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by William F. 
Mahler, Director 
Emeritus of BRIT, Bot,ink,U"acul 

i o|)VM.ilit .'unu 
- itan ( al Re • in 

Table of Contents 


I ii 1.1 Piiii u N ii lin ii | in i i () i T i 65 


Sarcostemma cynanchoides .*• . 

Rhynchospora le 

»f Alocasia, Caladium, Colocasia, a 

-i i A ii -175 

i Tami.h Basin, Cai:iorn:,a 

64 114 132,136,148,184,188,194,204,223 

\ . . . 234 





otanical Research Institute of Texas (BRIT) 

> > Box 3 

14, Lae, Morobe Province 4 1 1, PAPUA NEW GUINEA 



esented from the Josephstaa i orcst Management Agreement Area, 

niny e> ilu iln-n tut -ill - I , t i - 1 i it it hjim k u>s I nui new. 

ibed: Aglaia saxon Ml , - f ma josephstaalensis 

• , 

i <■ ' I u 1 i i i ' 1 uhaceae).A substan- 

records and discoveries f ran ixa are reported. Compilations of 

s t mduscs£ 

ire also included. 


fi, *<Dt&Wtfi*.3C^lZfjt)tlTl\&. t ft t>(D%\m\* Aglaia saxomi(-\z>?> 
fsf), Barringtonia josephstaalensis (+*-# U / W4), Calycosia mamosei{7ti * 
14), Psychotria mayana(7fi*tt)ThZ> a SH£|fc|llCliE%:S*fiJffl£j£fc<!:<D 

TheJosephstaal Forest Managerr 

tish biological baselines ioi the concessional aieas.An ecological reconnaissance of the 
If i l i ) ill I I'll if i I I mil ii ■ i 11 1 1 I lni a | a i 

ber9and 17, 1998, followed b\ i u-i i Ml n n ilu r In > t the period from July 
26 to August 25 P II I II | i| i i | tr i hm findings from these 

The survey tract is located ioi the mosi patt.wiihin leu itoiv coveted by the Annanberg 

topographic shec r, bin .4 < th h h 1 i ' I. 4 it h i iik I Nubia map units 

(cf. Australian Survey Corps 1 973, 1 974a, 1 974b, 1 974c: 7888, 7889, 7988, 7989). This gen- 
eral area is part of if 1 " ' li journal- , T ih i T.r.'u , p bsion of Bogia Subdistrict; and in- 
cludes the principal 41 ' I in lui i I nm i umirap,Wadaginum, 

The 1999 survey was based at three camps established sequentially at map coord 
nates (GPS) 9504500 N hi 40 4P u h r\j x 280100 E;and 9498679 N x 284829 E;at 
elevations from ca. 50 to 160 m. There was maiuh h . th footliill forest at Camp 1, and 
alluvial terrace communities at Camps j kiMl | -\ \\ ]C latter base-', provided convenient 
access to both riverine and foothill vegetation. 

All investigated site n ithn n i f a 1 1 luit I f Jul lowland forest 

occurduring Januan \ t i i|i Imi tint i t il md the dtiest in 

MaytoAugustwInu outheasterly tradi be. mn offecti (M< \lpine et al. 1983:65). 
Even during the relative tliy season, nvviauo monihlv inintalls ,ne still generally around 
200 mm, so the vegetation is only nfrequontlv subjected to soil moisture deficiencies 
under normal cot I categorize the 

project sites as tropi 4 i i ui m i ' ] h 4 4 descriptor empha- 

sizing the oveiall 4 . "i - t ,,atei I th it 

fhe sur il ne tvpi f i I f I I I n n it n 4 in Imi i ntl n 

developed by ( xlt i| Lslion liom station I I n tl | ntattv Although 

the nearest station with pulh h 1 1 'I I' i 1 line I I 4i innual rainfalls with 

moderate seasonahh - -i Iron bit | M ,> ,fi t . to o. ■ ui in this general region. 
Episodes of widest .ii iltm >i \\ es' Pmi.t u in e been documented (Johns 1986: 

F in 1 o. i ( 4 , [ t - ti ^ * I il | h t 4 I i ii , m I to ice represents 

the crustal remnants of an island arc which collided with the Australian plate about 10 
m.y. BP (Pigram & Davies 1987). Parent substrates are generally derived from basalt 
volcanicsofthis now disnpin unci an (Jagm Pol >inson I PP I J) Severe earthquakes 

oecui fu qui nth ithiu il i u II ill 1 1 l< f i | ^ t [ In it pe< itiou is piob 

ably being encouiai iod I \ I no 4 id. u. 4 1 I i Pi 1 1 1 1 P 4 nos particularly at the 
higher elevations (cf. Balgooy et al. 1 996: 201-02). 

IS 1 till B MM lid 1 Stlic t If 

icluding Josephstaal) are dystropepts 

i group characteristic of lowland r 

labitats and comprising the most cc 

n Papua New Guinea (Bleeker 1 98 

3: 98-9).There are no ultramafic or c 


Botanical work on the Josephstaal flora has bem p i i I, it I iw 'ottunistic in nature, 

the largest collection .. i tit- Mill kr dm- In i i | u d September 1 to 15, 
1958.White obtdiiu d i i t i M l i n I i ml it i I r Guinea Force) series, 
from which three collections were later desu in, ih i h| | mi n \ i , m u m 
limbo Kosterm.,/k"'< ',-;). Mm < ■ ,, • tci -se Hartley & Perry). 

ut' uif di ii tK ii ii i I i in lit nt I in I i e been generally 

complicated by tin tttutin linni I m i|i il I oh botanists in Papuasia. 
The K I Win to numb i n il . n I i in In i t n i I i die period, since for all 
sheets the point of reference is clearly speuf 1 1 i | kma t L 4 45 S and Lat. 145 

00 E.The elevation is t mi o tt nth n na tm mi |i r llv indicates riverine or 

alluvial habitaK Pen I n i -nun m m nnl mm li . k.< m mentioned on some la- 
hels tog, //ms/onm ksm//y\ in NGF k.k ,kbw to ,n , „iii i / P Vo, and Cryptocarya 
weinlandtt\nNG r ' i | ti It mtb n in li i ill MiVVInti lethm 

inqs, IheiL nmon i I mmbiimt In i liimnm l t basin connected to 

tin iiiimh m | t I i r | n' 4 il tun- " > | i I il I 1 ni| anm 1 I ho 

Robbins,axsugi|i ted I n on. n \ h mlmit i urn n o ' meosc? in Robbins 1625), 

which is cross reton i I 1 , mo ' ' • < 1M I In I n bo to the latter as a 

duplicate. Robbinss lm< -iht\ .pnilnmr, P.gil .ml le mi in ,u Atitau area,Madang 
boti t ' ' m ,, i, n, u ,- L / i i4 I in. In o i osephstaal 400 ft.' 


:o them by specialists fndk ate the sort ol taxi t hat wo 

from theJFMAAc 

m in in mil i ImuhuM mil iiulou 1 \u al otounds.The \ 

tions have been ir 

scorporateci into the survey documentation (Appendi. 

species involved i 

ire present within and immediately around TNC's proj 

During the h 

■ il it nil o Mr ph . | mu II tli h 1 i li 

amined from the 

■ vicinity of losephstaal. A typical label bom Pullet. 

keen e i|i t ■ the collection site as lb tm south o" | h i , i an si up. Alt liocmbi tl 
Pullen numbers ate from comparable lowland habitats, 1 1 lev wet e collected slightly ol 
side the project cnea am: are mentioned o mi i v mcioeiitak in the following s 
any event, eonifoirativelv few Allien spec 
bauum (LAL);the NGF sheets comprise a c 
Other botanists with collections fron 
B.S. Parris, and J.P. Croxall. the Parris nod c roxall numhets ate specialist pteridologic 
? along the unimpiovecbnad to t oph t i u 1 [] y-i\ Must of tin it let 

removed from the project site but are notable for being among the few sets taken from 

By far the most comprehensive specimen series for Josephstaal are the collections 
from the TNC-sponsored botanical surveys of 1 998 and 1 999. A total of 973 numbers 
were added to the national herbarium from these efforts; 62 from the 1 998 ecological 
reconnaissance and 911 from the recently-concluded expedition. The combined tally 
increases the plant documentatiot irly an order of magnitude over 

what was previously available. Due to the surveys' exclusive focus on the elevational 
interval below 400 m, the herbarium coverage for the JFMAA is now among the best for 
any lowland wilderness in Papuasia. 

The 1999 survey consisted of general exploration and collecting around three expedi- 
tion camps, primarily using establish' I i | it r Towing the secondary channels 
comprising the Guam drainage. The botanical collections were conducted by an inte- 
i)t UmI it im >n istino r )| f i nut. M i i ] n in. I Wiakabu. During the 

selection of specim. n I I 11 n m . lit I u u| usually spurned by 

botanists because of their inherent repellent qualities or other difficulties associated 
with their processing. Palms, aroids, stinging nettles, grasses, alien weeds, etc. were se- 
cured when suitable sp. miik n ^ n in mi in i I in nti.isL to the general reluctance 
for collecting such plants. Multiple^ ' nil ti mm I -it i, i • mi aUo made, when their 
significance was already apparent in the field, in order to allow evaluation of population 
variation. Survey protocols were consistent out tl - f ■ r i r . pi L -_tive of developing a 

Ethnobotanical polling was com kit tori separably by survey biologists J. Wiakabu 
and M.Gorrez,through group intM i i- i Nitni and uses of specific 

plants were recorded,with spec > taxa. All vouchers 

were field-pressed in 70% surgicT i 1 i I I iumhI transported to the PNG 

Forest Research Institute (PNGFRI) for processing and determination. Materials for 
exsiccatae were often accompanied by bottled, carpological, and xylarium accessory 
collections when these were necessary for identification. 

The Lae National Hert 
Distribution of duplicate s 

Rijksherbarium (k), and Harvard (A), are the principal receiving institutio 
will be allocated in conformity to preexisting agreements or in complk 
TNC requirements. Whenever possible, specimens were named using t 
nomic revisions, or from a combination of authoritatively annotated she 
descriptions. Some collections could only be assigned with doubt to a < 

ml NkRAI 1)1 nCRIklk'ih. Of MIL Viol lAkOP 
Iwo pniv pal Ion t mi in ,| li i ■! ill mI I i t 111 uui' (.'traces, and foothil 

1 ilk d . iiiiui ill. ii in mi. i In ill. i ' ' ' .n Ml in ill 

showed thai in ii i n n it I |m n h I I I ml Ml t Inn Mi h 

The margins of large streambeds in the Josephstaal area are marked by a distinctive 
riverine fades domin.iii'd by liaiumheliophvtes.and rhoonhviu tnxa. Ihis edgecommu- 
the interior alluvial^ m 1 in i | i in . i I , 1 1 j | i.t Jon dominant, and 
with poor vertical development. On better-drained alluvia the forest becomes more flo- 
ristically and structun Ii i - .-mill t n n , t mied community with interlock- 
ing canopy layers and clear understoiios. I hem ,nn intoimodiate communities appar- 
ently linked to dimn i Inr nli | I tn 'mi M i I'. . . n i li • s with a linear series 
of staggered terrace n. int mediai ne ini tpretable as a successional sequence 
resulting from progis i i m i in n hi u ixon,pm mim.). Superimposed 
over the matiK ot hi it i n mill ini t ii n t i <■ t i. nowth, which have 

alluvial forest is for tf i i r ii n < t n tun I ki n Hammermaster and 

Saunders (1995a), tl i i irly a mt mum t ihiuiii ithin this category. 

ridgelines was initially regarded as homogeneous. Herbs and subarborescent plants 
appear to range thu umlml i hill iibiiii ti it ib n m li ml mtional separations. 
However Wetennq ip t mm.) noted pi ui I ui i 1 in mopy compositions 
between ridgelines and lucted timber as- 

sessment. His observation is supported bvine known aui ecological patterns of arbores- 
cent genera, as foi i h i| I t 1 \\ i i n li| t t it ( u 1 im test environments 

lion ol the foot knl I fon nation into subtypes, thouoh the 1 1 ii 1 1 'inn tin I ion may be primarily 

the near-ground com| mtot htii i. h it.n t tl I I. t n t immunities will re- 
quire surveys of grealei intensity than the one just com kinked, employing a combina- 
tion of transects atmt i i I in I tn ilk 'ii i n if | i ted by recent findings 

The Josephstaal foothill communities fall primarily umk'i forest structural code'Hm' 
and are known to inti ,u I n 1 1 i In i it in t llnnti m i tei K Saunders 1 995a: 
1 1).The'Hm' category is Urn major met* hantnble forest unit in the existing JFMAA 
(Hammermaster & Saunders 1995b: SB 55-1 Bogia overlay).On the earlier classifications 
of Paijmans (1 975) and Saunders (1 993), the project sites are placed respectively under 
structural codes THm' and 'Him' Due to similarities in leunmolouv employed by each 
milk >i tin kttemi t t | i i m ill mi i k i in I iptions.The J. epl I i il 

tract is essentially a t , | il m< .liunt . 1. 1 • m- I I. >i >t tn >m > . I. ,ti si en itonmenl 

assignable to specific subunits. 

The plant names provided by respondents are derived from the traditional Maia (Maya) 
language spoken by villagers within the project area. Clan elders Francis Muoimuado 
and Josef Sigagopa were principal sources for the information summarized in Appendi- 
ces 2 and 3. Ethnobotanical questioning usually occurred in the presence of a village 
audience, with the clan elders st it i i i -in 'it t ti thtator. A consensus 
was thus established. The group nit it id I uneans for identifying the assets 

i from ecoforestry operations. Because the survey objective en 

rapid appraisal procedures wpi n i i|l i mi -inalytic methods. All 

aspects of the ethnobotanical ingun !l -:qu ^ ■ uti al ^valuation against established 

the Adelbert Range. 

Despite these limitations, certain patterns are evident from the compilations. 
Unlike the nomenclature of formal hi d i ously applied bino- 

mial protocol, the botanical c r 'i ''mpk. i n ■ ■ . lagers is multifari- 

ous and idiosyncratic. Most Maian iit, M [i , hi ] r'ptively based, and if 

translated will probably be seen as alluding t pai ilarl iture Df a plant, in the man- 
ner of pre-Linnean botany. Although m I i ul not a two-element nomenclature, 
Maian plant names can occasionally c^xhiliit funciional rr\rmblances to a binomial sys- 
tem. In such instances'generic unit ii nil ii i in d with a common designator at 
the front of a complete name ed with a gualifying phrase 
or word following the generic mark. Examples n* I i i 'iubu-nganam,"warubu- 

chondrocarpum, Dysoxylum biu-, n I < ' nun Various taxa in 

Strobilanthes (Hemujhiph 1 ,) an imiLnl i> man I i , variants of 'sagag;' i.e. 'sagag- 
gosmun,"sagag-u-goga-umun an I hi'mi i in mentions are comparable 
to findings reported l> - tir ^t il i n > i r . , r n a study conducted in another part of the 
Adelbert Range. 

In most cases, similarities in gross appearance are apparently the major criteria for 
application of names I III n i ii I \ I I r il ^'pisanthessenegalensis 

(Sapindaceae), Ixora sp. sect. Hypsophyllum (Rubiaceae), and Phalena coccineo 
(Thymelaeaceae), which are all identified as'kibi-kibale'despite obvious contrasts in their 
fertile aspect.The gross equivalence in habit and leaf form is seemingly sufficient for 
combining these taxa under one concept. The Maian 'maberu' is similarly applied to 
Cleistanthus sp.aff. papuanus, Erythrospermum candidum in I h *" t> 
showing again that names are assigned on the basis of superficial aspect rather than by 
awareness of specific ' i tural ; .tin uons The rationale may be less clear however, in 

regards to general form. Other than the fact that they all represent pmnately constructed 
fems,Microsorumnh i W/<; Bolbitisquoyana, 

and Pleocnemia maaodonta, are obviously diffoient plant /et m relegated at least in 
parpto the one n im h i i i II tint i t in noil t t i nt d dagol-dagol'are 
even more striking, inch idu i nl Imn tiM as A-rlmmm ( f .imboinense, Lindsaea 
obtusa.andLipans " ' ' ft tutt^M Mi it th . f I . t m ,u obviously differ- 
ent, the Maian cla ificati nt I i u ti ui I I uiulm tit >l taxonomic equiva- 
lence in the Western sense, hut must he pun ending (torn some other logic; possibly 
involving a principle of utilTv. 

In tloustically tic h etivttonments such ,is utc generally present in Papuusia, only a 
fraction of the bci in il <i it II nt I It u ( n t u n< .s Many plants re- 
corded by the recent ui t t 1 I il < I on t it ea t th eg n 
dents were unable to ptovide one.OI those- taxa whit h find tokples assignments very 

small percentage at< n t nil it -tin it • > t m > ' n . u In 3) When a plant is 

n' I i i i igi a kh t i , i le I ut i i I in u it I m- pit inetic root for that 
name is often not ira listened toot hot ptants. In general the convene relationship is also 
true;Maian names with hetetogerii >u groui m mbership are generally composed of 
'useless'element tin t>i Imht it it nl ippltcations. Where 

plants of diverse appearance are placed rogethet nuclei i ommon designation, the spe- 
cies involved are not et g uti ihi ilu> > t- i m- mMr nil ultutal perspective there 
may be no imperat I i It tit i i lit i it n a\ Maian botany is arguably 

grounded on pout Ipunupli mint in iu i t in it expends efforts 

toward the identifk itt i t n out . I \ ith ultuu i| gin atton, and tends to 

consign everything - I i t I ^l, I In It tit t ut a mtu n tradition 

such economy is prooablv tin essaiv to t m s 1 1 i <. t theamouni ol ethnological data to lim- 
its amenable to otal tuiuot n i t imlntei u " Inl tl ' 01 1 1 | bant classification is 
typically aitifictal and it i| | i ni . m i hut tt t milt tu insearch, it appears 
to represent a system closely adapted to local interests and reguirements. 

The fact that very different species are (ten pla I undo tin m, Maian nami 
will complicate INK ut ut t tt i u t t t t t t i n i litional landowner 

groups. Especially in |> i fatnili u i it it I i nci Annonaceae,whereiden- 
tiftc iti<jns itebig I I i ui it i ii i tt i u I i I j tt e structures, Maian 

botany will be unable ' 1 1 it n I nhm-m In i n uiti n i n at generic level. This 

discourages use ol tl n t in i i b i i t t t u tu il it miction, and neces- 

t it- nttodu i si if Western concepts into the ttainin t t i 

The frequent lace ot lokpies spec ificity also argues ega'ust reliance on villagers tea 
plunl i '. b ■' i 1 1 1 1 c alien , in I'Mr 



Understory shrub; monoaxial or branched, to 2 m tall, entirely glabrous. Branchlets terete, 
apically and discontin m I ' i i 11 mi f iniderm crustaceous.exfoliating 

in flakes. Leaves spirally congested in terminal rosettes, blades herbaceous or fleshy, 
adaxially medium green, abaxially light green, obliquely ascending in the lower halfdroop- 
ing in the upper half, linear or ligulate, 47-84 cm x 16-38 mm (200-310 x longer than 
broad), attenuate at both ends, margins serrulate, [he serrulations with an antrorse pro- 
cess inserted on tlv l« Jifm, m mh II it i i for ward margin;venation pinnate, 

ing freely beyond the commissural loops, tertiary nervation conspicuously and bifacially 
areolate, prominulous, midribs prominent on both surfaces; petioles slender, adaxially 
plane, rounded beneath, proximally swollen, to ca. 9 cm length but obscurely distin- 
guished from the lamina and occasionally with the decurrent leaf base nearly reaching 
the stem; stipules linear-acuminate, typically 9-1 8 x 1 .5-2.0 mm, falling early, costate, 
theribexcurrentlvpiol in i t ii'U i hi i ' t , encecauligerousorramigerous, 
cernuous, raceme om a rachis 2.0-3.5 cm long; 

peduncular bracts stipuliform, to 14 x 1 mm, involute; bracteoles minute, linear, not or 
barely exceeding 1 mm I. n ill | I , I ' mm irtu ulatedatthe base.F/oiA/ers (mea- 
surements from teh\ I, jn 1 1 f it t i i ii I i r . i nli hio vnish-purple, later 
green and red-suffused;ca!yxtubeturbinate,notangulate noralate,the limb membrana- 

with an apical orifice, rupturing at anthesis into 2(-3) subequal lobes, these approxi- 

3 oblanceolate,to 24 mm long for buds nearing anthesis, concave, venose; 

I ] i i'm nl i ii i i i 1 ' . ' 

the tube rim crowned by a frmuinu . u tl >t t .m modes ca.2-3 mm long; ovary (. 
celled, ovules several per locule, apically inserted, pendulous, irregularly obovoid; < 

corolla, thereafter persistent, basally dilated into a stylopodium 3 x 3 mm;stic 

ecoloqy. — Known thus fat onlv from the Josephstaal tr 

■ ' ; 

Etymology.— The new binomia 
Paratopes: PAPUA NEW GUINEA. Mada\gP'- vn<i: losephstaal FMA area, Guam River near expedition 
' .ir,f . lu ^-U\» -tru'iin i, tin hilt . ■ n- rhPS coordinates 9497596 N, 2801 00 E, 
50-100m, 13 Auq l'i - ih |1 r ' >^(LAE,NY). 

The connate calyces clearly indicih' up in u if in section Barringtonia, within which 
the new species is easily distimi ii ill n n I < r it is not as certain 

whether B. Joseph -.t < • • i i iin .II mi - I - m t fie collection was made along a 
forest track so the branches may be the result of bayonnet reiteration. 

