 

if the trnLF region. With uninformative characters excluded, length = 47; CI = 0.7660; Rl = 0.8854; 
ncy = 0.6782. Bootstrap percentages above 50% are indicated above the branches, and jackknife sup- 
is indicated below/ the branches. C = Cupmsus,}. = Juniperus. 

this clade excludes the two Bridge Gap putative C arizonica specimens (Fig. 1). 
The three specimens oijuniperus deppeana plus the two Bridge Gap specimens 
form a clade, though lacking strong support (63% bootstrap). Juniperus 
deppeana is a species present at Bridge Gap (Karges & Zech 2001). In context 
with the known flora of the Davis Mountains, chloroplast DNA data clearly 
suggest that the two Bridge Gap specimens are not Cupressus arizonica, and are 
most likely Juniperus deppeana. 

The first author examined the Bridge Gap cypress closely in July 1999 and 
made the following observations: The specimen's trunk bark is divided into 
rough squarish plates near the base, characteristic oi Juniperus deppeana var. 
deppeana (Powell 1988). At a height of about 0.5 m, the gray-brown bark is sepa- 
rated into longitudinal ridges, a key character f^or Juniperus deppeana var. 
sperryi (Adams 1973; Powell 1988). In contrast, the inner bark of Cupressus 
arizonica is often described as reddish (Wolf & Wagener 1948; Powell 1998), 
maturmg often into thin fibrous strips on large trees (Correll & Johnston 1979; 
Bartel 1993), although the bark of Cupressus arizonica sensu lato can be quite 
variable. Martinez (1963) notes that J. deppeana var. patoniana forma ohscura 
may have the lower bark checkered and the upper bark furrowed. 

Although other characters may be of use, in most taxonomic keys the ma- 
jor feature diagnostic betweenjuniperus and Cupressus is the presence of fleshy, 
fused cones (Juniperus), versus woody dehiscent cones (Cupressus) (Correll & 
Johnston 1979; Powell 1988; Bartel 1993, Watson & Eckenwalder 1993). Of the 
two specimens of the Bridge Gap putative Cupressus, one (Karges s.n.) is devoid 
of reproductive features and the other (Karges & Hedges 2480) has three sepa- 
rate orbicular-spathulate scales that are 5-7 mm long and 6-8 mm wide. These 
scales appear to have been sessile at the proximal end rather than peltate. 
Cupressus arizonica has 4-5 partially peltate proximal pentagonal scales ap- 
proximately 13-15 mm long, 12-14 mm wide, and 2-3(-4) distal oblong, trun- 
cate scales 5-8 mm wide and 9-11 mm long, completely peltate and valvate 
(Table 2). Morphologically the scales present on Karges & Hedges 2480 do not 
appear to be produced by C. arizonica. 

One of the two putative Cupressus arizonica specimens (Karges s.n) may 
be misdetermined because of a lack of reproductive characters, while the other 
(Karges & Hedges 2480) may have been determined as C. arizonica based on the 
three woody scales collected. Although the location of discovery of these three 
scales is faithfully recorded, we cannot be certain of their identity. Although 
unfortunate, this is not the first instance of a mixed collection being mistaken 
for somethmg more significant (Thomson 1991). Given the above evidence, the 
present known distribution of Cupressus arizonica in Texas remains limited to 
the Chisos Mountains of southern Brewster County. This may change if a speci- 
men exhibiting clear characters of C arizonica is collected elsewhere m Texas. 

California: San Diego C 
Texas: Brewster Co. 

The authors wish to wholeheartedly thank Sharon Yarborough of SRSC and 
Steve Boyd of RSA for making herbarium specimens available; Bart O'Brien of 
Rancho Santa Ana Botanic Garden for making live material available; Travis 
Columbus, Naomi Fraga, Clem Hamilton, Mark Porter, Mike Powell, Linda 
Prince, Smita Sanbui, and Gary Wallace for providing very useful comments 
on a draft of this manuscript; and we are especially indebted to Nancy Refulio 
for providing the resumen. Two anonymous reviewers provided excellent feed- 
back. We are grateful for a Mellon Foundation grant through Rancho Santa Ana 
Botanic Garden, which supported this work. 