The monadelphous androecium is marked by the unusual presence of both an 
outerand inner staminodial whdil lihoughtl innei cip it ihly reduced,the out- 
ermost structures are conspicuously longer than the fertile stamens. In Payens's (1 967: 
164) revision, the staminodes of ill i i n tl n il and only disposed 

in adaxial whorls. The existence or mi- 1 much, il ,r l m i e , leir prolongation in 

B.josephstaalensis.are thus highly unusual elements. Together with the linear leaves, the 
character combination for this sp p ^ Inited Although the plant's appear- 
otherwise cons ,i ni • i'Ii L t' 

The new species i hi i I ti n i ii hi i I i it , bark is reportedly 

used to poison fish in the mannei t ' - I ■ 1 - ih Ethnobotanical application of this 
sort had been reported previous "it -mi iin , nqtoniacalyptrocalyxvar.mollis 

(Payens 1967: 212). The latter taxon is identified by the separate Maian name 'gaira- 

Barringtonia !0) before reach- 

ing an impasse. Itcan I i ii Ii' II i it ill i II ing couplet between 
the existing couplets 22 and 23: 
E Barringtonia josephstaalensis 




FMA area, ak 'i i .'nit;', r < 

foothill forest, between GPS coordinates 9 

504560 N,281407E,, 

m, 1 Aug 1999 (fr, carpological), W.Takeud 



u mi tl - I h i Ii .in if pressed or obliquely 

Drawn from the type b 

patent, sometimes crisped, hyaline, arms 10 or more, acicular-setiform, ca. 1 mm long; 
minor hairs compact, rays coarse, congested, 0.1-0.2 mm long; stem surfaces early gla- 
brescent and then entirely glabrous below the leaf spray. Leaves imparipinnate, 3-5 ju- 
gate, spiral, terminally congested, sessile, 41 -57 x 42-70 cm at maturity, rugose, herba- 
ceous or papery, adaxially opague dark green, abaxially medium to light green, upper 
surfaces with hairs restricted to a costal channel and resembling the large hairs on ra- 
chisand branchlets,undersurfaces pustulate, indument lax, the abaxial hairs usually 0.5- 
1 .0 mm diam, following veins, intermixing with smaller stelliform scales especially on 
the midrib, dark glandular pits bifacially scattered; leaf rachis with indumentum like the 
branchlets; leaflets opposite,decrescent,heteromorphous,the terminal one oblanceolate, 
basipetally elliptic-oblong then ovate-orbicular, the proximal pair auriculate and 
amplexicaulous,auricles ca. 1 .5 cm diam.,subapical leaflets often the largest,oblanceolate, 
1 9.5-33 x 6.0-1 1 .5 cm, shortly acuminate at the apex, basally cuneate;venation pinnate, 
inconsistently camptodromous or (brochidodromous), secondaries in 16-25 pairs on 
major leaflets, 5-7 pairs on small leaflets, diverging 45-75° from the midrib then gradu- 
ally arcuate toward the margin, partial intersecondary veins freguently present, tertiary 
nerves scalariform or not, reticulum coarsely areolate, veins impressed on upper sur- 
faces,the midrib immersed, beneath with all veins raised;petiolule absent or the leaflets 
subsessile and costae swollen at the insertion to rachis. Inflorescence unknown. 
Infructescence axillary, emerging from foliate nodes, rachis 6-11 x 5 mm, with hairs like 
cteate. Fruits indehiscent, solitary, rarely two together, obovoid or glo- 
) 66 x 48 mm; style semi-persistent, stellately hairy at the base, gla- 
brous above; exocarp completely obscured by dense tomentum, the vesture initially 
orange-brown, later reddish-brown, mealy to the naked eye, only with magnification 
discernable as thickened stellate tufts; developing fruits stipitate, the sepals foliaceous, 
disintegrating, adhering to the exocarp, covered by appressed scales with pale setiform 
rays resembling cystoliths; pericarp woody, indurate, odorous, 6-7 mm thick, locules 2, 
each cell monospermous;seed surface distinctly sinuate in transection. 

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

The plant's stature and distinctive features make it a conspicuous component of 
the Josephstaal vegetation. Its susceptibility to proposed logging operations is unknown, 
but as a fairly tall tree species, A saxonii would probably be adversely affected by selec- 

nology.— It is a pleasure to name the 

new species after Dr. Ear 


>gional ecologist for the Nature Conse 


oject's senior sci- 


PAPUA NEW GUINEA. Madang Province: Josef 



nCamp 1 ('Kumamdeber'), mature growtr 

i foothill forest, near GPS co 

ordinates 9504560 

' K 160 m,29 Jul 1999 (fr), l/V. lakeuchiJ.Wiakab^M.Gorre^&AJnwali 

1 3,462 (CAN B,LAE); 

:\;mv; ; j:, i [[■i,:kaim.iViA::'ii<.:;K-A. mmU I Am', [\ I Ai ,NYi. 

The sessile leaves of /\. -> m untni ■ lutt Iv distinguish it from a 
ners.The thickly woody pericarp is also atypical. As noted by F 
species generally have mittle to comn eons pericarps, but the fn 

,>u ,l\ I ioi ii lu'd and i. - 1 1,, i • I ' ihi d. Mid effort to section with 
I he novelty's affinitv is to A. M/AmuAs of Borneo, but the I 

larger infructescences.thin pericai p,anb unilocular-monosperr 
Aglaia saxonii will key to fork 1 36 in Pannell (1 992: 56-57). It 

:i,VLAhl VVi ,ll!N: A iVi ■■,■■■■■■ ' . \. ■ : kv.epr.t.iA 

Understory shrub A > imtall - f | n tt f lum ul t . at the ends, twigs 

ten te mi no conq i I It n I iilm i ' u i 'I pun aril in mI 

cio .dui reddish !m n mil < lifoti i ms< nil in i ietei >lher stem surfaces 
puberulous or glabresc ent, internodes usually 2-5 cm long. Leaves decussate, elliptic or 
oblanceolate, 23-38 x 5.0-9.8 cm, apically with a short acumen to 1.5 cm long, margins 
entire, base attenu it ml il i i' u nil i nt i i nnptodromous, sec- 

ondaries 18-24 pan n ii i' i i if i I i 1 1 i in i ' ' ' 1 1 Mil the midrib, majoi 

veins raised above,more promiiii in I i reii ulaii n |i mi ikui on both surfaces; 
blades fleshy, ada-i ill >paqm er\ did green ii i e ilt qieen, frequently 

discolorouson dmiii i i i ' i i I i i M i h led underneath; 

upper sides glabtou mi m i | ul,r i ul n I i ule-, furfuraceous on 

pi i ik ipa veins, otherwise 1 appressedly scalelike h.iirv on the remaining surface; petioles 
2-5 cm long,pubeiul' i ii id -nil < tunnelled oi pl.nn out I til nealh tipulm kijiiii 
nate, 22-32 x 9-14 mm.basally connate for ca. i/4 the overall length, caducous, often 
disintegrating irreouh l^i n in u in r i ill ; ih ulousorglabrous, 

adaxially furfurac it tl . i u« -' ■< c . .i| itu'lei i illy monocephalic, 

terminating bran< hf i mi mul il imm i pi Ii I i nt il mi depauperate, heads 
sessile, hemisphem il i I. pit i dl\ |lol i in ' mm n m . hen fully developed; 
receptacle discoid i s I mi h t I ml i u lull \ n t oiange, numerous, 

Fig. 4. Calycosia mamose/'Takeuchi, sp. nov. A. Fertile branchlet. B. Flower from side. C. Corolla limb; one lobe remove 
3 mm. Drawn from the type by N.H.S. Howcroft. 

toliths, the outer in i i d I t i i I m ' I m n i it. I i ulat, 20-28 x 14-27 

mm, usually deciduous before fruit set, internal bracts persistent, highly variable in shape 
and size: linear, elliptic blomp hroad'y oblan ■obii< I l .26 1.5-10 mm, lanate on 

line of hyaline filan it h irat In mi n m I uted by the bracts, 

glabrous on all exterior surface-,, pedicels 1.0 1.5 mm long, pilose; calyx synsepalous, 
infundibular, 5.0-5.3 x 2.5-3.0 mm, lobes 2 or 3, obtuse, equal or not, 0.8-2.5 mm long, 

acute, 2.1-2.5 mm lor i nn it'll | il > - I i i M I thioat; stamens 5, in- 

long, filaments 2.0 2.1 mm, provided with ndmmni like ihei orolla throat;ovary bilocu- 
lar, completely ml n i tin fill I I I i n h mm I i um stvle 8 x 0.2 mm, 
glabrous, filiform, simpl Id. ml mp ■ ■'.' ^ . j'< . - mooth,obovoid,8-9 x 
4-5 mm, exocarp op i ]u I II n, n I .jkihrnupn n 2, planoconvex, dor- 

sally somewhat uiepuLit but t i I nl n I i< nil- i r ii i face with two linear 
imaginations into one h seed, albumen lacking tuminations. 

' i , i '-■.'. 1 mie adapted species of mature 

forest undertones. Numerous indi i ml i -u nth- I it' t mtrvey, particularly in 
the elevational nti i il'mn i n ni m p< , jM t , , t p Inned substrates but 
i ikm one ,io i .II inind on . isoml rl .. . k . I ji nn I 

The newspeut i n i ii t III ith i i i i i ii n im I 

by collections fron ihhn lit pil n I pn Provinces. Although abundant 

at Josephstaal L > m i I ill n nm r n it th r ilities of occurrence. 

' lyn <c)\ I h« ' | lithi I mlli Is th. pi mi pn fit's I m nnnq. 
Other specimens examined: PAPUA NEW GUINEA. West Sepik Province: Bewani Subprovince, 1 - 
2 km N of Bewani al u Im t I I I i r 1 Pong. 141° 10' E, 160 

i UibtSepik Province: 

rainforest, lat. 4" 01 s hi P If 1 1 I nl tltu >■ I Pni Madang Province: 

Bogia Subprovince l in i h i I I i nl 1 1 t I i n th I r Hum tk i ih k w imp 
area lat 4 J 45' S long 1 ! 4 k i U, 1 ' ' i P I > is in LAE 57,325 (L.LAE); 

t, GPS coordinates 9497596 N, 2801 00 E,ca. 50 m 11 Pig 1999 fn fi ikeuchi,J.Wiakabu,M. 

z,&A.Towatin8-\\ 1-1 in.pd I ,,k I f-1 m. i .minkinh i.ige above expedition 

•> mil Dumnkin II , t m il it ill tl ill ' i 1 tin 1 t f 

Wlrii 21 Aug im" In n n ' , - P LAI 

fmVnosm is distinguished bv kmm leaves, mknivelv long Pun. iibular calvces.and a caoi- 

nilon s mi i mi ui I' mum u I mi If i n Ii i tegarded as ranging 

in Samoa to the ol n n kit in P i > I i | t u Ii i ttun innbnties t 

Cephaelis, and also i| f - h < under Sohmer's broad concept of 

t genus.The Jos. | h md \ I i i pi. 

. , m u ill lm ml nl I I o ih d. r in • .' ^ m it is 2(-3) lobed. 

ik III ib in 1 ii .11. m i . oil in i i leportedl li r i mil uhid 19) but 

signed genus on other characters, particularly with respect to the lor 
d the capitate, numerously bracteate inflorescence. 

The only other species of Papuasian Calycosia is the Solomon end( 
arr.& Perry, from which the new species can be readily separated by a 
-s,the most obvious of these being differences in indument, stipule si 

5) I, IVI'i M ill ][ ' 1 i i i ' mi, 

espGPS coordinate u > tJ 1 i u, 1 u i "Hi i , - u ,t 

M.Gorrez,&A.Towati 1 3.940 («o "vpe: LAE; i ■ ec: A, BRIT, K, P). 
Species haec ab P.melanocarpae Merr.& Perry fiuctibus a Ibis differt. 

Branched understory shrub, or subarborescent to 5 m height. Branchlets terete, 3.0-5.5 
mm diam.,glabrescent,subapically smooth and green, on exsiccatae collapsing and com- 
pressed, fuscous. Leaves fleshy, rugose, adaxially very dark green and glabrous, abaxially 
medium green, lamina discolorous with drying:on both sides orange-brown to rufescent, 
rarely olivaceous, underleaf indument subappressed on costae,otherwise mostly patent, 

oblanceolate, 22-33 x 6-12.5 cm when mature, apex shortly acuminate, at most 

mersed-rugose, manifestly prominent beneath, secondaries equispaced, 1 2-24, on the 

rib, supramedially arm u i> mm h mil in w| in t .ih c losing commissural 
loops, tertiaries subscalariform, obliquely directed at the midrib; domatia absent; peti- 
oles 2-5 cm, adaxially plane, convex beneath, glabrescent; stipules valvate, caducous, 
lanceolate to ovate, 14-20 x 4-10 mm, bifurcately cleft, each lobe 5-9 mm aristate, ex- 
ternally marked by medial ridges insensibly confluent with the aristae, coarsely shaggy, 
inner surfaces denst k jppi I I in i h i ence strictly termi- 

nal, to 1 1.5 cm long, ebracteate, paniculiform, ramifications verticillately developed 
through 2-3 orders, the ultimate rachillae cymose, peduncle 1 .5-4.0 cm, cemuous, all 
axial surfaces entirely white, with i nil Peeve v. in of pa pica le oi subulate hairs, 
these mostly spreading, often crisped. Drupes globose or obovoid, 9-10 x 8-9 mm, sub- 
sessile, nitid green, opaquely white when ripe, exocarp glabrescent but with lax hairs 
persisting at the apex and has ^\ ,-■- u I pi bur, it- 'S 2, equal or not, 
planoconvex,iacking dorsal nee, w I | mi ibi citral lumen, ruminate by irregu- 
lar transversal folds. 

Distribution and ecology —Psvtii w , i ( mall tree growing in stands with 

multistoriedcanopv It i ipp w nil » n h I t. > tb« le - | ,b n n t and was collected 
only from the foothill zone. 

Etymology. — It is a pleasure to name the i | itt < i < -. >nez,a biologist 

currently serving with the Washingtoi i free Ml Jature Conservancy. 

s: PAPUA NEW GUINEA. M.-.v.-,.; 1 - .. -. ■: Josephstanl I MA area, along trail to Morasapa Wof 
ion Camp 1 ('KumamdeK-i n ,1m- n , •• - «r ill, n-st, near GPS coordinates 9504560 
37 E, 160 m, 30 Jul L tr e & A. Towati 13,585 (BRIT, LAE); 

>taal FMA area, Guam Rivei ne.-u -\< hi i I i I p t rut, il growth foothill 

i, 12 Aug 1999{f() l WTakn ' 

Among Papuasian representative i , th it I i il 1 , assembled in stelli- 

form fascicles (or also basally branching) is a character shared only by P.melanocarpa 
Merr. & Perry. The large fruits to ca. 10 mr ■ stipules are also 

characteristic of both species. As suggested by its epithet however, P.melanocarpa has 
black fruits, while in P.mayana the entire infructescence \s white. Psychotria melanocarpa 
is presently known only from Western Province, and P.mayana only from Madang Prov- 
ince. They are apparently geograi hi II aral ti i species from opposite sides of 
the Dividing Ranges. 

Psychotria mayana will key to couplet 87 (Sohmer 1 988: 24) and to a group of 5 
species consisting of P. sphaerothyrsa, P kan ^ . < dolichantha, and P. 

ramadecumbens.but is not related to those species. Because P.melanocarpa differs from 
P. mayana in fruit color, the two arc- fa if irt >n th > inn U and their relationship is 
fins not immediately apparent. 

The new plant can be incot|)i t ji i t tl i in iti i^ it bv interposing the 

following couplet between forks 86 an nuing on with the 

Underleaf indument of I lliform h r u|ui irl h nisial. Psychotria mayana 

Rauvolfia moluccana Markgraf; coll. 14262. Hendrian and Middleton (1999:457) cite 


Alocasia lancifolia Engl.; colls. 13852, 14097, 14216. A common aroid species, but not 

previously recorded for Madang Province (Hay & Wise 1991:522). 

Homalomena magna A. Hay, en I i u I i i it! ^' tainty only from W. 

The species is readily identified by the : i> ml inn j tous male flowers 

:oll. 13700. The plant is a hinhlv teslm led en 
dermic, previousK hi t ti m n \ in I'm n tl ^ i >l and South Naru 
drainages near Madat i hi n in, p m u I i i^n i i the survival of his- 
torical populations i.( nui« u << twill,, . t nhst i il pw mm u i leptesents the only 

U. Gideon (1998: / !l -h ! tegaidod 'his species ,is an 
undescribed Tapeinochilos endemic to the central part of northern New Guinea. It was 
pieviously known iioni ioui locations. I he kmophstnal population extends the lange 
i uili nil mI 11 I ami i 1 1 u 'itth K in nt i pi 1 1 m K i tl i. im hi 

The Adelbert fo 'hi , m I 1 n i i u n| I h him mi thought to con- 
tain only T.holhun // i ' < m ami m 1 1 hm, o m\ of an undescribed 
Tapeinochilos from accessible tmrain is vm another indwition ot the comparatively un- 
explored status of the Adelbert Range. 

Cleistanthus sp,aff.?papuanus (Laut.) Jabl.;coll. /36/2.Possibly new.The collection will 
not keyonanycombin it u t him I mi i it in I ' >i It much resembles 

Glochidion chondrocarpum Aim Mm mall coil < > IM imislv known only from 
several specimen' o! mm I it r 1 t- , in hi i i -it haw 1978: 372-73). 

hobhlv rare. Now weoided on the mormon ode 3 oi t hi t ■■ mahlawh 

I he jostg d istoo collecWon is smoke to (;. ohowm h opwm, ixit is mmifloious, unliKe 
any of the species in tht (.- i /iomoo. ,/m ,m, ,/, ,, , lh ,, m (] toup. The survey 
voucher is also vt tn mt I mini , huh i I | i hi c G chlamydogyne. 
In its ramiflory the hwpl t , H j ^ • ■. n t t i i ■ t m Hon between all the 
preceding taxa an I tl it I m tl in mi i I present a new spe- 

cies. I lowever the » I07A) have fruits in 

dense clusters emerging near tin mm t n m u it mti it nihil t m 
lumping may eventually become necessary in this complex of species with similar fa- 
cies,so it is prudt nt t. |n. iiiiiiimmI i n i ir> mi r> -I tn. h< t ta on to the present 


CaseariaerythrocarpaSeuuwi h / 'n ,mhl. haown only from the type speci- 
men collected on the Fly River (Sleumer 19b4:87) but more recently discovered in the 
Oomsis-Gabensis areas near Fae (i.e., Henty in NGF 16501, and Takeuchi 71 14). 

The specitm has not been ogxwted m the hern'mie since trie time of the i lora 
' ' out i it hi t I it n nm i t i In 1 1 1 1 i I militarily make the 

rather than simple en 1 ■ 1 im h I i o in i < h taa! population, the 


Helicia affinis Sleumer; coll. 13997. An arborescent species known only from lowlanc 
environments in Madang Province. As a restricted endemic, 1 1, affinis is the kind of planl 
likely to be endangered by introduction of logging operations to the management area 

The expedition voucher is a fruiting collection and thus cannot be keyed on exist- 
ing treatments (Foreman 1 976, 1 995; Sleumer 1 955). It resembles both H. latifolia and H 
finisterrae in aspect,buttheappressedly puberulous underleaf is more similarto H. latifolia 
The surface scrape on the drupe is conspicuously purple, a feature exhibited by severa 
Papuasian congeners, though not | i. n >i II >uhi f am uhmg- 


Psychotria dipteropoda Laut 1869, 14045, 14200.Psychotric 

dipteropoda had not been seen t it r 1 itm n til it icdiscovery during iheTNC 
surveys. The type collection a btaim I in theGogol drainage in 1890,but was subse- 
quently lost during the WWII destine lion oi Berlin I lerbarium.The most recent of the 
surviving collections was obtained in 1907, even though many botanists have visitec 
and collected from the plant's former localities. 

Psychotria dipteropoda occurs >i r t 'I irowtl tands beneath intact canopy,or 
the alluvial flats adjacent to flov. r t n i ' m \ u I t | in 1 T elv to be highly sus- 
ceptible to anthropogenic disturbance; firstly, because riverine borders are environments 
easily altered by human entry into' Id u In! mi ,t and secondly, because its consis- 
tent association with advanced growth shows this plant does not flourish in serai situa- 
tions.Though not a rheophyte, P. dipteropoda is apparently adapted to conditions in the 
seasonal surge zone on river verges. The Gogol and Ramu drainages have been seriousl> 
impacted by habitat alteration since the earl ' " i I tin almost certainly the 
cause for the plants disappearance i m l n I meal range. 

Although the Josephstaal colonii repn sent thi only known occurrences of the 
species, it is moderately frequent within the Guam drainage. Most sightings were of ster- 
ile individuals, but could still be identified because of the undulate blades and the plant's 

The ripe fruit on P. dipteropoda n , - t i Mow Fruits of Ptalasensis have a 

a P. dipteropoda yellow 

apparently U 

Psychotria sp. ?nov.; colls. 13451, 13756. A vining specie lh> h it ai < , a '' t 

Utll I I I 'I I HUM 1 I i Hi | in III 'It ll( <1 lu \ ill 1.)M I I I 

Pi losephstaai collections are unusual loi their e itemely membranous leaves and 
the oblong cystoliths densely marking all surfaces, including th( infloresccMi In II o 
ers are small, sessile,and glomerulate on lax rachides. Although possibly new,the status 
of this plant can be definitely established only through revision of the i ng tuxa 
Versteegia grandifolia Valeton; coll. 13405. A rare species from the pachycaul alliance, 
previously represented by three specimens from West Irian. In their synopsis of the ge- 
nus, Ridsdale et al. (1 972: 340) had specified the plant's distribution only as 'mainland 
New Guinea.' Colloc ii n '3405 kc\ directly to the above binomial and conforms pre- 
cisely to Valeton's (1 91 1 : tab LXXIII) plate. Although Lae Herbarium has no specimens of 
this species for comparison, the taxon's distinctive characteristics permit identification 

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

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


Wenzelia dolichophylla (Laut.& K.Schum.)Tanaka;colls. 13594, 13623. Wenzelia is a mem- 
ber of the subfamily Aurantioideae and remains imperfectly understood despite the 
potential horticulture I ilm ne ■ I Ii i> n p Only six collections of W. dolichophylla were 

Swingle (1 967) established two subgenera but was unable to assign W.dolichophylla 
into either one due to lack of adequate material. In the ripe fruiting specimens now at 
hand In 'in Josephstaal the 'hit seeds it irregular hyaline i n hi make it oeat thai 
W.dolichophylla belongs to his subgenu , 1-15) did not pro- 

vide a subgeneric affinity for W. dolichophylla and expressed doubt whether Swingle's 
seed characters were sulhc lent foi tecognition of subgenera. 