Adams, R.P. 1973. Reevaluation of the biological status of Juniperus deppeana var. sperryi 

Correll.Brittonia 25:284-289. 
Adams, R.P. ]993. Juniperus Linneaus. In Flora of North America Editorial Connmittee, ed. 

Flora of North Annerica Volume 2:41 2-420. Oxford University Press, New York, New York. 
Bartel, J.A. 1 993. Cupressaceae. In J.C. Hickman, ed.The Jepson manual: higher plants of 

California 111-113. University of California Press, Berkeley. 
Brunsfeld, S.J., RS. SoLTis, D.E. Soltis, RA. Gadek, C.J. Quinn, D.D. Strfnge, and T.A. Ranker. 1 994. 

Phylogenetic relationships among the genera ofTaxodiaceae and Cupressaceae: evi- 
dence from rbcL sequences. Sys. Bot. 1 9:253-262. 
CoRRELL,D.S.and M.C.JoHNSTON.1979.Manualofthevascular plants of Texas. The University 

of Texas at Dallas, Richardson. 
DoYLE, J.J.and J.L. Doyle. 1 987. A rapid DMA isolation procedure for small quantities of fresh 

leaf tissue. Phytochem. Bull. 1 9:1 1 ~1 5. 
Eckenwalder, J. E. 1 993. Cupressus Linnaeus. In Flora of North America Editorial Committee, 

ed. Flora of North America Volume 2:405-408. Oxford University Press, New York. 

outperforms neighbor-joining. Cladistics 12:99-124. 
Felsenstein, J. 1 985. Confidence limits on phylogenies: an approach using the bootstrap. 
Evolution 39:783-791. 

Friar, E.A., R.H. Robichaux, and D.W. Mounf. 1996. Molecular genetic variation following a 

population crash in the endangered sWyersword, Argyroxiphium sandwicense ssp. 

sonG'w/cense(Asteraceae).Mol.Ecol. 5: 687-691. 
Gadek, P.A., D.L Alpers, M.M. Heslewood, and C.J, Quinn. 2000. Relationships within 

Cupressaceae sensu lato: a combined morphological and molecular approach. Amer. 

J. Bot. 87:1 044-1 057. 
GouGH, L.J.and H.J.Welch. 1978. Nomenclatural iransfer of Chamaecyparls obtusa (Siebold 

& Zucc.) Endl.'Sanderi' (Cupressaceae) to Thuja orientalis L'Sanderi'on the basis of 

phytochemical data.Bot.J. Linnean Soc. 77:217-221. 
HsiANG,T. and J. Huang. 2000. The use of RAPD markers to distinguish among juniper and 

cedar cultivars. Can. J. Bot./Rev. Can. Bot. 78:655-659 

tains of West Texas. Sida 1 9:71 9-721 . 
LimE, E. L. 1 971 . Atlas of United States trees. Volume 1 . USDA, Washington, DC. 
LtmE, E.L. 1 978. Important forest trees of the United States. USDA, Washington, DC. 

Universidad Nacional Autonoma de Mexico. 

Minnich, R.A. and R.G. Everett. 2001. Conifer tree distributions in southern California. 
Madrono 48:1 77-1 97. 

PoRTER,J.M.,M.S.KiNNEY,and K.D.HEiL.2000.Relationships between Sc/erococrus and Toumeyo 
(Cactaceae) based on chloroplast trnL-trnF sequences. Haseltonia 7:8-23. 

Powell, A.M. 1 988. Trees and shrubs of Trans-Pecos Texas including Big Bend and Guada- 
lupe Mountains National Parks. Big Bend Natural History Association, Inc., Big Bend 
National Park, Texas. 

Powell, A.M. 1998.Trees and shrubs of Trans-Pecos Texas and adjacent areas. University of 

Rambaut, a. 1 996. Se-AI. Sequence alignment editor. Version 2.0a7.2. Department of Zool- 
ogy, University of Oxford. 
Rehfeldt, G.E. 1997. Quantitative analyses of the genetic structure of closely related coni- 

(Cupressaceae) complex. Amer. J. Bot. 84:1 90-200. 
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ranges. USDA, Washington, DC. 
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Dayle E.Saar 

Department of Biological Sciences 

DeKalbJL 6011 5-2861, USA. 