The Josephstaal plants have wide leaves with prominent and anastomosing sec- 
ondaries similar to those from the Sepik populations. However the bicolorous blades 
with divaricate laterals otherwise agree with features more generally characteristic of W. 
dolichophylla.Jhe red hesperidium (salmon pinl <>n a '- mJ i nkish-red on 13623) is 

dolichophylla originate from the Ramu-Gogol drainage, it is very likely that the survey 
vouchers are correctly placed under this binomial. 

Zanthoxylum conspersipunctatum Merr. & Perry; coll. 13636. The tree is a montane 
species from elevations above 1 500 m (Hartley 1 966: 205) and is primarily known from 
the Highlands Provinces.The expedition gathering is a first record for Madang Province. 
More significantly, the elevation of collection at ca. 1 60 m is anomalous and represents a 
significant extension of the species'vertical range. 

The Josephstaal voucher has a number of atypical characters which initially ob 
scured its generic identity. At the time of collection, spines were not visible on the 
branchlets,the leaflets were epunctate, and the foliage showed no evidence of resinous 
content. In sicco, spiculate excrescences only became evident after collapse of the 
branchlet.The taxonomic concept for Z. conspersipunctatum sensu Hartley is that of a 
polymorphic complex, since the species consists of numerous distinctive forms which 
cannot be assembled into discretely r 
men keys to Z. conspers 


Antidesma katikii Airy Shaw; coll. 13729. The species is represented in herbaria by few 
collections, having been discovered only fairly recently (i.e., 1968 by Katlk in NQF 32762). 
All gatherings have originated in the Ramu-Gogol basins. During the 1995 Bismarck Mts. 
expedition, large populations were recorded and documented near the 600 m level and 
the species did not appear to be as rare as the small number of collections might sug- 
gest (Takeuchi 1999a: 763). With its discovery at Josephstaal, the distribution of A. katikii 
now extends across both sides of the Ramu drainage and the plant is certainly more 
common than previously supposed, though remaining endemic to Madang Province. 

The recency of its discovery, and the uncertainties over the conservation status of 
A. katikii, are circumstances applicable to many other Papuasian taxa.This situation is a 
natural outcome of the uneven state of floristic exploration in PNG. Botanical collecting 
in Madang Province has been heavily focused on the Gogol and Ramu basins, yet the 
Josephstaal populations are within a mere half-day walk from the principal coastal high- 
way. Clearly, there is much work remaining to be done in the floristi 


Microcos sp. ?nov.; colls. 13469, 13562, 13732, 13830, 14104. The Josephstaal specimens 
apparently represent a new species distinguished by a bilayered indument of erect simple 
hairs with smaller stellate hairs underneath (most Papuasian taxa are lepidote).The plant 
has been collected in several of the north coasl distrid and possibl> also from the Gulf 
region of PNG. Although undescril ed, thi ittril utable to taxonomic neglect of the 


EtlingiT.i . n ^ ies Polyanthae) 

genusbyRM t nit 'St m 1 n in 

Most of lot- kapuasian species were tt 
were depu u il i II n II i i as 

The flowers of the Josephstaal cc 
in- th i r il i li 'I in mm's. sense Valet 
i,l • ' i i .1 it i * the top of the : 

homm m 

Although the Gogol basin near Josephstaal represents one of PNG's 

healitu mm i i t it mi h i It in Oi n i -I I. im k il in recent cleca 

logging. During the severe drought of 1941 ,i naps f mi t of the n e J 

fire (Johns 198b 351 it is iea n I k im thai, a q iti mt but unknown pari 

ofthe former flora has already U . n hint h I \ ill I i | i t < vents are consid- I- th. I ii n M km kam em lining near Gogol, and not unex- 
pectedly contains many taxa recet litem I Rti il h f II iccoids for Psychotric 
dipteropoda Rh\t i , < n 1 exemplify the simi- 

larities between the fl ti, m i i i Hi I miR iti an be understooc 

in terms of new evidence showing thatdistnb 
lated to qeohisforica phases of lenane accret 
the basis of the geological relationship, additi. 
Josephstaal and the adjacent Gogol-Ramu drc 
of the same accretional phase in New Guine 
thus ivoie likely to he rediscoveied at Josephstaal th 
cai habitats m tin 1 Madano Ramu area. I he spin ies k 
tions dating to ca 1'Wn mil l"M k)l Du\l|i I"m, en \\ uh further exploration of 
the project trau otl i i i iJm hi mi il k also befound.There 

hienu mifiihr i t i plo Is mm nil -j \<lalesiana series, Balgooy et 

al. (1996:198) conduce M'ih> in > m I i Im m | ' I n 1 1 \< sthern NewGuinea 

h i j h i t h in 1 1 1 1 1 1 1 1 i 1 1 1 1 1 i t II i 1 1 1 1 i 1 1 1 ' h , 1 1 | I i 

Miisot KeM' Guinea emlenm sarccone 
a (Balgooy etal. 1996:201 and fig. 16). On 

anal linkages c 

11 I" ,111th If Mt' U i. IM t . 


e the localities vveieal pan 

a | 1- 1 t • 

■y. iiiiiiCiCt no; oeeMeeesi's K 

I i i i iik jii eyed p.i'ts ik i 

unknown taxa. Th 

ie discoveries from the a 

Mil Ilk ui i \ an mi 

lelbert's position in 


M 1 id 'Ii- i i i ' ■ i l- 111 

orthern New Guir 

lea as a whole (ibid). The 

re an ecneia gi k ( i 

ture exploration v 

vill yield additional discc 

mm . ; i .ill in i- M ' 

ait Josephstaal he 

ive not been examined. 

Most I'apuasian endemics 

ties, though from! 

the past emphasis on mc 

)ntane exploration, it is ap- 

parent that the higher-elevation percentages are overstated in relation to the lowland 

partly a conseguence of the poor attention historically devoted to lowland environments. 
Because many endemic taxa are shared between the Gogol and Josephstaal locali- 
ties, it is natural to ask why the new plants had not been found during previous surveys 
of the Gogol and Ramu drainages. The novelties are visually conspicuous and with at- 
tributes which should ordinarily have ensured earlier discovery. If the new taxa had in- 
deed once ranged into similar habitat'-- in the 1 Gogol-Ramu basins, they were probably 
eliminated there by the environ il > ■ . i >ttne I Os, otherwise they would have 

tions ever being detected. 

fhe l \ urv. p >in1 to a m I ill il ,i i i eph taal nvii m nt it luoi 

his presumed status. However the percep- 
arising from the recent surveys also need to be weighed against the madeguacy of 
ng information on tht- Nr nut i i , I - t t I , status as a center for biotic 
sification,PNG hasthedulnc.u i . 1 1 r . in t in i u il'.'ii la's worst surveyed 
ins (Stevens 1989: 127). On a regional comparison, only ihe Celebes and Sumatra 
comparably low colleclic in il i I I u n m i HI needed that cer- 

nountdii u in i< at- ho' [ ' ( i II i I i i in llu eb atic o Mt 

ot easy to identify. When doci 

? that presumed connections between Josephstaal and adjacent areas might be 

formation weie a\aibi >n th. uuounding region. 
A total of 1 39 families, 445 genera, and 730 di< 
d at Josephstaal (Appendix 1). An unknown proportion c 
mented.The41 alien species (exclusive of cultivated plan- 

i the Lakekamu basin and Bismarck-Ramu Range were n 
protocols (including attention to weeds) as the Josephsta 
'alent search investment of one-month duration. These \i 
en naturalized count of 8 species (i.;% of the total) for Lakekamu (Reich 1998; 
Jchi&Kulang 1998), and 9 species (1.5 %) for Bismarck-Ramu (Takeuchi 1999a). At 
r Mt.a more intensive surve\ t i i iln , 3f - ipi i I o°c of all recorded 

itably integrated inlo lutuw schedules ,o an indopendont means tot determining the 
relative quality of evaluated habitats, nvvstioators have an understandable preference 
for focusing on in ii i t hi n i t nil I ut tl I un n i i tu he pr 

vide a more practical indicator of habitat preservation and isolation. 

None of the vm d r i , i at i [hi up m\ threat to the environment, 

with most of the mti lull mi i I m je herbs restm t i to repetitively dis- 

turbed ground {Piper aduncum excepted).! he presence of anthropogenous plants is an 
unwanted condition in, up hdiine bur ibr , n c rarer u <>s am inevitable when- 

units comparable u I | h t i ihr n ill im-lti Immi in nls and thus logis- 
tically difficult to botunm Ih o es I il t t u I Inih >lu< habitats is a marked 
contrast to these tl i ues Ith jhpioximitvi bar nter causes greater ex- 
posure to unwanted I n im 1 ma ■ i i n iti 1 i n • .id i ommunity-based 

development, bee ause of thi omparat mi I ( itior it' >rded by convenient 

i i Tin i miiI in iti i I i Ii ill t i ill i i n tu will enable con- 
sideration and impletriMiiation of a wide tanne o! planning alter natives for the project 

■I lib ■'• I 
Voucher source New Guinea Force 

senes,P&OBS Parr nil t It hi In l r ihl r tiez and A.Towati, 

T&S = W.Takeuchi and E. Saxon, sn - sin numero (without number), SR - sight record of 
taxon known to the projm l botanist. Other collectors indicated by name. Determina- 
tions by Wl unless otherwise noted. 



Adiantum philtppense I .; T 13436, 14254 , , , I limie) J Smith; T 13919 

ASPLENIACEAE ' !l '<P lt " j(1 drr ' ' ' ' ' ' lunu ' mrth;fh( 

l ; " ,ll " " , ' .]l()!l|> ^ l{li> 

t opel.iriil I e| - o >>\ Sledge 1 K> r "l ! ' > un 'I ' ' "' 

408); T 13490 BLECHNACEAE 


group'; 7 14255 DAVALLIACEAE 

Asplenium nidus I iininh' ':> Davallia solida (Forst.) Swart 

. - " m 1 "M I mi |i > i <■• e throughout aeo 

/ )//'/, >!(} d'urvilluci (iii 





> < , ' n hi H ' 


ui folia B\ume;T 



->retusa(Cav.)Maxor);T 14138 


v.?/!.; k icju.j.; c"hi---:n; r ? -?->- - 



un\ m,„,, ( i , hi t | u , n , . 



hilotum nudum [ 

i Gaud.; J 13626, T 13953,, 


cies',T 14256 

'< ■'••,■>;, . llt | .,m) ' 

, >U / I llll I ' - C\.l 


/ygort/u/n M '>07).'iL.j/(jrM!ikjrm.f.)! 

Lygodiumdimorpi , pel.; 7 


- ,uJ ' < M [I 111 ' 


Nephrolepis biserrata (Sv 


. Pleocnemia macrodonta (Fee) Holttum; 1 134'. 


U'ctanodiiiiiuununudy^nst.) C. Chi.; / /-fOC 


/(■•( tariann-Tiyiininidi, I'^sl; CoueL; / / >■•/'>••/ 



lA:, J 14299 

Icdarin rppundii " i 1 Holttum; / > ? 

'urn L; SR, foothill 

7ecfara sp., t jff. ?/. v,-j.«. ., ,u, 7 , G (Aidcrw.) C . Chr. 



: a'es (Hooker) Roos; 7 



Aglaomorpha heraclea (h 

;unze)Copel.; f/40/0 

(Gepp) Holttum, 



;um (Cav.) Wagner & 

spoouonfop/nno p knon to) uunnfn/nnno 

1 \ 1 -.1 1 1 ^ 

/im'mi/:;; pomn 'movo; '\Mioio / /P.'/, 1 

^^^|/)WJnfws5erlSLl^nL) 1 ^V/nM;^;/>/! l ^nf/mLi//fc/M 

il ) 1« 8402 

iNues)hVill.facies;7 ?3889,7 14/47 

74007, 14144 

•/ * ^ '3S87, 78679 


Saurauia conferta Warburg; T 13981 

\pluiciastcpluuh>\ unilus (.! .) Moltluni vat. 

luhtuin So mi a i\)\< 1 i)i iff '420 

s,;i/; i / l 'ii,is|o,seriO':-f )(iu;//u/ l ;o l( ill,'Oi : ; PopP/nOn 



, ' ■ , - • t .1. ■ ni ,h 


1 )54; 605); T 13902 

ferrugineum (C.T. White) Bloembergen; f&5 

, • ,' , < >■ > ! lit 11 lOJ / If 




Amamnthus dub/us ThelL; T" 73874 


' 1 f inn 1 In mi 1 in 1 

is hi.mam nina 1 ant SR, ridge near 




alsoUGV/onnoPo dPPS dot. U. rrodin 

GnefL/mgnemonL; 7/3758 


i ,,'ir/ii/?) inu/mnoino 1 i n in i 13439 

/Pnsnou/no Guiilenn. P\P. While in NGI 00300. 

(,nrtumsp:,K.I.WhilrinN( l l i0005,O\ec\ in hoi 

dot. Camp 2 

barium log but not found at LAE 

,' i op ' u -in < 


, • \ ,'i ^ n< itk us K Schum.; T 14217 

' " i i . 1, 1 , il mkb 


Podo, irpusd.rumphHB\ume;T 13492 

Cananga odoralu 0\ n i ' ■. inn M Su, 


Cyof/ioca/yxpopuanus Diels,or aff.; T 13507 


"-chum ' 

NU /ddu'8, dot. K.Saileh 

\p\ i/ ", ," : : /m ■ i ii V ;7&S '3030. 1 1084, 
T 13720 

Hon' /jt?4i). / WK. N'PP on pp ! 4 !86 
oonoo/wnons p;> .; /.SS 000>8J H6°5.!4Q93 

'hio ■ to '07-'' . -n ooO l.S iieffei an 
Heusden; 773835,74233 

i i l "i ,, / L.uim \ ,1 1 ?n . ) 

i ; , o >p/ o |i ill ' . i, ;nn \ Schum ' 
Z8754, 73769 

/ , | iff , ,, o I 111 i. hldl M7 

, , . , ." ' 1 i ii \\ n' i ' 

Pu'nPoouoo spu, aff. ArnMopo' (Diels) Men; / 

/SrndennPnmnol pO i '. 1 ,;'■■'< O'So Q:Onp'M.M'i:,U 

pJ d »l S ,A;r, 3 75,.,i S 57., W 6,,M07» 


Pseuduvaria ^p.B; i&.. 

S13054J 13423, 13733, 



akiana Martelli; 7" 7 



aria (B\ume) DC :,1 


cf. Xylopiasp-T&S 1 

'3064, also KJ. White ii 


Blumeo rip, 





.J 13907, 74207, alsc 

,i Wh 



1 - [hij ni - Lit 


^MariTrafT 1429 


itha (K.Schum.) 


KosletJ&S 13085,1 14761 
• 'ri(l)DC T1380i 

horsier (1992:528 529); i'&S 1 
Voacanga grandifolia (Miq.) Rolfe 


. < • . ,,'"•>, I l . hufn ii 
iiJt TO. ?50t57 , ^tT 735 f 
. ' v . -in ( - , .i Takeuchi; T 7 ^ 


Osmoxylon (closest to) st 

Philipson; T&S W353-BJ 

Schefflera sp.; SR, high epipp 

Pararistolochia schlechter 

' 13536.14789 
• in NQF 10323 

./R.Br.; 5 74229 


Bixaorellana L;SR, cultivated 


. - ' . ' 7 L 5R, foothill forest 


r ■ ' . ntosa Lamk; 5 74 


■, , ,tffo//L/m(DC.) 

NGF 10254 

'7'indriumviiiense A.Gray; I&S1310- 

i ] « "t/VGF 70325 

' "3\indus (K.Schum.) 

■ " iL indihl. 


l-\uihuuiiiimpl(i Span.; 7 14044 AlsomitraimiCHxarpa{b\un^) Roem.;SK,( am 

,< > honduc(L)Roxb;J 14042 3,cf.PullenW96 

( l l t ^, l lin!!:iil S'jm.HhlU.I W I I i I It < I III Ha p / 13871 

from Camp 2 Trichosanlhes spjonqiflora-bracteaia group' (c 

Harms 1925: 159); T 13910 
Intsia bijuga (Colebr.) Kuntze; 7 13692 


tWGF /2095, det. B.Verd 


W/7orumUvnh.;/ i WW, M(?(W 


/Wan/7foosc/ieffef/K.S(hum.ftllo!liung;7/.W ( ( , _ viiq.) Martelli ex DurW 

W8W '40/ / I (1 , p., ,'^s W050,7 Z4268 

CARICACEAE Tetracera nordtiana F.V.M.; 7 /3905 

Carica papaya L;SR, 'naturalized DIPTEROCARPACEAE 

CARYOPHYLLACEAE torta ^sso/c (Korth.) Blume; 7 /4208 

SR,Guam R. Diospyros papuana Valeton ex Bakh.; 7 1357} 


p,,,/, ,7, ,.,,,,.„, ,„■,., ,,7!iW)//,'<7M > Wpp. ex Miq.; / Wi'MswmprMW WvWWikW;/aiWW7m/!i'VW /(Wo 
13804,13977 " Diospyros rostrata {Men.) Bakh.; 7 / 3764, det. r- 

Miq.; 7 /404 /, also KJ. tWW/te /n NGF 10244 ELAEOCARPACEAE 

CHLORANTHACEAE H(k , 0i jip , , ,, 
Chloranthus erectus (Buch.-Ham.) Verdcourt; 7 Camp 2 

'^ 75 ^ Sloanea oijcici Wet t 


ile individuals throughout area '',, ( 

Garciniaa.celebical:,T 14159 (Waihutu m • 

111 « 'n/VGF/03/3, ^^^ 

det WW. Stevens n«gp 


, hum 1 1 ' u - i pW im (I ) Blume v 

det.M.J.E.Coode moluccanum (Decne) Muell. Arg.; T 137- 

Euphorbia heterophylla L 

Vcrdcourt (1 979:304-305); J 13823, 
Mucuna cyanosperma ICSchum ,1 
Mucuna novoguineensis Scheffer; T 14174 
Phaseolus lunatus L; 7 /4257 
Phylacium bracteosum Benn.; 7 /3909 

quadriglandulosaj 13960, 14062 
Macaranga subpeltata Laut.&K.Schum.;7 
White in NGF 10260 

<> ' t ^iltiqlandulosa (Reinw. e 
Reichbf.&Zoll.; 7 74036 

Omalanthus novoguineensis (War 
Schurn.; / 73956, 14071 

? F.M. Bailey; 


-; T&S 1. 


Casearia erythrocarpa Sleumer; T 13481 

i ni Kill I )i ,]ff 

Casearia sp.?nov.;T 14150, 14173 

h\thi >,t i m (Becc.) Becc; T&S 

/306ft T 13495 i\ n i 11/, ,, i . 

and /0326 

rm/sRoxb T 13502, 14176 
ht>i ,,< " ,, (F > b) Bomh I llu i 

f ir i II i t „ < ni on J 13940 
Osmeliapfiiiippina ( lure:/.) Ronih.; / / 392? 72Mn, 

74090, also K.I.WhitcinNGFlOPSl 

u <" i " ■ Remw.; 7" 73646 


Cofy/anf/iera reniv/s Blume; T 13524, 1391 1 


;/ Warburg; I 

siphon;'! U/H, 


; DC; T&S 13090, 1 


Polyporandra scand< 


in NGF 10298, de 

Pseudobotrys dome 

Rhylicaryum longik 


3; SR, foothill forest 

>.t u< > , , ,'! blLI IK .4<X> <> 


■• ■ ., ■ ■■■: . ■■■■,, ^ >jiss.; 1 / vn.<e 

///!>.■',, ,■;-. ,':/k7^L^ 1 .;',!■;, Hansen )1 Wanudah 

Wagadab transect 

inNGF 10297, det. P. Fryxell 

1 1 r 

\ ' • , , 1 h tii ► |! 


V(,/ 10 h '0 (k.dets A Kostermans 

hhliiii idrciiiii mtliik i'ijTost 1 n irkvkkkfr/N ,k 9 

70269, det. A. Kostermans 

11 m| itu, Ml 1025:116);? 1 341 S, 1422,3 

1 ' ,< - T'l< ' . . ' ( , h (' m I'-J'^' 

Akk'ik; sp.,ak/ t v; :i v):' ( ;ukk!u kikci f.; / e?5oP, 

i>Vh,ti \ '77 l.lyp< ' '- 1 ". >:< l \ 

736 7 7,73867 

/n./MMr/rYJ wiMm^-J h t t il 

' 1 ik ^ 7^lvns1625 

1 1, 1943 439 <~&3 73052 

Otanthera bracteata Korth.; 7 kkek f477/ 

genus indet.; 7 73763 



kkr/e nr^jloma-.u-.i km & Pirv, -.'' kikk 

form; 7/3685 

Aglaia argentea Blume; 7&S 73077 

, , 1 ' ' "■ )\ ' - < 

I 1 uin l 1 mt Tl l)t> I 

WGF 70248 

, , ,,. Mim , i l4 70 13584,13694, 

1i 1 J 14032 14201 



lihjuieaceilanica I hunk; 773673 

1 Ikms 7&5 13092-A,! 

hnimca elliptic n KoxkkkJoothN forest , ,inu 

k^emexo/kkkeukiik 13462,13712,13765 

fegraeo racemoso Jack ex Wall.; 5R, alluvial for- 

Aglaia sp.,aff.agglc e/oraMen 1 m T Hi 

vai rupeitnr.l 13484, 13538,13593 

Aph,iii:i!)ti\i>noiv\hh hvii (Wall.) K.N.I'aikei. 7.^ 

131W.T13501, 13582,13736 

'1 [ - tk 

WGF 70253, det. D.Mabberley 

0299, det B ( onn 

C7p/M)Ji.'ro/i/ov(> l( j//>us(Miq.)V.)k>ton;7,^"73.n.' 