While collecting material from wild populations of Dahlia to expand natural 
history information and for cytological and molecular analyses of the genus, 
several undescribed species from Mexico were encountered, one of which is 
described here. 

(Fie. 1). Type: MEXICO. Hidalgo: ca. 7 km NE of 

Plants to 1.5 m (type locality) or 1.2 m (paratype population) with l-3(-5) her- 
baceous stalks arising from perennial tuberous rootstock; internodes glaucous, 
hollow, nodes solid. Median leaves 19-24 cm long including petiole, bipinnate 
with stipels frequently at the second (occasionally second and third) rachis 
node, becoming pinnate without stipels to simple at the base of the flowering 
portion; petioles solid; rachis with prominent tufts of light-colored hairs at 
nodes, pinnules opposite on the rachilla, primary pinnae 5-7; leaflets dentate 
and ciliolate, blade bases strongly oblique, veins lighter-colored, sometimes cre- 
ating a dusty appearance at first glance. Capitula to 12.5 cm in diameter includ- 
ing rays, borne on unbranched peduncles 6-19 cm long and projectmg above 

the foliage; outer in volucral bracts ref lexed at anthesis. Ray florets sterile; ligules 
lavender with a spot of yellow at the base, veins darker below, tips mucronate. 
Disc florets hermaphroditic, yellow. Cypselae elliptic-oblong, 8.6 mm wide x 
1.2-1.6 mm long; black. 

Etymology and distribution.— Flowering commences in August (in 1995). 
Plants from the type locality grow along and in full view of a busy highway 
(Mex 105) near the large city of Tulancingo. It is surprising, therefore, that this 
showy species has not been described previously, hence the specific epithet, 
neglecta. The known distribution for this species is presently limited to the 
holotype and paratype localities. 

Paratype: MEXICO. Hidalgo: at K-8, 2 km SW of Mineral del Monte, along route Mex 105, zone of 
Cupressus and scattered sfirubby Quercus; road cut with dense vegetation on SE-facing slope, elev. 
2800 m, 11 Sep 1995 J.P Hjerting, D.E. Soar, & P.D. Sorcnsen 95-91 (DEK, MEXU). 
The combination of glaucous stems, tufts of light-colored hairs at the rachis 
nodes, strongly oblique blade bases, and mucronate ligules sets this species 
unmistakably apart from all others. Sorensen's (1969) key would place Dahlia 
neglecta in section Dahlia. Within the key for this section, D. neglecta pairs with 
D. australis and D. sherfjii at couplet "N" (page 325) and incorporates some 
morphological characters from each species. Dahlia neglecta is similar to D. 
australis, in that the lower leaf surfaces are conspicuously lighter green or sil- 
very green, but the former differs by having flat (not revolute or obscurely so) 
margins, a smooth (not rugose) upper leaf surface and glaucous (not glabrous 
or pubescent) canes. The fruit is of similar length but only 1.2-1.6 mm wide 
versus 1.5-2.6 mm for D. australis. The smooth upper leaf surfaces of D. neglecta 
are similar to D. sherffii. Dahlia neglecta differs from D. sherffi i by its glaucous 
(not glabrous) canes, bicolored leaf surfaces, and narrower fruit (D. sherfjii = 
2.2-3 mm). The unique combination of characters mentioned earher also dis- 
tinguishes D. neglecta. None of these species are sympatric in their distribu- 

A molecular study of the genus which utilizes combined molecular se- 
quences from the internal and external transcribed spacer regions (ITS and ETS, 
respectively) of nuclear ribosomal DNA, shows good support (86% bootstrap 
value) for placing Dahlia neglecta in a pectinate clade ("Variable Root Clade") 
that includes the "tree dahlias" (sect. Pseudodendron: D. imperialis, D. excelsa, 
and D. tenuicaulis), sect. Epiphytum (D. macdougallii), two species from sect. 
Dahlia (D. rudis and D. apiculata), and an unpublished species (Saar et al. in 
press). Within this clade, D. neglecta is most closely allied with D. apicu lata and 
D. tenuicaulis, but support is minimal (57% bootstrap value). Dahlia neglecta 
is not sympatric with these species and both D. apiculata and D. tenuicaulis 
have lignified perennial canes, whereas D. neglecta is herbaceous. 

The description of Dahlia neglecta brings the number of "wild" or natu- 
rally-occurring species of Dahlia to 35, but does not include the cultivated forms 


often called D. variabilis or occasionally D. pinnata (see Hansen & Hjerting 
1996 for clarification of the latter Latin binomial). This total does, however, in- 
clude three previously undescribed species currently "in review" or "in press:" 
D. hjertingii (Hansen and Sorensen in press), D. campanulata, and D. cuspidata. 

I thank Nathan A. Saar for preparing the illustration and Portia Gallegos for 
translating the abstract into Spanish. Paul Sorensen and Jens Peter Hjerting were 
invaluable throughout the field studies in Mexico. Guy Neson and an anonymous 
reviewer provided helpful comments to an earlier version of the manuscript. 


ANSbN, H.V. and J.P. Hjerting. 1996. Observations on chromosonn 

e numbers and biosys- 

tematics in tlie genus Dahlia (Asteraceae, Heliantheae) with an account on the iden- 

tity of D. pinnata, D. rosea and D. coccinea. Nordic J. of Bot. 1 6:4 


ANSEN,H.V.and P.D.S0RENSEN.(ln press).Anew species of Dah//o (As 


from Hidalgo state, Mexico. Rhodora. 

\AR D.E., N.Q PoLANS, and PD. Sorensen. (In press). A phylogeneti 

c analysis of the genus 

Dahlia (Asteraceae) based on internal and external transcr 

ibed spacer regions of 

nuclear ribosomal DNA.Syst.Bot. 

)RENSEN,PD. 1 969. Revision of the genus Dahlia (Compositae, Heli, 


Rhodora 71:309-365,367-416. 


Herbarium-Div. of Biology 

in Liatris elegans is presented, Liatris elegans var. hridgesii, 
t Eocene sand formations in central Texas. Liatris elegans 

n of Alabama and Georgia to the 

s elegans vzr.flabellata is reduced to a form of L. elegans var el 


Se presenta la 

descripcion de dos nue' 

v'OS t 

axa infraspecil 




maciones arenc 



y Georgia hast: 

a el estrechode Florida. 


nuevos taxa de: 


blanco-crema a amarill 

5. color de anto 






s.Uatns elegans v.r.Jlc 


alaseha reduc 

s poblac 

Liatris elegans Michx. is an obligate psammophile, occurring in well-drained, 
loose to somewhat compacted sandy soils in open to sparsely forested habitats 
across much of the Gulf and Atlantic coastal plains and in the southern Inte- 
rior Highlands province in Arkansas. Its colored, elongated, conspicuous phyl- 
lary apices in combination with white to yellowish-cream corollas distinguish 
It as one of the most recognizable species in the genus (cf. King & Robinson 
1987). All other species have more or less herbaceous, relatively short, incon- 
spicuous phyllaries with exserted, lavender to pink corollas (Gaiser 1946). Thus, 
m L. elegans the phyllaries rather than the flowers provide primary visible col- 
oration of the columnar inflorescences. Throughout most of the range of L. 
elegans (herein, L elegans var. elegans and L elegans var. carrizana L.O. Gaiser) 
populations have phyllaries that are predominantly shades of lavender, pink, 
bluish, or magenta, colors that are presumably generated by anthocyanin 

pigments. Two sets of populations, one on the western margin of the range in 
Texas, another in the eastern Gulf Coastal Plain m Alabama, Georgia, and 
Florida, mostly lack classic anthocyanm coloration at anthesis. Instead, these 
have primarily yellowish, cream, or nearly white phyllaries with some popula- 
tions variably cream to faintly lavender. The Texas populations were discussed 
briefly by Gaiser (1946, p. 174, as L. elegans forma/isheri Steyermark), but accu- 
rate description of the variation is lacking. In addition, the well-defined pat- 
tern of distribution of the eastern coastal plain pale-phyllaried populations has 
never been addressed. While phyllary color alone may not be taken as a reliable 
indicator of genetic cohesion, the geographic integrity and populational uni- 
formity of these unique populations suggests that selection for phyllary color 
has taken place within these populations. A recognizable suite of other mor- 
phological features supports their recognition but the presence of intergrada- 
tion of both sets of populations with var. elegans further suggests that recogni- 
tion at an infraspecific rather than specific rank is warranted. I have chosen 
variety to indicate that intermediate populations occur along the margins of 
these novel taxa. 