13806, 13827, 14056, 14247 
, >■ L m. vnhile Harms; 7" 14298 
Dysoxylum sp.; 7 74306 

C -' .- . r > iva (L) Merr.;79376 7 

i - >>etiolare Hooker f.&Thoms.; T 14047 

1 r I I lit lit I 1 I 

Archidendron bell 

Archidendron lucyi hv.M 

Entada phaseololdes (Li M( rr.; 7 '360 

Ficus hystricicarpa\Na\\m\ , ^ 

NGF 10308, det.E. Corner 

9/cusp/iaeojyce Laut.&K.Schun ' 

9/a/i polyantha Wai burq; /SS /3 703, 3 37 76 / 
Hcusprimaria Corner; K.J.White in NGF 10274, c 

Ficusprimaria Corner, or aff; 7 74/62 

1 ir hint T 13880, 141 1 

hicussubulata Blume; / V 73993, ~, •« 

9", ■ "< ^ - j L tb Corner 
Reus W0550 Roxb.; 7" 73549, 73760 
F/cussp.A,doesnotkcy;7 37099 


s iS£ 

rsufo (Warburg) Perkins; 
3sp/fans (Becc.) Kaneh. 
3 73822 


c, P iensK.ScU Um ,mS 

/39o^pu5 CO, 

Timunis J.R. & G. Forst.; 

>rachycarpa de Wilde; /C. 

" -'i 9 \ \</fpo Miq var. sub 

(Elmer) Corner; 9 13634,1 42.25 

" king, 973993 

Corner; / /49/2 

Miq var pnteli te> ~ ,tl m i 

■'/-/sf/ro cylindrocoipa ,. Sine:!.; 9. 7 C'hfe 
97333, dct939nd9r./ oh. Dnoretnar 
','75 < /a; //.si /39/o d c W h o :-, s p./ o s /'7/ora; T 



,,. [ ,,,| ,.[• 

T&S 13106 

'.hi, „;ih> „i -s., 

Mez; 7X5 1309/, 


/ \\,ispamum hn 


lh\aspermum nc 

,' >>'>', ," 1 ,HH. 

/As 13087 

ianniiD\e\s;l 14 

^■/yqium aeoran 

act - 

, m,'- cf. amp 

lum Hartley & P 



Opiliaamentacea Ro 


Adenio heterophylla (Blurm 

Wagad ib 
Passiflora foetida L;T 13961 

Piperaduncum L;SR,Guanr 
Piper betlel:,SH, cultivated 

Piper caninum Blume;7 '13t 

i hvbr'ivkim (I nit K. Schum.) 
Retry or aff.; 7 /35t il i 
' i' let TG Hartley 

Syzygium longtpes Merr.& Perry; T 13875 
, ,■■ " j 7j'\?en<;e Hartley & Perry; K 
mnS '0300 (type) 
Sy/yqium nutans (K. Schum.) Merr. & F 

,, ,," p i ■ '/'"ui,''t ll aut h\ k Si hum 

h ' ; 

imboinense C.DCJ 14084 

:.DC.;SR,GuamR. ' 

,• | ' ' / ptohablyconspeci 

'.v.'Wj/t;tf! sp.; K.I. Wiiiie it; N(,i 10? ^9, cited 





branifolia Bartl.ex DC.;' 

Valeton; T&S 13098, T 
ychotria mtcrococca 
Valeton; T&S 13088 

Kalkman;7" 13842, 1411 

13449, 13504, 13969 
13082,1 13551, 13586 

i .'■■B:29yJ135h 

Valeton; 7 13930, 141 69 

i i ■ Pcivon.; T 14153 

^j^r-rcauichiincina (K.Sc hum.; Valeton: / i.^Si 
13749, 13788, 13864 

<s Becc .; / 1AQ05, 14.W3 

• , i ' let S Datwin 
orinda bracteata Roxb.; K.J. White in NGf- 10?: 
orinda umbellata L. var. papuana Valeton; 

ussaenda cylindrocarpa Burck; T 13627, 1370 

I&S13061.I 13460, i vm 
l rrr ) /(Vi/!Lic/ra,'!V/o//aVdlelon; / 

> i '0 ^ > > 



Sfer ^|^oyen 


ounctatum Men. & Perry; 

J3542, 74096 

ma (laut.) M\\dbr.; T 1346 

/ Shaw; T/3729 

i Beus.;5R, foothill forest 

Antidesma rhynchop. 

y/umK. Schum .;f/ 4033 

■J 13897,14165 


Phalcriacoa nca (G 

aud.) F.v.M.; 7" 73499, /35C 

:um (Laut. & K. Schum.) 


W37, 74 7/5, 74202, 1423 

l 13469, 13562, 13732, 1 
Vhite in NGF 10231 
vteinNGF 10231, 1027. 

■la Rad\k.,or'rigidiusculo cor 

nplex' 14156 

1994: 593); T 14305 

iceaRadlk.;7 13681,14307 
'ora Rad Ik.; T 13759,1 3821 


„„, Celtislotifolio 


I I I ^ 

i, ypr,oiophu\ d.numtmilatis Winklri; / J404 lJ 
: . ■■.;■ i ' .;■ . ' -a//< ',;:".'":■ (I aut.) Chew; 



' ,' „ " •, ' '.inMi i < h- . ' •' V 
/\7!f/fD, .'!/,/, ■f,v/ !l /f l 'n'./\(Mi(i.)tlK'w;/ / M^ 
Elatostema cf. beccari/ Schroeter; 7" 74055 

I Lih)^ci!iam.k!n!>i;vl!un! Rionqn.; / 142/4 

. ,l ', lUhn < -I 

111 II 'i ' 

"' " I ill/. . I iii'. I 

• hott T 13580 

:-J::::.'i)idophoi\r,rr',!o.\i.l; ■-;qiH. m^ 
i! - <• H ",Vio7vGF 70290, det. I 
Rhaphidophora sp.; 7 73707 

ii - f sp AT 13410, 13876 

■■u>n\\:.:ndcUK.I.WI)iti'itiNGh 10246 

Womersley,alsoin , n 



Mm ui in r > ' nimushollrungiiBecc 

■ 1 1 n ir I h ti ^ , peditionCamps2a 


oso (L.) A. Che- 

, c - l: :;,;;; ; . 

albertisianus Becc.-J 13641 

hollrungii Becc -T&S 13059,1 


hiana Man.;SR, throughout | 



7 L.;SR, cultivated 

fa (Becc.) Becc; T 13522 

i ;;■■/< ' '. 

microspadix (Becc.) Burrett;] 

. - 

»pe//7Blume;SR l Guam R,ste 



Aa/aonemo marantifolium Blume; 7 73534 

T 13643 

■i.^ ; .is,'noec?u/7o/jLvN.i.lir.;7 13595. 13854. ! iF- '.'. 

il • I-:- - ' ' '- det A. Hay 

T 14252 


K.J. White it ' indf i. i 

Amischotolype mollisstma Hassk.; T 13 

« - T ' i ' .' w f lull ' 

Andllmavitiense Seem >T 13885 

■Vn< ( ^Wand/cj//rjln.|l.;/ 1W2, 1409/. I471t> 

Aneilema sp., aff. Ihumile Warburg; T 13860 

Alocasla lauterbachiana (hngl.) A. Hay; / /3S46, 

>>",,\ ' <n thi82 

t , 1 II i 1 , NJ ll - '/ <f 

Amorphophallus a , ' < , 1 f 1 ' i H - t • 


qcnus indet., but probably Floscopa scant 

■\.'?.>. )!pr,ophailui pueamiUiU-i (Dennst.) Hie or,:- .; 


SR, road to Roumirap 

t ' fa (1 ) Schott; T 13640 


>hh/hioiiiysb£cair-:r t r.q\:.l !3478.1385i.i.i-;/-^ 

Costus speciosus (Koen.) J. Smith; F 73702, 74 

Tapeinochilos recurvation K. Schum.; 7" 7370( 
i ' ' ' f uov 7 13/43 



Dioiionuiescuienui (lour.) Burk.; SF 


Drui >• < < ,ingustiloih.i Roxb.; I ' - 


,;IS.\ ! 



',',,,' ^ ,i\ )()L / ' M 


" - ,, / , ,'\'< h .1 hell.) I . 
i ;)??!/.';-,;,; i. i.f?)!,';o.' V.ilinon; / / ^ 

I J.'srOO'tJiiVw'/j/i 


. U lhh-,U.JIr-'M 

/VmrWu> ?/)W/)n, 




r ;4008 

< " t - 

■ ," . '.; ' : 

1 low i mil; / 

/3403, / 



' i ; 


Zeuxine polygon 

uihb :s 

genus indetj 13505 

< :> , lihpiiHohtum '■-, - 

-; ni;j,'/c6' (lorsk ) Stapf; SR, Guam R. 

1053-A1 13568 

Pogonatherum paniceum ( 
Polytoco macrophylia Bern 

ria ot 



- i" ' ii i \ ' 

/ //;■'.'!, y : j .'Y.* sp. 'nov'.;si."i.", / , ;i; , v ; . , .')!/]je;; / /LV,'^ 

Smith; 7" 73406, 7346: 

'I I I", 'I 2 

;. Spellings are phonetic. Several taxa arc apparently r<. pi im II Ml n t ' 

mtheMaian Ian iu i i | il ill n m i in i II kuai n 

Scientific Binomial 

r ':!>.'! .leruniorrjr-:: 


Aglaia lepidopetalo h 

J/u, ; mv iTjk.'u. 


(Elmer) Corner 

la ; lago 

Jsp/tn r- ct ,->,/ ,- 

la i ■ la |i 

dago lago! 

//pun's i ondylobalhoi 

!' , 1< 


W i , it ' I 1 i 'i 1 ! 






etr . • tin , iuU " h h a m Cii i 

: ■ iang la| 


Morinila umbellate! 1 .var.pjpi/u/ifj Valeton 

Rub a< eae 


, ' ' < ' 1 alii l.l II it >ll 


Begi n at i a< 

Muii , llhiir 

\g } i lgg >m< 

■l;',-;iS,Si inipClieli: 


Dysoxylum sparsiflorum M. 



•neg>]!i'<r>i\ (Poir.) I <. i 


Aphanamixis polystachya 
Psychotria amplithyrsa Vale 
Elatostema novoguineense ' 

StvjxA i///>ui magniti 

ll.i. MJ'ti 

Pseudobotrys dorae Moeser 

,,. .N 

,. \W 





pat ciay;:l ;:a.:ol 

■ ■■ephiiummiaopinrnitum Hobtum 


■ • .'-irif:/ tinqwiib van Royen & Steen. 

DIM ■ :■..:- 

'.' . kinitivtii debica (I laniv,) Pbilipsnn 

. t !f ' Ui'sati 

f)r.raum gratissimum L. t :p:ik,is 

Harpullia ramiflora Radlk. 

nib... ■:■:!-- :i.i- : 1 

Cryptocarya laevigata Blume 

Begonia papuana Warburg 

Bi g in ii eai 

.." i : '■ . . , A irl ..- | 

.,".,' < < < - 11. mi 

i 'vrtandm sa, section « :'ni!OSip!i<>'i 


sa b kamb U 

Eiatostema sp„ aff. maaophyliLnv l.i mm. 


sagag gosmun 

i > H . , , ,'<■'< 1 J 'butt 

s>agag-u-yoqa umur 


;a \a |-ugo in 

-■ b a :ih ;.l 4 t v? ,•„•• ;p.i 

■', anth : eae 

: lai ar| i eai 

riacourtia inermis Roxb. 
Phyllanthus rubnflorus J.J. 
Agloia agglomerate Merr. 

Polytocha macrophylla Be 
Cephalomanes atrovirens 

Kv> i.;.:i.-aceae 
■ ,i ■ acc-ae 

I I. I' l' | '" .1 V. 

Popowia sp. 

"wihiir.ona sp. 

( nunoihtilamu!: s 


I taispungens Rcanvv. < 
Antiaropsis decipiens K 

voa h:KV: :;j'";;<rri 

Dichapetalum sessiliflon 

' ' at r' ' , ' 


■■■.kiniitoo^riicHni K.'.i hum.& H 
' ■.,uihoojiv<rni!><:'ii]ii': DiHs, or 

II It ,■>>•.■ 

widasag Glochidion granula 

1 . .uk,: 

( immo kyllingia Endl. Cyperaceae 


tndospermum mnh.i: r aaaaa^ Riititu Kuiy Euphorbiac ut' 


Artocarpus communis J.R.& G.Forst. Moraceae 

Wedelia biflora (1 .) DC. Asteraceae 


, ' ,. f < Maitf Hi Asteraceae 



ileton Rubiaceae 


1 agraea ce/'/an/'caThunb. Loganiaceae 

laxa represented by M 

aian orthogiaf hi in i It m i i utu 


i il 1 uiiequiloba 

N.E.Br.,mang g ig ii i i mil n 

Valeton,babagalum ii,] i i u >( \ n ,\ uui i iinnnnn ilnul i i 

i tnlvpemoll 

kiui-kiva, sepi-sepib ' t 1 i i G i i i i il i tionia pseudolateralis 

i ii , i ii .... h,i .In 


rubu,wo-sayep; i pi al >n im : itrovirei : Presl, ;ingi singg p, /ve len 

, t v gafoBk 

1 1 ni \mon'a-'-\i Mi ;.:.,:"., a -a -MMOa.'a „!:■■ ua,a;mm/5 PK7l ', a<. ; a, ; . -.. 

[ ill ii-diri f 1 i M .1 Uiii ii i r It rup ', 


mm tnaiucrni: I ,mI . da i!,i da dag, uiagu a; i a\maaMMM ruprslic J.M. G G. 

n i n I I ii Ii I I 

beccarii Engl., kolaiv, ma- 


bip, wange-warap; Lindi 

■ mv OMMu .,. 'maa g.i i d, jo ,.aan nana. • a ;:.m MMMMaGduuRa 

lasa-lasa, singi-singgip;/ 

I 111 I I I | HI ,ll ' I I - pi',' 

1 ubsp./anc 

'■Mi: | I h > ;amM. , , < mm il III h 

Phrynium pedunruhit m 1 i ill 1 mi i ihih i ' nbnflorus J.J. 5m., esg- 

i -"let bugu Gf>t f 

' I UN | I I III HI l I II I t I l I II I II I I II • I )l 

macrodonta (Fee) Holtti 

ul Mi al in ■ , ' , !'■ . ; i ■ in bun mu ul ul il u! 

osag-rep; Psychotria may 

j Band, ex DC., 


iPsychotriaphaein > a-, I it •. I m , • I il ii-lun ir uni| 

irap;Tabernaemontana aura 




Scientific Binomial 

Ageiiitimuc)i>y70ides L. 
Aglaia agglomerate) Merr. 
Aglaia agglomerata Merr. 
Aglaia cuspidata C. DC. 
Aglaia lepidopetala Harm 

Aglaia sapindin 

' , i n ■ ■ i 1 


P /podiaceai 

I ill ii ■ . 


I uilipirir.piniik-hi i 

: -MIC 
• mi 

( ri>l)(li<>nniiil-::, a 
( cphaloinanesci 
: rihcinfloiibunc 


vigata Blun 

. : rta B un 

bago bagol 

1 ,' 'J - ihl,Ml 1 1 'I 



hapetalui ■ siliflannn 1 mill. 

: ha| el ila< i ae 

/V;is,wr;s ;;<;/. ■:».'•' w Vali.Mon ex Bukh. 

Ebr nai ear 

;-.o.m,' " pa;) -■")• A'alrluiu \ Kakh 


; .,'),' ::.j (.'!-".,,/„...■ (Boivi i Chi 


Panax cannaefoi mis (r or st t) K Si hum 

iuir,, 1 , i.i ','.',''■ 

Dracaena angustifolia Roxb. 

[)yso;<yhi!>)hir.-:v : : Mcir.& IVny 

.'Wild. !'<>.' 

■. •..-.■." ■'I'wianiiin I.M.KiHry 

Dysaxyhin) -.parPlaium ibi m Icy 

//Vjfos/(v,')«.')(;yoyL//ntwiH'Wdrbuk),oi ..ill. 

suiTibi. >/..-,n: ; i!) 

/ iatosicnia novoquineense Warburg, or aff. 

rub. i r.:l\i 

Eiatostema sp.,d c f.macrophyllum Brongn. 

!iat!osia(hyu)hiiquinervis Racllk. 


wui':::.i rci,:!ii 

Lndospermum moluccanum (Teijsm.& Bine 

i.) Kurz Euphorbiaceae 



„nrlY 1 



Gin: hi: !i,,!!; 

Gin: hi:ii.->:: ., 


irum K.Schun. 

las x- h offer 


; ; ,;;|- ;';;;; 

;. ■•/ha Schelly 
^^ ;i 5chettcr 

vogerdak Ho; 

lomal-nqanam Hamuli* < •,',,, \ nil- 

,1 , ,' |, ;ir 

efide Fabaceae 

• : ,' ''. ( ' ( ,: t I I It I I < III 

' j-i'/i, , i ( j u ienw, il'oit lit. nh 

. ;; hlSjlis 1,7. fo/tJfti (Ro-th.l R. lit. 

w/ci i f.i 0/ ; < ' ftc I ■ ■■■■■ i i a 

C.-no :.!■;:;: 

in ,1 l I il| 
■ iinii m::.|i!I|' 

i.J „ ,,,„ 
it,. I ,n ,i >',l 


AliMul'H ''i,= 

Piper decumanum (Rump 

Piper mestonii F.M. Bailey 


Piper pullibaccum Treleas 

Pipturus argenteus (Forst. 

i :..- :,j 

Pisonia longirostrisTeljsm 


<k)<Ui Uimr-i 

. / 11 ///>/- / l h 

■ •. ^it)l()tiispubiw 




^,'c ,-• e/'5U 

: HiMC 



p ■■' . ' "umphii Schott 

:v.t la i' '1 liau >l 

Pioncphtiun} micropinihitum ■ Dl; 


kola v nganan 

uiuWIIU ,p., ill ?W7SfiV(, 1 ■ 

voce) g;;- cat: 

' " , < ,'1 '" ; !ll II 

' ('■ .l, - '< 'opocta Laut.&K - 

. -.iioiiindipicropoda Laut.&K." 

ng ibu n : ibu 

'V" IV,"l.i \' , Ml , II 

h ih.l.. !:!:-. .M 

■ . hol> <i"hl\u i i lah u In 

■■. holiitimiiyuihi lakouc hi 



ngayom nasag 
babaga un 

• , ihotiij, t ?n<-vmv L . 

!'■ :■ aceac 
■ ; ai eae 

masubrepandum (K.Schum 


O/OPidi >\» 

:.els i I 

Sc'dGirr, ti i'. 

sagag .1 1 ir 

bi igang dap 

Termmalia impediens Coode 


• nitiiVki n!i<\ 1 



rropidia disticha Schltr. 

zegiacauliflorai' • !er- 


■eg/'o auliflora iK.'h hm 


si hell.; leaves edible '' '^''-'ru'T^^-f ^''''ihM " '' 

a vegetable,fedesp< 

i as a vegetable . . ' ' 


? Ha ;!• tin ie id It 

is/s (Gepp) Holttum 

ds eaten as a vegetable 

sumatopteris sp„ aff. 

lismatoqiottis sp.;youi 

ig leaves ate edible 

< Hluine;. hewed ,i 


!r!:i;ginCUm I .Mill I I I t I 

used for house rafters 

house a 
digging implements Psychotrio n 

i tt I it i ' | i ts used as 

to, house building {Dioscon 

I In haiiflaluiri wssililhuiitn I < vnh,; used as ties Psycholna s\ 

! ' < , icnnopformis I > t I I S< hi it n n n ■■-. 

used as rope for house building lieved to 

is used 


■hihu)nh)rt)hadrynarioides<-r,yr,ki'i\Roo',: earns 'it/ijir- shoots av d'po.M i-to rhe sap 

i '' ' ' , > 1 1 n f < ' - if r t r i I -:, I t [ t juice extract used 

used for decorative pup i i . Iiti i n i i i iikj cfJVGF 

bilas 10278 

1 i mi f i/MJ.,sm<' hoi mi, lU t ' i i,u ,, nibbed on yam before 

dren to prevent cryin. i i in r i u i J ed in magic rituals 

used m hunting n 


(Elmer) Corner; matu 


Heliciaaffinis Sleumer;fru 

'• , h , n iihu 'iin "" up < 1 nit, tiuits eaten h\ 

Neuburgia rumphiana Le 

[ et mature fruits 

/u,"(, • . - -' fi r v,il i mi ill Iruil i citcn In 

Pangiumedule Rein w.; see 

Pipturus argenteus (Fors 

1 hum fruits eaten by 

C'Mthocuiy. (Kipuamh Piols, or aft.; ripe fruits 

- /o w ,," , , . , , i , M , . r, , h imii 

by possums 

lyzygium aeorai 

-. ■■.■-■ • .'■';' V"- !v',.Mi.;< Pen 


)iiiu| l< no im l',i, t) r> ' , I n it pi In I. h i i fi il I 
ji i ih. j . n in , I it* m ir. i mi in -h' h mi 

jq's nostrils to ( lypiotinxy, h^vinai.i hlume;fed to doos 


:;/o Blume,7/'rora//5 < omplex' i ilypti calyx hollrung Bee 

I r 7 vuh'", L-i i it IjI ill 
be a particularly good firewood for cookin 

>ixbchrymajobi L; fruits used to make: neck'aiv 
■Tmnst'o cfm/ncVaicton; leaves used asvvrai 

j. fresh meat caught in the 
3 (Poir.) Chew; leaves used 

used for wrapping sago 

young leaves are mashec 

diving goggles to prevent 

f'kin(l\oninpop\iono Knuth; b 

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

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

The Latin diagnoses were corrected or written by John Pipoly, and the illustrations 
of new taxa were prepared by N.H.S. Howcroft. John Pipoly also responded to my re- 
quests for information from the BRIT botanical library. Several identifications of difficult 
collections were successfully made by senior dendrologist Kipiro Damas (LAE). My ecol- 
ogy associate Hitofumi Abe of the Japan International Cooperation Agency, kindly pro- 
vided the translation to Japanes [i| i I i i in i i h i i ting with ongoing 
workonthespecimensincludePI I i | i h n> N.H.S. Howcroft (Orchidaceae), 

K.L. Huynh (Freycinetia), and P.F.Stevens (Clusiaceae and passim). Referees P.F Stevens 

1 980. The Euphorbiaceae of New Gu 

ii 11 i r Sin i C I if 1 J- J in i 

sheet 7889 (reprinted bv the National Mapping Bureau 1985). 
1974a F'apu i i f mm 1 I I n in i K -p miu| Mi um y Map. Annanberg 

sheet 7888 (reprinted bv lb- National Mapping Bureau 1981). 
1974b. Papua New hum ll ' M i imi I n Map Adelbert sheet 

1974c. Papua NewGuinea 1:100,000 Topographic Survey Map.Manam sheet 

m - im f unit I I Um 1 1 it u i Oif | in i Mia hi I '-Mi 
aigooy, M.M.J, van. 197 id 'Lint gmxjiaphy ol tin- Pan ilu . Blumea suppl.4:1 -222. 

P. Ho mi ind I vVi i l < ' I t i ii I tin Pacific — floristic 

and historical distni utnii itu i in f I m In I j t a I Miller, eds. The origin 
and evolution o "id 

processes. SPBAcacioniK Pi ess, Amsterdam, the Netherlands. Pp 191-214. 

BmP l 983. Soils of Papna 



and Australian N, 

ational University Press, 


C a i, B J 1 Ml Documentation ol 


3 0fN 

ew Guinea. In: C 

-1 Peng and CH.Chou, 

eds. Biodiversity and terre 




liliil ' 1 it hi A Mni ) m .1 M 1 ,' 'U i 

graph 14423-156. 

Coo i and, T.B. 1949. Aspleni 

and Ble 

:eae of New Gu 

inea. Philip. Journ. So. 


Croft, J.R. 1981. Hernandiao 



. 1 inndboom o* i 

he flora of Papua New 

Guinea 2490-201. 

Darwin, S.P. 1979. A synopsii 


ie indige 


genera of Pacifi. 

c Rubiaceae. Allertonia 

DiPARiwiNt Oi D if- i A I i > " 'i I ii t i I iii a\ ot Papua and Ne 

Guinea. G.W.Reid, Actmo uov;. Piintoi. Poil Mm" ",1 iv, Papna Now Guinea. 
Duyfjes.BEE 199f Hernandiao u Dora Malesiana ser. 1, 1 2(2)737-761. 
E in, F.B.I 977. The pah Hon >i N Gun , Minn n mil i Papua New Guinc 

Office of Forests, Bot. Bull. 9:1-39. 
Foreman, D.B. 1976. A l i> m iin tu i\ -MM i-nu <• M mn Mmloa< memn M 

(mum i uM \u ii ill i Mi I i i in Ii tut uti m and Mm f 1 tii i I i 

vr'i''. ; n, hi Mov : "noiand, A' midaie. Now nouih Walno 
.1995. Prot i Ml > mn.ed.loindl) M < tin 'I u of Papua New Guinc 

3:221 270. 
F i ii 1 I AMn i i u mm i Mi f Mli i n oi 1 10(2) 1 5/ 251 

|| l | III'',', ' I! , ' , IM < I- Ill U 

Gideon, O.G. 19 i 

Hammermaster, E.T. and J.C. Saum i I 11 ih tt and vegetation mapping of 

Papua New Guinea.PNGRIS Publ. 4, Canberra, CSIRO and AIDAB. 
, and 1 995b. Forest resources and vegetation mapping of Papua 

New Guinea. 1 :250,000 vegetation map overlays separately issued as working copies 

to PNGRISPubl.4, Canberra, CSIRO and AIDAB. 
Harms, H. 1 925. Die Cucurbitaceen Papuasiens. Bot. Jahrb. Syst. 60:1 50-1 61 . 
Hartley, T.G. 1966. A revision of tin r i i i n I ' ^ Rutaceae). J.Arnold 

Arbor. 47:1 71 -221. 
Hay, A. 1 999. Revision of Homon •■ , > n i I nil i i u i^ in New Guinea, the 

Bismarck Archipelago and Solomon Islands. Blumea 44:41-71. 
and R. Wise. 1991. The genus Alocasia (Araceae) in Australasia. Blumea 35: 

Hlndrian and D.J. Middleton. 1999 Ri i m J . ' luxynaceae) in Malesia. Blumea 

Heusden, E.C.H. van. 1 992. Flowers of Annonaceae: morphology, classification, and evolu- 
tion. Blumea suppl. 7:1-218. 
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Office, Singapore. 
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75/12. Geological it - Hmnl ■ Guinea. Dept. of Lands,Surveys,and Mines. Pp 

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etation. PNG Office of Forests, reprinted 1 984. 


Malesia, proceedings of the Flora Malesiana s 

C.G.G.J.van Steenis.Kluwer Academic Publishers. Pp 121-132. 

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lian National University Press, Canberra. 

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,and 1 943. Plantae Papuanae Archboldianae, XIII. J. Arnold Arbor. 

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•sand 1:1,(KH), 

vegetation map. 

eel. 1°' hl<- .mil' - M o i Mn I'litilimb 

ational Umve 

Press, Canberra. 

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. iMeliau\m).l 

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8. Useful plani 

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■al Bot. 

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- F II ii - I i Ts 55-144. 

Peter D. Stii ing. 1 999.Ecology:Theories and applications, 3rd ed. (ISBN 0-1 3-91 5653-4 

hbk.) Prenti<< Mill i f | m i Ml- I ill- lu^iri ' s A 638 pp.,b/w anc 
i oloi.ann line drawings. 

/ I f I 1 1 i k\ t 

and tin ones iti tin ti I I t J i nl i i n'l H i liu t n eat h of the subsec 

111 h i, I i .1 ,h I |ii t I I I i ii it ih ii ill u i 1 I i II | i .hi 

qenoial, (lien move:, through evolutional 
a nice presentation of ahioti. Ia< ti 




s It! urn 

ha discussion of the 


i . e. i i|i ii i, I el 

aviora ecology, pop 


ecology.and ecosys 


. Ii .Hill 

reservation biology. 

i them. 1 hose 

ugniiK.inl ioIc-M 



Paul M.Peterson 

Nancy Refulio Rodriguez and OscarTovar 

"[ , [ I II i ■ ;. , h llT' ' n n1- - 1 ll h 1 *h 

i Inl i I u i i , i >■ i, ) .'0 k:n r of Riquid near Hm.n i 1 near 
eCordillera.Thein | i m i i I > I ill i tl 

- M' i .-,-o ■ mm long .' 4 s ,, 


remo surde Cordillera Him i ,[ i ■ n i 1 n ,. i n 

, el "1,1 Kill 

ibas,al norte de la Cordillera Blanca.Esta mieva es 

eon tmmom 


:ulas mas cortasl 10-201 c m d< lai i >] . nm 1 

li lj[ 

inna .1 , 1 l 

ninutas y blanquecinas; lemas anchamente ovadj 

voize (1998), and Laegaard and Peterson (2000), the first and second authors recog- 
?d the unique features of three grass collections.These specimens were distinguished 
:heir dark green and plumbeous-spotted spikelets with a ciliate, flattened rachilla. 

new species is clearly a member of subfan ill I il il Mibe Eragrostideae, 

EragrostisancashensisP.M r i i Fig. 1 PERU.Dfpartami:nio 

A Puinudf' ii (nit I i |[i ir ill 'i I r I iquia on Route 02- 

01 4 on road toward lltnr i (10 u I ' k " \ • mmrn kvht I W P.M. Peterson 

&N.Refulio Rodriguez k? 793 fun crvmUSM!; ^.- ■■■-■. :K\ MO! NY! RSAITAFS! UC! US! WIS!). 

Ab Eragrostis magna Hit< in in tit uj null Mm - lui ilti kininis (6-)10-22(-26) cm 

3 6.1 min lonqis I- m it it i ill n| it ti i in itihtts usque ad 1.5 mm 

lonqis.venisqlumaruni I (. m i ■ tun | ulminiui pluuinqiir ukn< lulls khMis ok'Vdtis, ant hioris 1.2 2.0 

Caespitose perennials. (. tilms ,k> HA < in tall, emc i,teieie neat base, glabrous below the 
nodes; nodes mostly basal or 1 taml . ' ih i uMi i i 1. I i! tons Sheaths 6-16 cm 
long, longer that the In - t intoumde ii qt mt mostly o ii i><n ot with scattered hairs 
near the summit, the hatts up to 1.3 mm lono; maiuit'.s mostly smooth usually with a 

Ligules 0.4-0.7 mm lots] i hn« . A han si i In h tl - - huit . Omuling up to 2.5 mm 
long,thesebreaktiKiLl1 it i itmt I it n in km An} cm long, 1-2.5(-3.0) mm wide, 
flat above the ligule to tiohilv involute above, anii allvai timinaiu, usually densely pilose- 
above, the hairs up to 2..S mm long. Panic les It) 20( VA) cm long, 5 15 cm wide, open, 
the loosely flowi i the culm axis; inflorescence 

branches mostly 1 ^ 1 i i tn f u i t ml t I m i I i < m | i i Ii i i ,i i ondary branches, 

hairs up to 5 mm I nuous to flexuous, 

scaberulous. Spikelets 3-6.1 mm long, 2 4.5 mm wide, florets 3-8, ovate, compressed, 
dark green with small i imb. i | t 1 I II i tbm i u nil iensely ciliate along 
the margins, the halt ti| h I mm I n tit il it n tkil |iiimisftt I then the 

I- tiiin i 1 ill no i In i it i tl i ik i ti i i t 1 In i itiies 2-2.8 mm long, 

lane colali to ovate, muni nai u h lot than link . i F i m i hh mt ogual in length 
1 -veined, keeled cM t il il t t tl i u nk ninute, whitish, raised 

glands;apcx acute to nu uminuto, often muc.tnnate, Um mucro up to 0.5 mm long. Lem- 

scure, keeled, scabi i ul i n,k b I It- i m | ti « n o usually with minute, 
whitish, raised gland i| i it k t kit Minn I I k if i 18 ) I mm long,ellip- 

purplish.Caryopst sk ikitilnt tiuiuh n m n . k. . : I i oovedon theadaxial 

d ligule. C. Inflorescence. D. Spike- 

V. Flowennq in mid to Lite Marc h lliiough May, with caryopses in June 
{R.Ferreyra 14577). 

Distribution —En, e ' • > ku i nl ! n I | utamento Ancash in 

three distinct locations: m mi tne southern end ol die Cordilleia Planca (type locality in 
Provincia Recuay),neai I knyLis [LToviih la I luavinbeind near Bambas (Provincia Corongo), 
just north of the LmdilL i i 'n 1 i nl t n I in lie tun mi stt < | 

rocky slopes at mid elevations (2200-3220 m) associated with xerophytic plants such as: 
Aij,i\A', oeeimekm/b enemy /eye, lei hi, \ hiuico, ami nil km sluubby Astern car. 
Additional specimens examined: PERU. Departamento Ancash: Provincia Corongo, 7 km NW of 
YupanonroadtowdidsB.n hi i Me < f b < ' . ' ' 'iquez 13915 (d5,dSM) ; 

7 km NW of Bambas, 2710 m, 26 Mar 1997 .Peterson & Refulio Rodriguez 13919 (US, USM); Provincia 
Huaylas, between Cara in Muallan i DO )0 m lun 62 J i4,7 7 (USM): between 

Huaylas and el Callejon, 2-100 ,]S00 m,3 lun b)62,/?.bssnym 14594 (USM). 


{Peterson & Refulio RikI" i ' °' ) >n l 1 1 if oi n h k IIihiIok in illustration is not 
presented, since the i hi m> ii h, mi ti i i ni| .m t i I ui i lie sheath cells were 
mostly collapsed. 

The blades are typically loan/ t -1, 1 'c k like Iphospl ioei mipvmvntecarboxykinase or 
classical PCK type ck tin d i utibinl ml Ii ml n I i vnthetic carbon re- 

duction (PCR) cell ell i |ht it i i i i | i ,ence of cells between the met- 

I latterslevandWal'ot pm,i ndpn M i- >l P I < . II ill cb n . d lamella, in Hattersley 
and Watsons (1992 seim nee die chkvi n hvm i i| pears loosely arranged and guite 
regularly is contigu >u n t Mm m t lb i e, t ,, ^ r, ,| t / cells) between adja- 
cent vascular bundles. The lamina are involute with | nimaiy, sec ondary,and tertiary vas- 
cular bundles decreasiini in si/e. I he piimaiy vsm ulai bundles me well differentiated 
intoxylem with mebi ' in | I il ■ i i n i I I u k I it I i n tene md pan u 

chyma bundle sheath). However, tl f in I nil in I itt t the primary vascular 
bundles is interrupted on dieabaxial and sometimes the adaxial surface by a girder of 
fibers. In secondary vasi ulai bundles milv the abaxial parent hymn bundle sheath is in- 
tern pted bv a in I t tilt nti in m it bin | n i k i i i bundle sheath 
is contiguous. The t^ i tl itt. i ( tl mul I iL i m pibi i and the furrows are 
1/5 to 1/2 as deep as the thickness of the blade adaxiallv and usually less than 1/5 as 

a single primaiv e it ndle.Pei blade thei i I mm i cular bundles and 
20-28 secondary and I bin i ul n bun Ik dm 1 i U iiee secondary or ter- 

tiary vascular bundles placed between each primaiv vasniLii bundle. The xylem of the 


£ ancashensis 

E. magna 

Culm heights 



Blades, lengths 

"hi. /idth 


1-2.5i 0)mm 

3 7 mm 

Inflorescence lengths 


1.5-11 cm, spreading 

30-50 cm 
12-20 cm, 


Spikelet lengths 
Spikclet vvidihs 

2-4.5 mm 

2- 2.3 mm 

Rachilla vestiture 


strongly flattened 


eoflemma' 2 mm 

:>mrna lengths 
nther lengths 

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


since the majority of these species are PCK-like and perennial (Van den Borre & Watson 
1 994). We suspect that the closest sister to the new species is E. magna (Hitchcock 1 927) 
since it shares many features, e.g., dark green and plumbeous-spotted spikelets with a 
ciliate rachilla (reduced to a tuft of hairs below each floret in £ magna), very few culm 
nodes (one or two), and rectangular-prismatic caryopses. Eragrostis ancashensis differs 
from E.magna by 15 characteristics (seeTable 1),most notably: shorter culms (26-84 cm 
tall); shorter and narrower leaf blades [(6-) 10-22 (-26) cm long x 1-2.5(-3.0) mm wide]; 
shorter panicles [1 0-20(-38) cm long];shorter branches (1 .5-1 1 cm long) that are widely 
spreading;ovate spikelets (3-6.1 mm long x 2-4.5 mm wide) with a long ciliate (the hairs 

up to 1.5 mm lot 

1.2-2.0 mm long. 

Anothet possible sistei to the new species miqht be F. macrothyrsa Hack., men- 
tioned by Hitchon I- I i h in th i - i Ml t ' oue rothyrsa differs 
from E.ancashen A I ntill lit I i t ill n till (20-45 cm long x 
4-15 mm wide) lone p ml 1> nl n I h i pi 1 ' mm long), shorter 
lemmas (1.5-2 mm long), and shoilei anthem (O.n I mm long). I he distribution of E. 
p occurring in Paraguay (type), Brazil, and Bolivia. 

specimens from 7 km NW of Bambas (Peterson & Refulio Rodriguez 13919) and between 
Carazand Huallanea i/t ',.'ve ' A in t lllm mot- ml hi t individuals with less pubes- 
uiHt'ihin heitt ti I lei on the blades, sheath 
slightly smaller florets, and shorter hairs present on the rac 
from Provincia Huaylas (Ferreya 14577 & 14594) have less h 


. , I .. m 1 l n ' I II 1 < "i I III il II .1 lit- H [ I I 1 1 111 II jl ill 

-u NADP malic enzyme species and PCK or NAP malic enzyme species, Aus 
J. Bot. 24:297-308. 

evolution and don i ti iii n - I lipm mil m i I h i t it I m I u i i< 

Hitchcock, A.S. 1927. The grasses of Lcuadoi, Peru, and Bolivia. Contr. U.S. Natl. Herb. 24 

LAEGAARD,5.and P.M. Peterson. 2000.21 4 (2). Gramineae (Part 2-Chloridoideae) in G.Harling 

A I \nd( i oi ed \ELFcuadot in pn 
Pftfrson, P.M., R.W.WrBsriR, and J. VAinm-Rmv, 1995. Subttibal classification of the New 

World Eragrostideae (Poaceae:Chloridoideae).Sida 16:529-544. 
, , I 1 '", tun n IN til Mio,tideae (Poaceae: 

Chlondoideae). Smithsonian Contr. Bot. 87:1-50. 
R i S.A. 1 "■ i n in i de hi hove 1 I I hi da miens, I 

Tovar, 0. 1 993. Las Graminees (Poaceae) del Peru. Ruizia 1 3:1-480. 
V n i t B 1 1 1 A and I H 1 < M I it i n u h it nut ' il o i mi. 


Sherwin Carlquist 

1 1 i I j i . )" T\v p' 1 ii. n mil 1 Mi t 

atmes is uncle 

garded as specialized.The absence of borders o 

■ t " 1 i, i'i. .M i 

t .ii il in bT im Pli r j| in h 

-r. Quantit.jrivt 

n i Hi r <» ; hasthe A/ood ofa 

desert s'mu:u 

e Phytolaccaceae. E 

Earlier systems include Achatoc ,u | ii n . -i ill 1 >n > ulnn Phytolaccaceae sensu 
lato. The family was segregated by Heimeri (1934) so as to include one species of 
Phaulothamnus and nine of Achatocarpus, and has been thus recognized by most sub- 
sequent authors (Cronquist &Thorne 1 994). The ovary of Achatocarpaceae has two stig- 

occur in Phytolaccaceae s.l. if Achatocarpaceae are segregated. More significantly, 
Achatocarpaceae are not known to produce either betalains or anthocyanins (Clement 
et al. 1994), whereas all other Phytobn aceat I oniain betalains. All authors place 
Achatocarpaceae within CaryophylLilm,, but the position within the order is less clear. 
Placement of Achatocarpaceaejust outside of ul I t Phytolaccineae is currently com- 
monly accepted (Manhart & Rettig 1994; Thome in Cronquist & Thome 1994;Behnke 
1997), whereas Brown and Varadarajan (1985) place Achatocarpaceae outside 
Phytolaccaceae sensu stricto but inside Phytolaccaceae s.l. 

The moderate b^ i t n it iiut m placement of Achatocarpac^i i u I n 

kind of data, include \< it [in In ' I n ' m ilu il I ' n m dilution of tht 

phylogenetic reiatu i I i| tit m it I i n it n d t >T 4 Achatocarpaceae 
have been contributed by Metcalfe and Chalk (1 950) and Gibson (1 994). Molecular data 
are likely to providt u n u - f n > <l > < m >| h II A in i . t relatively sparsely 
singled will: respect to M\A 'oatuies. 

The stem of species of Achatocarpaceae has a single cambium whereas several 
genera of Phytolacca^ eae have sm cessivc t ambia.Onlv a single t ambium is known in 
thephytolaccaceou • i m > , ■ m I T nchostigma {these 

genera would fall into Rivimmone if Phytolaccaceae reduced to Anisomeria,Ercilla, 
and Phytolacca). J\v guestion of whethei successive enmbia oi a single cambium are 
plesiomorphic or uf |l n I I u i n n I Hales as a whole re- 

mains to be resolved. 

The concept of A myophyiMlos offeied hy Aiongmst and I home (1994) or Behnke 
and Mabry (1994) is used hew. Plumhaginaceac and Polygonaceae are considered 
outgroups when tl if i I i it I It i 1 wg However, the data of Will- 

iams et al. (1994) have led a working group (APG 1998) to recognize an expanded 
Caryophyllales in v hi It I hml i nn i I I i i i i« f mi hi ^eae, Droseraceae, 

Nepenthaceae and all I Thome (1994) con- 

cept of Caryophvllal ihail ii i i r n II il In I tern in it 'in of 

Caryophyllales, when .ill families Imve been surveyed, niav refle< I the new classification 
oi ma\ tend to sh ubg uif | ilt< u 

[Ik ft s. in | if u m [ >a 1 1 )i i ut t t I , it i if it gjhvllales that has 

included CaryophylLu one- (C oe h p not 19MS), I'oiiulac a« ear and f fe< torellaceae (Carlguist 
1998a), and Basellaceae (Carlguist 1999a). Among the genera oi lamilial segregates of 
Phytolaccaceae s I tudn I io Mt< n t r 1 - , m.f t M |uist 1998b), Agdestis 

(Carlguist 1999b), Sh\ ge/mne nlguist n imsi 1999d),and rivinoid 

and phytolaccoid Ph ml i i . iln it | II Ml Mmilies of Caryophyllales 

in the broad sense (APG 1998) will ultimately be included. 