orma Jisheri Steyermark, Field Mus. Nat. Hist 
TfZIu^^!"' highway, 4 mi S of Copperas 


ug 1934. George LF.sfier 

var. eleganti apicibus phyllariis petaloideis f 


^urneisvelpallide resets 

Corms globose; stems sparsely to densely tomentose; lateral heads usually 
densely crowded, divergent-ascending to spreading at right angles, sessile; phyl- 
laries parallel and appressed below the petaloid portion, the colored apices re- 
curved, abaxially pubescent to tomentose, initially pale lemon yellow, then fad- 
ing to cream or very pale lavender, rarely rosescent at fruiting; corollas 
cream-white, tubular, the tube scarcely broadened to the top of the limb, the 
lobes scarcely flared apically; stigmas cream to light yellow or nearly white. 

Paratypes: U.S.A. TEXAS. Anderson Co.: George Engling Wildife Management Area, Upper North Pas- 
ture, W Side, 1934 Burn, 18 Sep 1958, Marsh 55-8 (TEX). Bastrop Co.: 1.8 mi SE of McDade along old 
highway by railroad track, 30 Jul 1992, Mayjield & Westlund M86 (NCU, NY, TEX). Freestone Co.: FM 
1848, 5.8 mi N of Buffalo at jet. US 79, lower Queen City formation, N 31°32'46" W 96°03'22", 6 Sep 
1992, Mayjield 1557 (F, FLAS, GA, GH, NCU, NY, TEX, US); 14 mi S of Fairfield, 15 Aug 1955, Turner 
3904 (BRITTEX). Grimes Co.: along main highway E of Brazos county line, 4 Sep 1993, Mayjield etal. 

of Poynor, grassy open oak woods in sandy soil, flowers whitish, 20 Oct 1963, Correll 28558 (LL). 

0.9 mi E of FM 1733, 21 Sep 1991, Mayfidd el al. 1123 (NCU. NY, TEX); 2 mi S of Grapeland, 12 Oct 1937, 
Cory 26136 (F), Lee Co.: NW of Lexington, on county road to Phears Cemetary, by small pond, Carrizo 
formation, 500 ft, N 30"27'30" W 97''03'15", 15 Sep 1991, Mayfield etal. 1117 (GH, NCU, NY, TEX). Leon 
Co.: 5 mi NE of Buffalo along hwy 79, 11 Sep 1958, Turner 4433 (TEX). Limestone Co.: 5.5 mi E of 
Kosse, 9 Oct 1964, Skinners 30519 (BRIT). Nacogdoches Co.: 4 mi W by N of Garrison, 7 Sep 1948, 
Cory 54734 (LL). Robertson Co.: county road 330 ca 2.0 mi E of its junction with FM 2246, 500 ft, N 
6 Sep 1992. Mayjield 1559 (GH. NCU, NY, TEX). San Augustine Co.: sandy open 

1 lake on Upper Iron Ore Cr, 1.2 mi 

e N side of the road, 22 Sep 1991, 
s E of Sam Houston St Un. 27 Sep 

., Mayjield et al. 1120 (E GH, NCU, NY TEX). Wood Co.: Le 

Forma as a category fails to represent the clear geographic pattern that occurs 
■. hridgesii. I have chosen to provide a novel epithet for the present taxon 
because the type locality of forma/isheri is apparently inaccurate. No popula- 
s of L. ehgans var elegans (Fig. 2) occur anywhere close to the designated 
tion "on the highway 4 miles south of Copperas Cove'' in Coryell County. 
While there are some thin, Cretaceous sand-bearing strata in this area as well 
as some river terrace sands, searches around Copperas Cove have not yielded 
populations or habitat suitable for L. elegans. Furthermore, no other collections 
he area were located in searches of three major herbaria in central Texas, 
among others. The establishment of a new type and name will serve to solidify 
ncept of the taxon. 