[he two genera of Achatocarpaceae considered here cite shrubs to small trees 
(Heimerl 1934) Tim \ n u.n ' m i nh in . i oi m t m nkedly so in the habi 
tats of Phaulotharui n > i h i i M 1 n in tth n ' lexico, less extreme in the habi- 
tats of Achatocaipi, hi hunt lino' 1 i t M i una ill i 1 909;Heimerl 1934). 
The relationship ofwood anatomy to the in ologyol this lamilv is a focus of the present 

MAI ( ;.;IA; s Ar-JI = Mr bKM)-. 
I he collections studied am as follows: - < i I it i n Mi v 

lot A ' 4 ' l i r - i I, I - i i i i i i i i i i m ■ in 

i |( ntin 1 1 itnpf P nn n inn o tw ',' ' Mm , g - t • , .or mm I- o 

22596, POM (sample '< mm in diameter). AHrt softening with 44. ae]ucous cuhylei 

tive features reported ar 
wood features accords v 
group is a mean based c 

when these differ from t 


ment length, 292 urn. Mi i, H n i i e pit mut mt n f dales nonbordered. 

Perforation plates siim|)|. I mi 1 ill 1 11 1 |p ih on 1 , el interfaces (Fig. 3), 

alternate on other vessel faces fFio 1 I m , 1 ! ' | < 1 1 of vessels minute,about 1.5 urn in 

diameter, circular in pu'hti- 1 i^m 1 1 1 ill II 1 1 pit iomtures. Imperforate 

tracheary elements dt^ hit until 1 1 1 ill 1 1 ^\\ iTiq 4 extreme left). 

Length of libriform fibers, 677 um ' m 1 In t 1 I III tin tin fibers, 2.5 urn. Axial 
parenchyma vasicentnc scanty, in strands of font colls (I iq. -!, to left and right of vessel). 

multiseriate rays, 365 um. Mean It! m 1 1 n ^r ells Mean height of 

uniseriate rays, 1 19 pti M ih m it ( m t -It nil ells (Fig. 3, left; Fig.4) 

except for tip cells (Fig. 4, lower left), which are square or upright. Uniseriate rays com- 
posed of procumbent orupright end I i II ill I mm I ill mostly simple pits.Ray 
cell walls about 2.2 pm thicl 1 not observed. 

Achatocarpus nigricans. Mean number of vessels per group, 1 .85. Mean vessel diarm- 

Mean vessel wall thitkiu • , ' pm m- m i I pit Inmeter, 1.7 pm. Mean libriform fiber 

length, 670 pm. Mean I I mi ititil I In i 1 ' pm Mean multiseriate ray height, 

330 pm. Mean width of tnulri Htir i I h 'II dl t-,u kness about 1.1 um. 

Many vessels filled with amorphous yellow deposits. Libriform fibers commonly filled 
with amorphous yellow or dark deposits. 

' i'ii Fi I | i I tmti iiii it t dial groups or soli- 
number of vessels pi^i run r ^ i i 1 t t length, 354 pm. Mean vessel wall 

Mean libriform fiber wall thickness, 1.1 ptn. Axial parenchyma is in strands of two cells. 
Uniseriate rays more, mini n tin 1 1 mlim u 1 1 I 1 1 o) Mean height of multiseriate 
rays,302 pm. Mean width of multOM it 1 o p 1 , ', m 1. iht ^f uniseriate rays, 229 


i, W^'ir 

i f i t i t 

conspicuous growth rmr h.u -m i " 'i IsMiuiu iv^nrpbis Tiate or umseridtc I iq 3 Radial section;anaxi 
chyma strand touches the left side and anotherthe right sideof the vessel.Fig.4.Tangentialsection;tip of bise 
left; alternate minute pits on vessel. Figs. 1-2, scale below Fig. 1 (divisions = 10 pm); Figs. 3-4, scale bel 

Mm. Upright ray cells common, but procumbent ray cells also present (Fig. 7). Bordered 
pits common on tangentially oriented ray cell walls (Fig. 7). Ray cell wall thickness about 
1.1 urn. Amorphous deposits pi< uitmiu I n I in hi t m n i hi 


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

The bark of A. pre proecox, however, both an 

outer and an inner cylinder of thick walled cortical sclereids are present. The more com- 
plex bark of A. proecox may be related to large diameter of the sample studied here. 

Of all families once included in Phytolaccaceae, the family most universally segregated 
is Achatocarpaceae. In fact, the molecular results of Manhart and Rettig (1 994) and the 
cladisticand phenetic studies of Pociman (1 994) showed that Achatocarpaceae are not, 
in most analyses, a sister group of Phytolacca! can, and might even be in a near-basal 
position in Caryophyllales (as defined by Behnke & Mabry 1994). Achatocarpaceae are 

true of Barbeuia. In two caryophyllaleai fun i > ohyllaccai and Molluginaceae, 
anthocyanins are present but betalains are absent. 

Caryophyllales:absence of borders on perforation plates characteii/es Achatocarpaceae 
and also most families ' it |l 'I - it - inn r|,u Pi /Iguist 1998b, 1999b, 

caryophyllalean geneia sf) iImm i I | m. i i m I() ,, loi the familial com- 

position of Caryophyllales or tin | in n I I n n| uiai nl in llie Caryophyllales. 
The libriform fibeis i t " I n ,- i i p oho M a ute in dicotyledons 

according to tradition il , m n ,, r t I'll n as tracheids, shown 

by Metcalfe and Chalk (I. c.) to be more primitive, occur in families often claimed to oc- 
cupy near-basal positions in tin tdn u . >ph Cceae (Carlguist 1995), Stegno- 
spermataceae (Carlguist 1999c), and Rarbeuiacoeie (Carlguist 1 999d). All of these fami- 
lies, however, have successive can h it | I II - v only in some genera), whereas 
Achatocarpaceae does not. Whether absence of betalains and absence of successive 
cambia are plesiomorphic or apomorphic is uncertain. 

The presence of vei m ill | it i i I Id p „ i. s a featui n t re- 

i Caryophyllales (Metcalfe & Chalk 1950; Gibson 1994).The minute 
ve of the distinctiveness of Achatocarpaceae as a family. 
f Achatocarpus can be chara* u u I \^~> \ i- i us Type MB, transi- 

g.5.Transection; vessels are mostly grouped. 
mostly of upright cells. Fig. 7. Radial sectio 
attop,uprightcellsat bottom. Fig.8.Transection, phellem at top,secondaryxylem at bottom;outer cortex consists 
)l parenchyma cells, inner cortex is comprised of pale gray fibers. Figs.5,6,8, scale below Fig. 5 (divisions = 10 urn); 

tionaltoHomogeneousTypebhil n r ] , pi I nutiance of upright 

ray cells in the Phaulothamnus specimen studied is related to the small diameter of that 
specimen and is indicative of a ju\Hhili ml- mi<, < nlqtn ,r I ism) and not phyloge- 
netically different from the conditions shown by the comparatively larger Achatocarpus 
specimens, which exhibit a rathe'- more mature pattern. 

of moderate to marked xeromorphy (Carlquist 1984). This is independently evident in 
the Mesomorphy Ratio (vessel diam ' • t i 'I dement length divided by num- 

tive efficiency.The values for this ratio are: A nigricans, 59; A praecox,99; P. spinescens,2\. 

The desert or near-desert habitat >\ > < < ixhn ith the low values for 

that species. Southern California desuM shrubs as a group have a Mesomorphy Ratio of 
20.9 (Carlquist & Hoekman 1985). The higher values for Achatocarpus are close to the 

coastal sage shrubs, 80.7 (Carlquist & Hoekman 1985). 

APG (Ar i n Ph i i j I •< n i , I i 1 m i tor the families of flo\ 

Behnkl,H.-D. 1997.Sarcobataceae-a new tarn t n .jT\ la, j s (axon 46:495-507 
and T.J. Mabry (eds). 1994. Caryoph la i >h it n m ind sytematics. Sprir 

Verlag, Berlin & Heidelberg. 
Brown, G.K., and F.S.Varadarajan 1985 Si idio' in Cawophyllales 1 Re-evaluation of cl< 

fication of Phytola , n t I i 10 49-63. 
f 'i uiSA^Uh e| ,|i Mipih | n l i I I 1 i | arueand relationshi 

imperforate trachea ry elements. Aliso 1 0:505-525. 

1988.Comparativ< * )Hi it i i i I i D erlm & Heidelberg. 

1 995. Wood anatomy of Caryophyllaceae: Ecological, habital, systematic, 

phylogenetic implications. Aliso 14:1-17. 
1 998a. Wood anatomy of Portulac at > u in 1 1 i< m i m I lit eae Ecological, ha b 

and systematic implications. Aliso 16:137-153. 

i cambial variants. Flora 194:1-' 


1999b. Wood anatomy of Agde 

•stis (Caryophyllales): Systematic position and 

1999c. Wood and stem anatorr 
ationships; nature of lateral meri 

iy of Stegnosperma (Caryophyllales): Phyloge- 
stems and successive cambial activity.lAWA J. 

Phytolaccaceae ^ u \ i I il I i t mm itm I u u unbia Mi 

, and DA H hi u I n n >< m 1 'hi woody southern 

California flora. IAWA Bull., n.s., 6:31 9-347. 

±emlni,J.S.,TJ. Mabry, hi W-i i R,and A.S. Di i ,i i i ,. 199-1 In: 1 1 I) Behnke and T.J. Mabry, eds. 
ii , i| lijl il. L hit i nid ti i it | in i. i 1 i F> ill i 9 Heidelberg. Pp. 

:RONQUisi,A.and R.F.Timi l99TNomei atui n I t 1 n mi history. In: H.-D. Behnke 

andTJ.Mabryeds i n |h i i l in i . n i iti | nnnct Verlag, Berlin & 

Heidelberg. Pp.5-25. 
Iibson, A.C. 1 994. Vascular tissues. In: H.-D. Behnke and T.J. Mabry,eds.Caryophyllales. Evo- 
lution and systematics. Springer Veriag, Berlin & Heidelberg. Pp.45-74. 
lLiMiHi,AJ934.Achatoc l irpaccac.lii:A.Lii(ilpr&H.H.iriTis,Di(.Miaturlk.hGnPflanzenfamilien, 

ed. 2, 16c:1 74-1 78 ih i u n i hngelmann, Leipzig. 
,G)MMiTTn >n Nom<nu n i l l| t> 1 Multilingual glossary ol terms used in wood anatomy. 

Verlagsbuchanst ill ' i I i I i in* Muni n i hind 
;ribs,D.A. 1935. Salient lines of structural specialization in the wood rays of dicotyledons. 

Bot.Gaz. 96:547-557. 
/Ianhart, J.R., and J. H. Ri i no. 1 994. Gene sequence data. In: H.4D. Behnke and T.J. Mabry, 

eds. Caryophyllak , I nti i m I i 1I1 f mi i i hint Ihildhtm, 

Pp. 235-246. 
/If rt au i, C.R. and L.Chalk. 1950. Anatomy of tht ; 'flu J m i idon Press, Oxford. 
Iodman, J. E. 1994. Gadi t and | h n u« siuuics. In: hi , ' Behnke and T.J. Mabry, eds. 

Caryophyllales. Evolution and systematics. Spunqer Verhg, Berlin & Heidelberg. Pp. 

Vai m r, H. 1 909. Phytolat caceae. I )as Pflanzcnreich IV(83):1 -1 54. 
Vii i iams, S.E., V.A. Albi v., and M.W. Cm- . 1 994. Relationships of Droseraceae: A cladistic 

analysis ot ,oh / sequence and inorplu imgical data. An am . Bot.M 1:102/ 1937. 


Chapel Hill, NC 27599, U.S.A. 

Richard D. Noyes 

Plant Genome raapping I aP'oiaioiy 

I I i'l f t I /i C lllf'M,) ll| ,, 

Athens, GA 30602, U.S.A. 

i ■ in | i 'v.;.i;-n r-, ,7 1, f 

Batopilasia byei (Sundi et i In i ' nt i >» Batopilasia byei at I 

)rphology. DNA studie 

I i ,■ i| I 
•Pin i iint'u / i i I i i I in se excluye de Erigeron y de 

lae y se trata como genero monotipico Batopilasia Nesom & Noyes, gen. no\ 
Batopilasia byei 

superficialmente en su morfolnq ,i I i n i i m ^ncanos fileticamer 

"I . u II- i> 1 1 ijui / at " i. l t i n . < i ml. < hloracantha. El nue 

ie original desctiptu r I I II om 1990), it was regarded as 

;t similar to E.ortegae Biake (= A i lm i i i id F oxyphyllus Greene, these 

■e species constituting I Pugnron sea. \pmiosi [F.ortegae.the type). A number of essen- 
jifferences have since been tea* u <d i "-t • ' , »m 1 1 ortegae, and the 

?r has been segregated as th>' m 'i | i mi - 1 m.icantha (Nesom et al. 1991; 
Sundberg 1991) Enge>o> ■ , L ' m I > n treated as a member of the primarily Cali- 

1 I l m 1 1 < i n '< < it Pycnophyllum, 

som & Noyes 1999). Nesom et al.(l<) ( )|) noted thai ,iltliou< ih "the evolutionary 
<s of E. byei may yet prove t utside of Erigeron, there is 

in an isolated pos hi : ' -.Mit ml, I tm l < < ul ' i the 1 tribe 

Astereae (Noyes & Rieseber origin of F. byei 

lies outside the phyloqeneti m u ol </pe.m>o md < ., ind their closest relatives. 

Relationship to Erigeron and Conyzinae 

Erigeron byei was originally plat ml in Ammcm bet uuseol its pen mved resemblance to f. 
ortegae and A our' . ' ' itt i tl i in ml ti i i r mi mPticm hip of 

the latter (within Erigeron sect. Pycnophylluni) and the minster of the former to 
Chloracantha.E. byei has no close similantv to any othet species e! / M^tvon and does not 

\ t it ii ciii\ ; n in. i in ul i ' ii ' i II i II tew flowered heads 

i ■ the species from the genus. 

Molecular studies by Noyes and Rieseberg (, I 1 ' « 1IJ1 < '"I'liiidud. n |'te 

Conyza;2Aphanobten 1 i h t n I a and delimit a mono- 

phyletic group that loin | md mill i >n im I II -m 1 |l H), with three excep- 
tions: (1 ) the North h n i in , ' Ii i ul the group, appar- 

ently in a sister relat i I | ■ , ' - I i pi in i i: h its close relatives), 

a species native to lhe< mda m Island thes. l\ ^ nmip in a tei telationship with 
a group of species indu in i ,,-:■• '< ,Aam m Pik an species of Conyza are placed 
within subtribe Grangeinae cither than Pony/mae; and (3) A dye/ is related to species 
outside of the Con\ 'ii ni A > leAeitedqh pi i ind genera of Conyzinae encom- 
pass considerable m< q u i il ' - i it but r s the only New World species to 
be excluded. 

tently green-glabiati i. in mid k i < . m, t i u I d i il i ph hum itlnu her 

ba< i ous ,jpu all\ ■ n t I tu th'hr |i it - nt it i urinous nerves), and 

deltate collecting ipptndii "tl t IN i I Soltonia also shares 

this combination of habital and capitular features. 

In the Noyes and f i 1 mi p ' i I i tl it i 

genu in » > m i, , m thi m tti i tnuiiii si ii I it I to >ymphyotnchum, 
Oreostemma, a ndqm i 'I It A > i tilth in > A? was not included 

iti tin ii litiah I it | I I i i t n n I h n Boltonia and ( 

earlier been suggested by i pi )NA analyses of Astereae -in the context of 
pling, Morgan (Amu | , j p, ,■ , > - , t > *> , is most closely re 
goldenaster group tan ml I , i u hi ih (1989) observed t 

from nuclear ribosomal DNA (Noyes& Rieseberg 1999), but the suggestion remains that 
Boltonia and Chloracantha are closely related. A long-standing hypothesis of close rela- 
tionship between Boltonia and th« ian g iu m ris was rejected by Gu and Hoch 
(1997) on morphological grounds and by Noyes and Rieseberg (1999), based on mo- 

Boltonia remains a genus strongly isolated in morphology (see Cronguist 1 980 and 
Anderson 1987 fortaxonomic suiiitiuii ' Im tit i it. mti Erigeron byei in having mostly 
cauline leaves, conical or convex receptadcs,phyl!aries with a thick,raised,orangish midrib, 
short-tubed disc corollas, cypselas broadly elliptic to elliptic-obovate, strongly flattened 
and broadly winged or unwinge I ill i i eat ich margin, these often with con- 
f i i ii )i in |i I oil Iu in i i t i i It itonir i til il t i'l 

bristles and 2 (-4) much longer, thickened, barbellate awns. 

Chloracantha differs from Erigeron byei primarily in vegetative features— taller stat- 
ure, thick rhizomes, pen iirii.i it lh i i ._[ i ith thorns and axillary budsjack of 

persistent basal leaves — but also in fin pi heads and greater number of flowers. The 
similarities of Erigeron byei and ( h, , - nth i uhierete,4-5-nerved cypselas and 

highly derived (apomorphic) fruiting features of Boltonia. 

In summary, although the diu i i mi i in jm nnnot' jcum byei was sus- 
pected at its original description I > i h u t ■_■ r - ■ 1 1 . >' n i ts r ■_ I iin <>lv generalized fea- 
tures make it difficultto placethe it Brail i iorphology,however,£.bye/'isnota- 

bly similar to the genus Chloraca>~ u n i- I- uln tdence indicates that it is closely 
related to Boltonia and perhaps l' 1 - < > . "uP basts of its morphological isola 
tion within Erigeron and its unamhn i m ml il \ \ In I « |mi< tu [placement outside of 
the Conyz 

Batopilasia Nesom& N(j\l j s ijtti no, I n- m ibeig) Nesom & Noyes 

A Chloracanthae similis ,> nm ti n mm I n npinMin .ilde redactis, gemmis 

| I ii i inn n | i i I ill m e ul mbus. 

Batopilasia byei Nesom &No v < < Fig.l "yeron byei Sund I 

Phytologia 69:278. 1990. 
Perennial, caespitose, glabrous heil ttnni ,tn I ihtn,lignescent,rhizo 
dex branches. Stems 7-20 cm tall, usually with 1 -2 ascending branches n<E 

Basal leaves pi i i.f'in nn n n t nt 

I. Habit and morphological details of Batopilasia byei. 

I III i 11 J 'II 11 III IH I 1 tllll '111 ' 

jed at least on the upper hulfthe innei sen 
k . n - P i II ' m " 1!- I' Hi ' tin 

5-7 mm long, the lamina 1.4 2 mm wide, coiling at the tips. Disc flowers 24-29, fertile, 
the corollas 3.5-4.2 mm long, not inlkited oi indutated, lobes triangular-deltate, erect; 
style branches with I Mi II duo i I m I mm I i Lvpselas sparsely 

strigose, cylindrical and t- 'td to kiotuk Hutt. m d I mm I jiuj, 0.4-0.5 mm wide, 

with (2-)4(-5) thin, orange nerves: pappus of 1 5 Tl barbellatc bristles 2.9-3.8 mm long 

Nesom 1990). 

ently is restricted Th< In u| uhi n I u u.din undberg & Nesom 

the town of Creel and in the general urea of the Bauunca del Cobre (Fig. 2). The plants 
grow on steep rocky slopes, commonly in rock crevk us and ledges in arroyos and can- pine oak wo i ml i ' u mum t i k ti u il unci May through July. 

Batopilasia byei 

bution of Batopilasiabyei. 

ts from Don Pinkava and 

i from MikeWarr 

her of Phytologia) to re 
i! for help with publicat 
iversity of North Carolir 

Qonouim, A. 1980. Boltonia. In: Vascular flora of the souths 

Asteraceae.Urm <M M 'iih mli'i.lh ' Chapel Hill. Pp. 1 
Gu, H. and RC. Hocn. 1 997. Systematics of Kalimeris (Asteraa 

Bot.Gard. 84:762-81 4. 
Morgan,D.R. 1 990. A systematic study of Machaerar 

Machaerantheia^^w i ' Fh ' lr t'tm t ini\ of Texas, Austin. 

NisoM, G L 1989 Inti i i n i n-iu n .fff 1F1 <i<ieron (Compositae: Astereae) 

Phytologia 67:67-93. 
Nesom, G.L. 1992. Revision of Erigeron sect. Linearifolii (Asteraceae: Astereae). Phytologia 

Nlsom, G.L. 1994. Subtribal classih ii u 'il i i i i > Phytologia 76:193- 


ereae).Sida 18:1161-1165. 

, R.D. 2000. Biogeographical and evolutionary insights on Erigeron and allies 

■teraceae) from IP_> se^ii-u i' dahi II S\ i ,ol.22u: l) 114. 

, R.D. and L.H. Rhsfbfrg. 1999. ITS seguence data support a single origin of North 
• in terc v (Asteraceae) and reflect deep geographic divisions inAsters.l.Amer. 

aae. Phytologia 70:382-391. 
x Phytologia 69:278-281. 


(hupcllliil.Ni .' ■'■.W-.VXt)(l\A 

3 species of Baccharis) by F 

Archibaccharis trichotomy 

Archibaccharis ca/oneura S.F.Blake, Proc. Biol. Soc. Washington 55:11 7. 1942.Type: MEXICO. 

Oaxaca: Mt. Zempoaltepetl, lower slopes, 1 9-27 Feb 1 937, W.H. Camp 2701 (i iolotypi : 

NY, fiche!; iso; ypr: US-photo and fragments!). 
Klatt's citation of the type of Ben i ' f i i In i I lebmann Nr.55.Herb. 

Hort. Bot. Hafn." Judging from the handwriting (by comparison with Burdet 1978), the 
type specimen apparently was fit t i 1 r n II huh I it > tin i is' Baccharis elegans 
HBK.? var.The identification as t < '< * I Ltt |v. umably was laterand 

by Klatt himseif.The provenance of the collection is not noted on the sheet or label, but 
other details of the pecimen, includirui th irh i m ii i I itii itu nh mh 
match the description and protologue published by Klatt. The nature of the three-parted 
division alluded to by the epithet is not clear. 