^mology.— This variety is named for Edwin L. Bridges, prolific plant col- 
and knowledgeable f loristician/taxonomist. Bridges' tenacity m the field 
and his mcredible plant recognition skills are truly inspiring. Our memorable 
■ips into the sand-bearing strata east of Austm, Texas, were crucial to my 
understanding of this and many other sand-loving species. 

Distribution, variation and ecology.— Liatris elegans var. hridgesii is best 
epresented in the southwesternmost part of its range from north of the Colo- 
rado River in Bastrop and Lee Counties northeast to Henderson County in Texas. 
Here, populations consist entirely of individuals with phyllaries that are creamy 
white to very pale lavender, and possess long-tubed corollas with relatively short 
weakly flared lobes. In the Post Oak savannas of Williamson, Milam, Lee, and 
Robertson counties, all populations I found had consistently cream-colored 
phyllaries. The latter begin the flowering season with a very striking, nearly 
lemon yellow color in the phyllaries, a color that is otherwise not present in the 
phyllaries of any Liatrinae. Later in the season, well into fruiting, the color fades 
to creamy white, sometimes infused with a rose or lavender tinting. Further 

northeast, populations seem to intergrade in terms of phyllary color but other 
features are consistently more like var. hridgesii than var. elegans. The phyllar- 
ies of var. hridgesii tend to be longer and more strongly recurved with longer, 
more narrowly tubular, pale cream corollas than in var. elegans. The var. hridgesii 
is similar to the next newly described variety in the pale coloration of the phyh 
laries, but it has a more compact inflorescence with more strongly recurving 
phyllary tips and sessile heads (see further discussion below regarding pedun- 
culate heads). 

The var hridgesii occurs almost exclusively on Eocene sand-bearing strata 
running from southwest to northwest from the north side of the Colorado River 
in Bastrop County to Wood County in the northeast. Its influence is also appar- 
ent m population eastward through Cherokee and Houston, to Nacogdoches, 
Shelby, and San Augustine Counties where the Carrizo formation is exposed 
around the western margin of the Sabine uplift. Post-oak savanna is the pri- 
mary habitat where this variety is found. Mixed pine forests of eastern Texas 
are more likely to have populations of L. elegans var. elegans (Fig. 2) but both 
occur therein and some intergradation is observed. 

Liatris elegans var. carrizana L.O. Gaiser (Fig. 3) occurs south of the Colo- 
rado River in Texas on deep, sandy soils of xeric sandhills (cf . Gaiser 1946). This 
variety is characterized by its elongated, napiform corm suited to the drier cli- 
mate and deep sands that prevail in this region. Its phyllaries are structurally 
similar to those of the var hridgesii, with elongated strongly recurving apices, 
but they are consistently colored blue, magenta, lavender, or pink, rather than 
cream. Populations of var. carrizana just to the south of the Colorado River in 
Bastrop and Caldwell Counties, Texas, may have more or less globular corms 
but are otherwise more similar to the var carizzana. In all other areas, L. elegans 
has depressed globose corms as is true of many species of the genus. 

Liatris elegans Michx. var kralii Mayfield, var. nov (Fig. 4).Type: U.S.A. ALABAMA. 

Li!!; Co.: Locally abundant in open pine woods 6 mi S of Auburn, 23 Sep 1899. F.S. Earle and 

L.5. Earlc 94 (hoi otype: KSQ; isotypes: F! GH! ND NY! US). 
SiimlisaLe/egciiKi var. hriJi^csJiapicibusphyllariisflavisautalbisvelpallidelavandulissedcapitulis 

Corms globose; stems sparsely to densely lanose-tomentose; lateral heads (at 
anthesis) usually loosely arranged, ascending to divergent, pedunculate; phyl- 
laries abaxially tomentose to densely lanose, divergent along their length, the 
petaloid apices flaring only slightly or not at all, initially light yellow or cream, 
then fading to cream; less often pale lavender to rose, corollas pure-white, tu- 
bular, the tube flaring gradually to the top of the limb, the lobes strongly flared; 
stigmas white to very pale lavender. 

6.4 mi N of the Barbour/Dal 

e County line along Alabai 

na route 51, ca. 