Archibaccharis trichotoma is known to occur from central Oaxaca to Chiapas, Mexico, 
where it grows in pine-oak, pine, evergreen cloud forests, often with oaks, 1750-2900 
meters elevation, flowering in October through April (Jackson 1975; Nesom in prep.). 
Liebmann travelled in Mexico and Cuba during the years 1840-1843 (McVaugh 1987; 
University of Copenhagen Botanical Museum 1999); he collected in 1842 (May through 

early October) at nuim mi \o< tliii in ntm inpion of i i (McVaugh 1987), pre- 
sumilil. hi il it t,p- :- iiii.'i,!'-'' /as gathered. 

Hit li iumiI \/ i i I i , n< m late, olntouith long at uminalt 

apices, petiolate, thk h ji, I il> m > it I ninn t t > n 1 n ition, mucronulate on 
the distal 2/3-1/2 of the margins, the stems are glabrous and evidently dark-colored, 
and the capitulesciii ■- i n i i n t m il i in n in 1 iho\e the level of the 
upper leaves. The type specimen I it i nun it h. il ' ' . Kins trichotoma is 

southern Mexico and i ntia m i tl n I i e tn t m c t stems and coriaceous leaves. 

Ih t i >iii mik probl i i I t n i ii M li iln i it ii il i in I i | ' mm n phot 
graphed by the Field f ] i i t it iltl i m< nil i m the type of Baccharis 
trichotoma Klatt. The label of the specimen (W 33218, MO-photo!) identifies it as B. 
trichotoma Klatt and indicates only that the collection was made in "Mexico"by Galeotti, 
will oliected in Oaxaca, 

including regions in tin 1 1 ' r 1 1 • 1 1 s i > r t r i ■ it- In' <• > ' hotoma is known 

to occur, from April 1 839 to early 1 840 (McVaugh 1 978). The two branches on the W 
sheet were taken from pistillam [Tints that am t haractuiistk of the species as known 
from the type and other collectioi L inmiti n til n I the Galeotti collection 

(not the type) led to a search lot the type, nltimatniy annulling in the reduction of the 

mi n it tul t j i ''I tf R\ tin im ho lo it tin pe, | pared Mined mag 
and sent it to me digitalivg denial! of f)S tot help tinting a vim! thou ,, MO for a loan of photo- 
graphs of types toTEX, John Stroll urn tot em out agement and comments, and John Pruski 
and Eric Lamont lot m nn i 1 e n tit n it nil t if it I f nil Tcations costs pro- 

vided by the Motion it Itlm n I ui I ill M u i n t n lina Herbarium (NCU). 

pi M 19 7 S.(. .inula, i ' b t ' ii ui i n ipl i 
a i m, on, J J ). 107TA mvinion of the pen io on /'Ann < oneo 
Phytologia 32:81-194. 

ting of tfieoiitainal v 

alu iti i >t ti I i ui i i ii ih tonus Phytologia 
i prep). !lie( onijoul ' I. ■ . • .!■ nmtic account of the f 

ereae, lo' Monnono. 


Thomas G.Lammers 

Department of Biol •> „ .' ; i < , 

i'"/. ■ ' •': < i 

1 ^ 'l' ")490/, (7.5.A 


:ion of over 2500 Sf^iniui 11 1 huhi il if- 3. e recognized, all are 

il Chile. Two are branched solid- 

i" I ni litre tit il i i i Hi i I is found in the dwarf- 

arub and sclerophyllous vegetation zones of the xeric north: Lpolyphylla Hook.& 

a Bonpl il I i i l h i li i , - i 

te other two are la 


nperate rain forest regions o 

7 Hook. &Arn. The species o- 

hi SUM I N 

j L.polyt 

Lobelia L is the largest genus ol Lobel ■ leae impanulaceae), comprising over 400 

species of annual and perennial herbs, shrubs, trees, and giant rosette plants (Lammers 
1993a).ltiscosmopolil i ii idi Ml ution indigenous to six continents and several island 
groups (e.g., Hawaii, N iland,th i it illes) Nearly 38% of the species are African and 

another 29% North Am i u i in I Australasia each have about 10% of the species, 
South America H ( ', cit i I hi n ■■ hile only two species occur in Europe (Lammers, 

The genus was lasi t n u i| I II hint r 1 ' h kind it i r it 

:e subgenera: Lobelia with two sections; Mezleria (C. Presl) E. Wimm., nom. illeg. (cf. 

imers 1999) with t html I run n( i sections. Further 

ture was indicate d b li id in j man f the sections ml subsection 

and other subordinate taxa (cf. Lammers 1 993a). 

Wimmer's classification of Lobelia was recently revised by Murata (1995). Though 
based primarily on seed coat morphology (Mut Mi I 1 ' h mo n .is on was also concor- 
dantwithother dato ti n ,l I n mm i i In lin n inomosome numbers (Lammers 

1993a) and chloroplast DNA (cpDNA) restriction-site analyses (Knox et al. 1993). In this 

of the second one should be replaced by subg.lsolobus (A. DC.) Y.S. Lian; Lammers 1 999). 

Among the sectioi t il In i n I I Tut i i t 

(G. Don) Benth.Wimmer (who used bo i illi ntimate name sect. Eutupa E. Wimm.) 
had construed thi s n i i n I i k li i | i ti i i tu< a South America, 
and the West Indies uothin tin - ton h - < <t . nnmer did segregate the Chilean 
species as "§1 . Species chiletises'hiiidoi the invalid siibwctiona name "Primanae" nom. 

nud.) In Murata s le In i ill it i > hi u \n < . in removed to sect. 

Colensoa (Hook f) J Mm h i i < > t ' l i 'm [ ( leaving only a small group of 
species endemic to central Chile in sect. Tupa. 

iteroparous(polvt t ii|n I min i i il t i 1 f)ink,or wine-purple 

non-spurred corolla v.uh nv >iu mu npho dYf •, Y I b.o coherent at apex; staminal col- 
umn shorter than il il anthers bearded 

cells with long lumitM p ' f 1 urHl L <' - md hexaploid (n = 21) chromo- 

some number. The lattei two features appeal to bo unique within s u bg. Tupa (Murata 
1992, 1995; Knox etal. 1993; Lammers 1993a). While hexaploidy is regarded as derived 

(Murata 1992,1995). 

This more restricted in m ti i i| | i > ti i , I , the phylogenetic 

analyses based on cpDNA data (Knox et al. 1 993). In the concensus tree, the species of 
sect. Tupa s.str. formed a monophylet'c group tiiat was supported by 10 cpDNA restric- 
tion-site mutations (bout w| il •' I o ilu hue n lining species of subg. 
Tupa examined, u e <> w t w I ai I < i i Roth ex Schult.) 
that had been includ I in > t ' I, Vimmei, fomn i its toi group. These plants 
were all tetraploid (n = 14), so fat a known (Lan mers 1993a). 


lighly toxic plant known to the indigenous Mapuche as tupa. Feuillee gave the plan 
e Latin polynomial"Rapuntium si>catnm,foliis acutis.vulgoTupa. "In Species plantarurr, 
inaeus (1 753) accepted the species on the sole basis of Feuillee's description and plate 
d assigned it to his genus Lobelia. In taking up the vernacular name as the nomet 
/tale, he misspelled it "trapa."This error was corrected to L tupa in the next editior 
nnaeus 1762). 

scribed from Chile. More than om 

--third of these descriptions (e.g., Sims 1810; Lindley 

1 826, 1 830; D. Don 1 835) were base- 

d -it ints cultivated from seed in various European 

botanic gardens.and were publish 

' I i In r 1' I < mi puf iFii tiic ii i , . ' 

Botanical Magazine. In the latter h< 

alf of the century, eight more species were described, 

primarily by resident botanist Rodi 

i I i 'fnti loPhilippi (1808-1904) in the course of his 

studies of the Chilean flora (e.g., F 

'hili| p 18^5, 1895). A number of varieties and forms 

were also described over the year 

s by various workers. Altogether, 35 heterotypic taxa 

referable to Lobelia sect. Tupa hav 

e been described, all from Chile and nearly all in the 

nineteenth century. 

Differences of opinion on thi 

2 circumscription and typification of genera created 

additional combinations in the grc 

mi' t> D ] 83d) believed that Rapuntium Mill. was the 

nomenclaturally correct name k t tn t n | i i i I m -t lobelia by Linnaeus, 
while Kuntze (1891) argued fot J. "*■ • • ■ 7 Hi ' « t t>- Don (1834) segregated robust 
species of Lobelia > th il T it II 1 in upa G. Don, a move supported 

by Candolle (1 839) but subsequently reversed by Bentham (1 876). These differences of 

in an additional 61 homotypic combinations based on the original 35 taxa, giving a total 
of 96 validly published names referable to Lobelia sect. Tupa. 

In the most recent flora of Chile, Reiche (1 905, 1910) synonym ized or excluded many 

sect. Tupa. In the most recent monograph of the section (as "§1. Species chilenses"), 
Wimmer (1953, 1968) recognized 20taxa:seven species plus thirteen additional hetero- 
typic varieties and forms. This classification was implicitly accepted by Murata (1995) 
when he remodeled the section, and was also embodied in the most recent catalogue 
of the Chilean flora (Marticorena & Quezada 1985). However, when I began to prepare a 
treatment of the Campanulaceae for Flora de Chile (cf. Marticorena & Rodriguez 1 995), it 
became apparent thai In is classification of the section i loss cum optima I, and that a 
thorough revision was required. 

Though tupa (sometimes rendered as trupa) is the Araucanian name for L tupa, the plant 

1965; Hoffmann 199'/). This 

i by SchuluM 1981, 1990) to be Ooliniiolv 
psyc hoactive/OOe ac (uaO cam pounds n ,ponsihl i it I hi', h t iy are not known. The 
latex does contain f nun Tkaloid including 1 in it 1 i liketo- and dihydroxy- 
derivatives, lobelanidine and nor-lobelanidine (Santa Cruz 1 932; Hill 1 970; Raffauf 1 970; 
Gibbs1974);however,these substances are n l|v In n . < MiOtes^ Hoffmann 1980). 
Lobeline has bei i the treatment of 

bronchial asthma s\ mi t m n 1 n ^ m i i i ii lit nl tmmBlacow 1972;Lewis 
& Elvin-Lewis 1977). In Chile, the expressed latex of L. tupa has been used by rural folk to 
relieve the pain of dental < ntim,and in compiosses to met joint and hoof pain in horses 
(Murillo 1889;SanlaCru/ I 932). I ho pharmac oloqL nl hast", lot such uses are not known. 
I lalluLinooenu. ot not,/, tupa is definitely toxic . I ouillee H714) reported that the 
odor of the flowet •! >n i i i i i t in in i ntit i I piM e m i! in tl I i 

not able to confirm th i* Im t l'mntoouohth . it i I. m means pleasant. In any 
event, beekeepers t oi , side i th- | Tint i numm. > a t u.s t n n ^s honey an acrid, un- 

below),l have observed large orange bees stealing ma tar from tne flowers via the dor- 
sat slit of the corolla, without contacting the anthers or stigma. 

Feuillee also repot tad thai 'a nail quantities nl latex nibbed in the eyes would < uuse 
blindness, a statement I can partly confirm. On more than one occasion during field 
work, I absent-mindedly rubbed my eye after handling material of L. tupa. Though latex 
was no longei pern. [ ill it mi I m i t iiu t 1 tan to sting and water 

profusely and m\ i t I hit hi h t it Ii It Mr face became swollen 

and numb.The effect pm ma e tot n- at an ami Munlf M.- 1 a teported that inges- 
tion of the plant ot it Me ul 1 i mm i , nit tun lie it m bloody diarrhea. Most 
recently, Matthews i I 1 ' ' I i inn ■ in d th. < i el i London gardener who was immo- 
bilized for 10-15 minutes after ac: cidenlally inhaling auborne dried latex of the plant. 

On a more pleasant note, the species ot MOO/ set t. lupa have considerable (toil 
cultural potentiaOO i I i i t it r q t O n | O i the nineteenth cen- 

tury, the resulting [Tin ie m 1 i ii I il lint ' im i i laideners and plant 
aficionados (Sims 1825;Lindley 1826,1830, 1833;G. Don 1834;D.Don 1834, 1 835;Lemaire 
1843; Loudon 1844). Though most were soon lost :mm cultivation, L tupa apparently 
remained in the trade (Voss 1894;Finnis )0 tilth. 1 9b8 Thomas 1990; Huxley 1992) 
and may have been the object of some selective breeding; Chittenden (1 923) mentions 
a cultivar fromStob num i\ n inn d Oilli it it II. »< m . «.i 'I the three other species 
recognized here has been reintroduced to botanic gardens (Jammers 1993b), which 

mtTi .V-Yk ■■■ 

I. I hif.ii L h- that i 

slli i up I i t. I , , I 

ami October I I ml .< t I ' >< 

ng the entire geographic, elevati 
(27 2(TS) in the north to Puerto (V 
.representing all speci 

visited and studied. I was also able to examine several naturalized populations of L. tupa in 

the Juan Fernandez Islands during an expedition in January-February 1 986 (cf.Lammers 1 997). 

During these stud particulai itt nlioi | iid I mi t of n rph I gi il 

variation within populations. Then jh ne of th ■ nfraspecific taxa recognized by 

lations,they do not merit taxonomic recognition [ghrm m >\ ,o placed on discern- 
ing possible geographic,elevational,or ecological correlates of morphological variation. 
In addition, material was also gathered fo The results of that 

work were reported separately (Lammers & Hensold 1 992). 

In the end, it was concluded tli.i! onl\ mm t - . i^ mid no ihfraspecifit taxa couid 
be distinguished in sect. Tupa: L.btldqesii Hook.T Am. [including / . hianda (D.Don) End!.], 
Lexcelsa, Lpolyphylla [including L oi/afa Reichc i i I [inTu hngf muaonata Cav.]. 

The rationale for each of these synonymization i I u - m 1 1^ respective species. 

morphology, and habitat: (1) a pair of branched solid stemmed si nubs (rarely small trees) 
with faintly striate oblong or ellipsoid seeds, in the dwarf and <erophytic-shrub and 
sclerophyllous vegetation zones of the xeric northern portion of the range [L excelsa 
and L polyphylla); and (2) a pair of robust herbaceous or suffruticose unbranched hol- 
low-stemmed perennials with minutely foveate-reticulate broadly ellipsoid seeds, in the 
deciduous forest and evergreen temperate rain forest regions of the mesic southern 
portion (L bridgesii and L tupa). 

The two pairs are largely allopatric, though populations of L excelsa and L tupa do 
approach within a few miles of each other in Prov. Colchagua, Curicd, and Talca.The 
members of each pair are definitely sympatric In the xeric north, the geographic ranges 
off. excelsa and L. polyphylla overlap by roughly 90%; the former extends a little farther 

high into the Ande. hi i i il ih> mesic onth '" ;u I i i cety restricted distri- 
bution that lies entin thin th broader range of L tupa. 

Not only are the members of each pair sympatric, they often form mixed popula- 
tions. Ofthe 17 popul i i tudi I in m m si nth - n ipnsed L. tupa only, 1 8% 
L. bridgesii only, and t mi I >i llu m population ml i in the xeric north, 
54% comprised / p< ul >i\phvlla only,and 21% were mixed. 

Despite this appaiont opportunity for miscegenation, I could not locate a single 
morphologically intermediate individual during field work, despite intensive searches. 
Furthermore, I have examined only one herbarium specimen thai clearly is intermediate 
i.The holotypeof Tupa kingu Phil Tected it il| iraiso 
nt L excelsa x L. polyphylla (see below). 

..Differences in chromosome numb< i ma\ \y i il< I ut il 

do not appear to be i i m - In .' ' I < oius to bloom a few weeks earlier 
than /. excelsa, there s stil veiv < onsiduuble owiLip n then respective phenologies;/.. 
'",,,,-, 3 * 1 1 : ' ,--1 7 hi i n o n iimutl, 

I he best hypothesis is that isolation within each pan is a pioduct of pollinator dif- 
ferenu i situ it n u nl n i ll n 1 1 | 1 1 i I i i I n I t ' I 

(Thompson & Lamm- I' 1 hi, itt i li | I I it \ \ -n tint one mem 

ber of each pair is adapted to entomophily, the othei id oi nithophily.The flowers of L 
bridgesii and L po\?> h n mil t It tm i , its by bees and other insects. I 
have not observed | in an i it i I r I mje orange bees were observed 

visiting flowers of 9 bm /mop i onUKtinq I he aniheis and slip mas with their backs. When 
the same bee visittm ' it is , i i t tl r n ii tar via the dorsal slit 

without toiu lunt] in I i i 1 1 , i r i i t ti it I hi t ill i t-xcelsa and L. tupa 
appear to be adapts | In i' n mum ii n i ih u hi i not observed this. 
Further biosystemati i n h I . I needed to test these hypotheses. 

Hhileth lent pi i II m t ii n'h hii ihhi o-d morphology, 

and habitat thisdot i | n I u i it i t m itl i i lehving phylogeny, 
as inferred from cpDNA restriction-site mutations (Knox et al. 1903). Though the south- 
ern pair (/_. btidqesii an i ' i upp it li i b It I tin 1 1 ned mutations (boot- 
strap value 95%, decay index - 3),tnenoitnern paii : s not. i nsteai t, /. polyphylla forms the 

branch, sister to the southern pair. The sister-relationship of L excelsa to the southern 
pair is likewise supported by three mutations (bootstrap value 93%,decay index = 3). 

Though no det nl I i ml ul il I ' | I m I it I i i| | n 

that morpholoqical I it 1 n r nlm it! 9 p[ pi I n Lobelia polyphylla, 
the basal branch, is the most discordant membei of the section, with its much smaller 

red (or pink) fiow. t lit! ithern pa laracteu i unbianched hollow 

herbaceous or suffruti; i e ten usual lecuuent leal maiqit snd minutely foveate- 

reticulate broadly ell p si o I i M qu fait . \rhin the (L excelsa + southern pair) 
clade are the pink corolla of /. bodges;// and the tan oi pale yellow latex, bibracteolate 

pedicels, and floral coloi , I ,im \. I - t t t ments regarding the 

apomorphyor plosiu i ill. t in i i ll i I ndinq better knowl- 
edge oftheoutgroup. 

ib 0.6 9 m t ill mi I m ill n ' i ill i i I u t | i nni il 

woody, solid, an i mf it I i i I i o 9 i hi i uttm 

pale yellow or tan. Leaves simple, alternate, exstipulate, pinnately veined, sessile, pubes- 
cent or glabrous; margin entire or finely callose toothed sum limes forming a sagittate 
base decurrent on the stem below the point where the midrib meets the node. Flowers 
perfect, protandrous,resupinate,epigynous,zygomorphic, pedicellate,solitary in the ax- 
ils of the upper leaves or these reduced in size, creating a terminal 10— 65-flowered 
bracteate anauxotelic (rarely auxotelic) racemose inflorescence; pedicel equalling or 
shorter than its flower, ebracteolate or with a pair of linear bracteoles at or below the 
middle. Hypanthium obconic, hemispheric, campanulate, or obovoid, pubescent or gla- 
brous,adnate to thn u , ,| . ,|,- il ,ate,triangularor narrowly triangular,pubes- 
cent; margin entire or very rarely with a few teeth. Corolla sympetalous, unilabiate, 15-65 
mm long, red (sometimes yellow and orange in I ', I, >t >< i > ill yellow throughout 
anthesis), pink, or wine-purple, lacking a nectar spur, glabrous or minutely pubescent- 
tube suberect, curved, or arcuate, slit dorsally to base; lobes 5, valvate, monomorphic, 

lobes,connate,forming a staminal column shorterthan the corolla and free from it; fila- 
ment tube exserted through dorsal slit in corolla, bearing small flattened triangular tri- 
chomes ventrally at base, otherwise glabrous;anthers dehiscing introrsely and longitu- 
dinally, the dorsal three longer than the ventral two, occluding the orifice of the tube, the 
surface of the tube glabrous or the dorsal three with scattered long hairs, the ventral 
two with tufts of white bristles at apex. Ovary inferior, bilocular; placentae axile, large; 
ovules numerous, small, anatropous. Fruit a half-inferior capsule, broadly ellipsoid, ovoid, 
obovoid, oblate, or subspherical, dehiscent by two valves at the rounded or truncate and 
apiculate apex. Seeds small, light brown or golden brown, oblong or ellipsoid and faintly 
striate, or broadly ellipsoid and minutely foveolate-reticulate; testa comprising a single 
layer of cells with long lumina (Type D of Murata 1992, 1995). Chromosome numbere = 
21 . Four species,endemic to central Chile. 


:orolla (38-)45-65 r 
he lobes 12-33 mrr 

. Hcnnist pcrenni. 

,t< in I -i 2 30 ii 

22 33 mm long 3.L.tupa 

U i i il i m 'I , -mi ill ,n inn ihi k ii tl,Htul'« 1 -1° mm long; 

Lobelia polyphylla Hook. & Am., Bot. Beechey Voy. 33. Dec 1 830. lupa polyphylla (Hook. 

Lobel. 29.1 836 Dom , ' II I -mi hi t i ..nil 1 Ih^llnCHIIK 

1 i | ,[ H ' (\ t fj 1 f'll Ml, I 1 - ' i ' ii i n Ml Infoi 

mation in brackets: \A n mtl,H,n il m mi H P c Arnott 1830). 
Lobelia purpurea Linol hi nl hi I'M I'M 1 I' l'" i 1 1 n hutu Hort. Breiter. 249. 

I8 ! /. I'upapuipuiea C.[ )on., Cm'm i Iim. 3: '00. 1 8 34 Alarm > Am a piapareti id. I ion) C". Pmsl, Prodi. 
Monoqr. I Cm 

CI III I \ il[ ,n nm r ( b 1 Js O.hO.l >," ii |0. i m.ih I I , \ imniu 1 953: 616]: CGE; 

■(C Presl) A. DC. in 

DC , Prodi 7 3^3 ]t - /»■ i , . '« m> , hi ith [ innaea 38:727. 1874. 