4.5 mi N of I 

)oster, Turkey 

Oak/Bluejack Oak/Hickory 

sandhill area, 495 ft, 3^40'! 

)5" N, 85^3810" ' 

^, 12 Aug 1994, Mayjidd & 

Fuertes 20]] (TEX). Covingto 

.n Co.: Dozier, 31 Oct 1941, 1 

ieed 2053 (TEX 

) Escambia C 

o.: dry, sandy 

area above low hillside seepage bog under powerline, on 

N side of waste, 

rn extension c 

)f Co Rd 6, 2.4 

mi W of Co Rd 27 and 2.6 m 

i W of AL 113 at a pomt 3.8 



ton, just W of 

Pond Cr, SH SWQ SEQ of seel 

, TIN R7E, Flomaton 7.5' quad, Soils-Plumme 




I'Ol" W, 11 Sep 1989, Orzdla 

nd Bridges 1181: 

5 (TEX); 3 mi 

s of Canoe, 17 

Oct 1929, 0'Nein6]7J(FLAf 

5). Geneva Co.: Sandy west 

banks of Choc 

tawhatchee F 

liver between 

Geneva and Eunola by Al 52, 

18 Sep 1971, Krai 44342 (N\ 

O.Lee Co.: 2.1 ra 



1971, Krai 44247 (NY). Mobil. 

368, Leio ng 4879 (NCU).Russe 


of GlenviUe, 26 Sep 1976, Ha^ 





s.n.(NCU). Santa Rosa Co.: pi 

ne woods on Juniper Creek, 

Rd. 4, 6 Oct 194 

ington Co.: vicinity of Falling Waters Sink, longleaf pir 

le deciduous sc 

rub oak foresi 

t, 17 Sep 1956, 

Krai 3574 (GH) GEORGIA Ben Hill Co.: 7 mi NNW of Fit 


1968, Fairclot 

h569J (NCU). 


)n outcrop of Altamaha Gri 

t, 17 Oct 1970, F 


(NCU). Grady 

Co.: Sandy clay ridge, open, : 

I mi W of Wigham, 26 Oct 

1 963, God/re_v 6 

3218 (l.L). Ma. 

con Co.: Near 

Etymology— This variety is named to honor Dr Robert R. Krai, well known as 
one of the most knowledgeable botanists on the southeastern US flora. Krai's 
personal collections amassed over four decades from the coastal plain have con- 
tributed greatly to the understanding of this and many other species. His classes 
I took as an undergraduate in Plant Taxonomy and Dendrology stimulated my 
interest in botany 

Distribution, variation, and ecology— Liatris elegans var. kralii occurs in 
two major population centers. Populations to the north, in the Fall-Line 
Sandhills of western-central Georgia to eastern-central Alabama, including the 
type, consistently bear phyllary apices that are white to light tan in color. A 
second, somewhat separated area of occurrence is further to the south in coun- 
ties along the southern tier of Georgia, northern tier of Florida, and west to 
Mobile County in Alabama. In this area scattered populations of var kralii oc- 
cur sympatrically (but not apparently in mixed populations) with var. elegans. 
The var. elegans is much more common in this area but relatively pure popula- 
tions of var kralii do occur, albeit usually with both white and pale lavender 
forms (e.g., Krai 37333, Mobile Co., AL; Krai ^4342, Geneva Co., AL; Faircloth 
5691, Ben Hill Co., GA; Krai 3514, Washington Co., FL). All populations refer- 
able to var. kralii have short peduncles bearing the heads and the heads are 
well-spaced and ascending. Although pedunculate heads may otherwise occur 
throughout the range of the species in shaded or apically injured plants, these 
are rare and unusual. Also, the phyllaries of var kralii are longer and narrower, 
with the petaloid portion especially elongate in proportion to the basal herba- 

ceous portion. Another recognizable trait of var. kralii is the divergent phyllar- 
ies with the tips ascending, rather than recurved. All other varieties have more 
strongly recurved phyllary apices, with the lower herbaceous portions closely 

Variety flabellata. — The type specimen of Yd^r. flahellata apparently con- 
sists of a single plant divided into two separate sheets. The only two sheets re- 
ferable to this taxon are at NY; one has a single plant with root and two stems 
attached and a third