Ih>itmann .Piesl) kum/e, RevrM ,on. PI.' '/,'. I8'H /mPC>i,CC/C vmAu^cnana 

(C . Presl) Reic he, Anales Univ. Chile 1 1 7:459. 1 905. Tmi : CHILE. V* takai-m, 2.1 km S of Punta de 

above the road, ele l i m J I ' 'm , , in hem desiu 

Dortmanna bracts IC r - I I hvlla var bracteosa 

(C. Presl) Reiche, Anal* ; Univ c hil. Ihl'ihu I leosa (C. Presl) E. 

Wimm. ( Ann.Natuil i i i i m . n ' v 1' T r , II 1 C m - tr petrosis collium, 

Oct [1 829], Bertero 

;/ tin > tlh IP i I I i 1 ' m( IP I) A Pi 

in DC, Prod r 7 3^ 1 ' ' ' ^ h. I i H I il >h 1849 Dortmanna 

: > < lih ui • m h i i-i ' . - - • ' t , sopitolta(C. Presl) E. 

VVlMllll.,/\nil II 11 I III I , I I '' IP I I Ml I, ': ' ' "MMMII I u 

In '^ ui m I • mii • _ ml" Pi Ii Ml ' ' ' ;>' 


i hi rofit he']) 

lyphylla var. latifolia A. DC. in DC, Prodr. 7:39 

iesignated:K!;isoLECTOTYPEs:BM! CONC! E! GH 

vpurpurea Vis., III. Plant. Nuov. 2:23. 1844. T 

/ the road, elev. 1 50-220 m, 2 Nov 1 990, U 
). Described 

1997) or elsewhere, a recent colk ti nth ta re 

Tupa ovata Phil, AnaL Uimmntim 1 n i n , M i , [ Mn urn ! ii ,t 3 700. 1834. Lobelia 
oi/atoReiche,AnalesUniv.Chile II 160 IdOS.Tm (Mill Carrizal bajo, Dec 1871, Kings.n. (ho- 
lo-ypi :SGO-043561 [three small branches on right side ot sheet]!; n , , - : BM! K!).The one large 
branch on the left side of the sh > t nth- n r i I L nri'iiM ^ • ] : hi h I wji , 

the manuscipt name "Tupa glabrata Ph." 

Tupa polyphyllavai (ui i it in i i 11 , h\llavar.coquimbana 

(Vatke)Reiche,Anales Univ. Chile II 1 '1 in r r U III B Pi l e LEIqui]: entrada al 

i mi i I f Ii, eral la Mrguesa In i ■ t u I I I j u. t Buenos Am I Oct 

1 963, Marticorena & Matthei 172 ( pe hen ignal ( ) I OS!). Because the 

h| Ii il il i In I hi i i i i i' hi i r en it tin pi it k iu< i I 

Tupa pofppiQiona PhiL.Anales Univ. Ami mm ^0: 1 8rn IS')',. I , -CHk I .« Jui Hot a, Germain s.n. ( n ■ - 
1 e designated I i i i I i i I i t m i I 

Ion it i *■ i t ri t design,. :-0 ,-.; it i the i in FTiilippi's ! i .uui 

* it i I i i t i th i i it I iuh 1 1 i th nit it 1 1 i ii i. in his handwriting. 

h I hil n il I In mil i I I I hil Fei h nil 

Univ. Chile 117:460 189 I t CHIII < oguimbo mmsm. (i - iypi here designated: 5GO- 
057182!).Theothem\nt \ im mi 1 ( i m I m -| 1 - ppisn (SGO-0571 83! 

i' ■ i 1'itiHi] ■ m .'d ] was ii- i annotamd I I i •' rm name. Although 

1 i m I | i p i 1 iti ill I i i i ii i t Lit i was listed. 

Tupa gayana Phil., Anak Uin mtn n I I Mil' - ad rivuios Serena, 

Oct] 836, Gay 1466(i , SGO! [phutograph:GH!]). 

ni ii M tub i i ^ 14 MIL Coquimbo, 1889/90, Geisses.n. (lec- 

one annotated oitl, Hi urn mi, I h i handwriting 
hrubs, 0.6-2 m tall, tun aoocI i [ i I in, I p i ent or glabrous; 

)ng, widely elliptic ell i t m I i i n i In n itelv pubescent or 

leaves, or aggregim I it t i ' 4 1 J 1 n i I u 1 t ~-S0(-75) mm long, 

2.5-28 mm wide, wid I h| 1 llipli n m I lliptu im hi )i ovate,glabrous or 
minutely pubescent; pedicels 7-17 mm long, ebracteolate (very rarely some with a pair 
of linear bracteoles 2-5 mm long at or below the middle), minutely pubescent. Hy- 
panthium 4-7 mm !onq,-1 8 mm 111 diameter, ob< oniq r ampanulate,or rarely obovoid, 
pubescent or subglabrous. Calyx lobes 3 8 mm long, I .3 mm wide, triangular or nar- 
rowly triangular, puU t nit 1 n im utn 1 t 1 1 1 1 1 ■ 1 1 1. t ih ap< < k uini 
nateor long acumifidt- > n ill 1 oiiti I ni 11 | t lit ' m|\ pubescent; tube 

mm long, 0.8-2 11 n I I Ham it toil I mm 1 in either tube 4 7 mm 

long, 1.2-2.5 mm in i 1 r 1 1 > t 1 n h| , 1 1 < < I ibi u 1 the doisal three rarely 

with scattered long pubescence. Capsule 9 -12 mm !onq,6 9 mm in diameter, broadly 
ellipsoid or ovoid. Seeds 0.7-1 mm long, 0.3-0.4 mm in diameter, oblong or ellipsoid, 
light brown orgolden bio n,fainil mam iMuuia i9:,,liqs.52 > 9 Chromosome num- 

Icones.— Lindley (1 830) [as L purpurea}; D. Don (1 834); Hooker (1 837);Loudon (1 844), 
pi. 66, fig. 3; Navas (1 979), pi. 42D-G. 

,' ' , ' , I 11 1 11 n if ntiali hile between 

latitude 27°S and CI l nth t< n m nn >\ nib ■ |l ti Im ih and sclerophyllous 

vegetation (cf Wall tl p ut i li n I | ml 111 ometimes in sandy 

soil, from near sea level (often in sight of the ocean) up to nop n stately as high as 1200 m 

above sea level, often in comp hi itl, M 1 1, through February. 

P/si ms/on I hi 1 1 1 1 1 in mi I m il It 11 h 11 it jin especially size and 

whether the plant I mn 1 oiiscmt 11 1 ite inllon u or I sars solitary flowers in 
axils of unreduced I. 1 Mm m flit t m ,l iln h m i >f the species' exten- 
sive synonymy. Tliouil tl mm h i |C t Tin m i i A limner (1953) as L 
polyphylla l.lmeanlolia and / . ovata] appeal most distiix til was m >ted during field study 
that many populati i i pn lq i i it t li i i | xt\ Individuals that 

differed considerably in leal ienqth, width, and outline olten vvete found growing side- 
by-side. Furthermoft Im th t imi ml h I i irl 1 1 the season often dif- 
fered significantly from those appearing later. In some cases, two or three of Wimmer's 
"taxa"could be found on a single in f i in ill i m | I. it - - nnetal 1 167 (CONC), 
tin I ' it ' - a t. I - tm - i ■ to var. latifolia. This 

suggests that at 1 

east some . 

af the observed variation is environmentally ir 

lated to rainfall 

r day-lengtl 

i perhaps'! tat her than undei genetic control, 

related to heterot 

In detailed r 


analyses of this species (Lammers & Glass 19< 
data, which were gathered from 64 herbariurr 

eluding nearly all 


mmts. Variation in foliar features was absolutely c 

with lid gaps by which meaningful N m hi 1 1 in i /. nvata} could I li tin mi h t I J 
correlations were detected bem.—n t h t [I n and any geographic, ecologic, 
or elevational parameters. Furthermore, the plants examined cytologically (Lammers & 
Hensold 1992) well represents I the di • i . t t folui morphology; all showed n = 21. 

In contrast to foh ir tin tut tl II i t mi nnrkably homoge- 

neous. They are distinctive ithiit, ti i e iu ,« )t then mall size and the dark 

pigmentation/The hue of the comlh i i ntt n d purple which compares favor- 

ably to the color of a good Chilean red ire (" no t into"). The only variation in floral 
features which appeared to be geographically correlated was anther pubescence, which 
showed a clinal pattern of variation: pubescent anthers were commonest towards the 

This species is unusual among Lobelioideae in showing some variability in the pres- 
enceorabsenceof bracteoles H' i J | imen found that had bibracteolate 

pedicels {Marticorena etal IV" ( )fj , I epical ebracteolate pedicels in the 

same inflorescence ' innldi I mil i< t m i i lit ink ate anauxotelic (i.e., 

incapable of renewing vegetal l h tl I tut I - i ith oiowth resuming via 

(Schlegel 2752, CONC.SGO) was seen that was clearly auxotelic,with an inflorescence in 

full anthesis surmounted by ca. 15 cm of n Mi'i it, th 

Representative specimens. CHILE. Prov.Copiapos -2 kmWofTotoral 'aylor et at. 10807 (ASU, 

MO).Prov.Huasco:La,leza, b o-s; ; i m ,' > ^>> r, \\ D); Mina Los Cristales, 

Marticorena et al. 1 700 i< < )N< j tin. o ' , in o , <\\< I i .<h del Pretil, Ricardi & 

Marticorena 3966 (CONC); 3-4 km SW ■ -. " ' "■ • Prov.Elqui:Punta 

Arrayan, D///on eta/. 5457(F); La Higuera, Jan 1 n 153 (UPS);entre 

La Serena yVallenar, Garaventa ' ft tt I n< aHtoguu sner 14 (CONC); El 

Tofo, Kubitzki 279 (CON< VALD Lutil.i H et „iui , A idNi F MU, OSH, UC, US); 

PuntaTeatinos, Lands/)) ' 1><<1> e r in IN, I n j ,iiih h > 0< t 1878, Philippi s.n. 
(SGO); La Serena, Sep 1898, /?e/tne r , , r , i- b , j, h jS > i larticorena 45647949 

(CONC, OS); 20 km Set In jfnne IN > Universidad deTalca); 

La Serena, Sparre 2793 (S,SGO); Net I , tie , < > ,\ I V Vu )) Prov.Limari: Talinay, 

5/7es 439 (CONC); Fray loin <s ,' I i I It I I i,t,t w Hmnilb Lammers etal. 6372 

(CONC,F);11 km Nof Manias I.' H .mill .,--„, -, , ,n L F , , ^ ) iB, CONC, F,MU), 7665 

NY, S); Fray Jorge, Wenh-n un> • ' - I I f , ' i Prov.Choapa: Pichidangui, 

Correa 67 (SGO); Puenlc N 

CONC,GB,F,MEXU,MUl nil N tl i' ifONC f, MU, NY);8.9 km N of Los 

Vilos, Lammers et al " V < )\l I 1 1 I il - it eta/ 7668 (B, CONC, F, 

TEX); 23.6 km S of Lo< Vile Lu e i, - r - i|.,_f ) .1 1, US), Lst.icion Ingenerio Barriga, 

Marticorena etal. 210 (i (m_>, Ann., nmli '," - e,i S5 (B), N of Los Vilos, McGill 1028 
(ASU); fllapel, Rose < J I It II I U </S i mson /6646 (BH). Prov. 

Petorca:4kmNofLongotoma,Lamm^ + • n I ' » J n > UBi 1 f in s of Pa pudo, Lammers 

,'i-; i 'i,_ 'J i 1 . Im N i. t n mhernmost road toZapallar, Lamnu < r, . " "'iNJCF). 
Prov. San Felipe de Aconcagua: Quebrada de Las Palmas, Sep 1965, Ramirez s.n. (VALD). Prov. 
Valparaiso: Limache, Garaventa 299 (CON 2 

/ 908 J (CONC); RenaLi I in t , N F) 6413 (CONC, F), 

6414 (F), 64 /5(F) f I M I [" N I I li 1 b km 5 of Punta de 

Angelesiomme^t' f M ^ n II ' 10 NY, SGO); Valparaiso, 

Dec1851 Philippine ( I r < i mt ' mn JO (BM,CAS E F GH, 

M, MO,S,U,UC, US). Prov. San Antonio: El U, , , * ) > >' i< ' Aanobo Lammers et al. 

7800 <B,CONC,F),Ouehi HI -l Region Metropolitana: 

PpisVeick . oun< s< / i i|h| i ,|| , ; n , , na Navas 2298(GONC); 

hum del i I iuu I "ki |u o ' t 

CULTIVATION. U.S.A. California: in - t it ' At da i, i-don Berkeleyjul 1963, 

2. Lobelia excelsa in I I i I Alt , nl - jm (Bonpl.)CPresl, 

Piodt f 1 n i.|i I ib, I - 'o . ' , , < lt , it n| I i uni .hi, oen PI 972. 1891. 

As no original in il> i .1 was In jt> 1 •• • plate | i.[ h L, I .ill " , | t T, , m< is heir oesiy 

oweA, meowee dms.Boi.Mag. ASm AAA 'h'Giirr hav.Awsilcs Hist. Nat..': h'4. I 8i ;:. m owW; 
salicifolia Sweet, Hon ul uil hi I 1 , ' n uweet) G.Don, Gen. Hist. 3:700. 

1834.Typf:CHIIF Val| i n 1 1 \ I i i i nit i EM' itm i n MO!), 

nercial nursery. 
he/ i itiiut i I in<ll n I in I i I i I i I I - , > il nd i G hon, Gen. Hist. 

3:700. 1834. Doitn i II I it II 111 hE A I I I IAIN. 

hi I did i V| ~ I tit I, nut, l I in IE I^p|r3i|\r mi il mm 

rial was located, the plain published with ihe pioi, ,h„iu, is here designated as the lectotype. 
A mn • ,,'M n hum tr, H r L ■ I himi, I 2A1H-MA HI' E I _ G- : 

elev.45 m, 15 No. I""" ' , <<>< < > n, nhm ■ <v i i i hem dm ninat, d OME 

isoneotypes:CONC'F'M 'h ■'" ' it I th h Hi in [ ml ,,i , vn in the botanic 

,]aitl, n it Turin It il hi I it I I it i u u ,in I m , , n i i h m in pe, im- n 

could I , Eh it ,] ,i i nt , || , t ,ii tint ' A in 1 tl ; i ' l> < m r I t d> n ,t d i 
the neotype. 

Tupa glaucescen*. Phil m t tu I I II ,n i I i I , t 1 -- hj 

s.n. (holotype:SGOI). 
Shrubs, 2 4 m tall, rarely trees to 6.8 m; stems woody, repeatedly branched solid jla 
brous or mtnuh I pu! it it ' m ii il I unit i IS em long, 0.7—3.1 

' in i' ' Ah ,h i i i i I ihl 1 1 i , I | i I, nt ' II 1 1 1 1 nl lanceolate, glabrous 

or minutely pubescent; margin serrulate, minutely simulate, ot subentire; apex obtuse, 
acute,oracuminat, i m n- ei p ht i iii i n i t it it 1 e tounded, obtuse, 
cuneate, or attenuate. I sowers solitary in the axils of the uppm leaves; pedicels 12-45 
mm long, bibracteolat< n the .wor tl id or rarely at the midc 
bracteoles 2.5-10 mm long, lineat, minutely pubescent. Hypant 
15 mm in diametei I , mi i o- is hi itl hemispheric, btoadly i 
broadly obconn tl ,1 i, nninit, 1 p , , ,t d I - ^(-20) mm lon< 

mm wkle, nia tin u la i or n.iitowlv hianqul.n, minutely nui en ei it ;apex acuminate oi 
acuminate. Corolla (38-)45-65 mm long, yellow and orange in hud, becoming i 
intln i I i t it I , |l ihl il i i n I | ill i ill ti nut h in 

mm in diameter, pale straw-colored, the dorsal three with long white hairs on the sur- 
face (especially toward apex). Capsule 10-15 mm long, 11-18 mm in diameter, ovoid, 
obovoid,oblate,orsubsphertcal.n J t i i M l n il I ijncj, honey-colored, 

faintly striate (Murata 1 992, figs. 49-50, 70). Chromosome number n = 21 (Lammers & 
Hensold 1992; Lammers 1993a). 

Icones.— Sims (1810) [as L glgantea}; Bonpland (1816), pi. 46; Lmdley (1826) [as L 
arguta);Munoz (1 966), pi. 1 28 [as L tupa]; Navas (1 979),pl.42A-C [as L salicifolia]; Hoffmann 
(1997),pg.218no. 1 [as L tupa]. 

ni^triinuionjiobiioi, and Phenology. : ndemic loxeric noilh mimyi c hiie between 
latitude 29°S and 35°S,in the regions of dwarf- and xerophytic-shrub and sclerophyllous 
vegetation (cf.Waltei 1 r ,Fh , m i , |, , i | , mil onetimes in sandy 

soil, from near sea level (though . I mi i ii i i1 ill > ^ m i i| n ) 1 200 m above sea 
level, often in company with / / ' M >mii<i > | 'Mini ) i through February. 

The report of this | n- n m -i h I i > i i i MS 3) is apparently due to an 
inadvertant switching ll I i i i piesentstheonly known speci- 
men of L. excelsa from Bolivia, wl '<.'< > i| >f losedly the only specimen from 

Chile of another lobelioid, Centropogon m" , I l'tu_e ' 'nu h is indigenous to Bo- 
livia (cf. Stein 1 987). However, duplicates of Rusby 634 at GH and NY are specimens of L 
excelsa from Valparaiso. Clearly, I ib> I i < I M it h< I it some point, result- 

Discussion — In contrast to/ / > m I hi x>dy member of the section, L 

excelsa is relatively homogeneou mil i in u uphology. One feature that does 

shrub, one individual was encountered in Prov. Limarf in 1 989 {Lammers etal. 6382, B,C, 
CONC, F.MU) that was definitely a tree.The plant was fully 6.8 m tall and did not begin to 
branch until 2.5 m above the ground. Its twin boles were 1 1.5 cm and 9.5 cm in diameter 
just above the base and were oi ;ufficienl ,i i th that the tree could be climbed to 

- ill. i i flo' "iiii'i bran< In : 1 1 f ■ ■ - n damage 

regularly bibracteolate pedicels, ami for the c oloi change of its flowers, which go from 

biological significance of such l )l h m -i i mied by Weiss (1995). 

Plants bearing pure ulfuivln. - i ill h trace of orange or red (and thus 
no color change) are known, huM nl ir i nh iImm.TI Ml Y a specimen culti- 
vated in Switzerland [anonymous s.n., 29 Jul 1858 (NY)] states "flor sulphurei." More re- 
cently, Eric Knox (pers.comm.) report* 1 1 i il | n 1 m t,M it i M ersity of Michigan's 
Matthei Botanical Garden from seed of Lammers etal. 6393 consistently bore yellow flow- 
ers over a period of three years. I know for a fact that the plants from which the seed was 

ii 1 1 i ti tin ill f H mi n ll i I n i it r i r I in i il plants with 

yellow flowers ever (j i in i it i | ■ 1 1 i mil mm I i I it conditions. 

Representative spec-it, -, CHILE. Prov.Elqui 1 ] iesta 1 

Buenos Aitewu/o/YU II' ii. tul.hui ii' i, , i J HA if CONC,F,OS,UQ; 

Mormon J6328 (BH). Prov. Limari: Mi. >nu I,. -I ndn i' i m J' )n. btada La Higuera, Jfe 

O3UC0NC) Cabrei i > M ON F , I , 7 (t ONC); 1 km N of 

efa/.7664 (BM,CONC,F).Prov.Choapa:l I nr U> tO t , n / K v,| n,o 2 km N of LosVilos, 

Lammersetal.6330 1 k I ' U ■ l ' OH i f - 1 O im • . il ef al. 6348 (CONC, F, 

MU,NY);11 kmEof Pin mi. ik in hi in i Rltofanelal n > << / .0 6353 (CONC, F,GB,MU, 

NY);5kmSofCanelal m .- L 1 1 I f i h,, Nl if I llapel Lammerset al. 

6362 (CONC, F, MU) On L I III i| «l ,- i + il III U m sWof \\\ape\,Lammers 

ef o/. 6367 (B, CONC I MM kin if Puente Am ^hn i > f < n (( ONC, F, MU NY); 14.5 

Matthei 427 (CONC) III ipO .' ", u iN I Ml ' n ON( ) Mon(,vo )()7 7 

(CONC);lllapeltoH'i Prov.Petorca. '.apaWar.Gardner 

&Page5085 (E) Pichum .< if!/ <^ I 'N In' mil t f ipudo hammers era/. 6399 

(B.CONCFMU) 1 km O , M I L i n 51 Un N of Nogales,/.ammm 

ef a/. 6408 (CONC, F, MU N, MkmN.itN i if , i-h, , - )ff F, MU,NY); 1.3 km N of 

(CONC); Papudo, Montero 8023 (CONi n n m , A '32 (SGO).Prov. San 

Felipe de Aconcagua I h I i II I II OS) Quebrada de Las 

'. '■ i ■■■■( ■ , ■ < '. Y ' Prov.Quillota: 1? i m from'iltil towards Olmue, Gardner & 

Knees 5446 (E);Olmue, 5 "■ « M - i In o\ ,inF. I i , -< i <( O o4/8 (CONC, F,MU, NY, 
UB); Parque Nacion )NC, F, MU), 6450 

(CONC,F,MU),6452(CONC,F).Prov.Valparaiso:\alj n i I la^Zonas.Harshberger 

1074 (NY, PENN); Quebrada de la lorluga M Feb I'm ,/-.,„ ui. Y ON< OS); 9 km N of Concern, 
LammersetaL6417( - • n Y - ,', ' CONC,F,UC);3 km N 

of Laguna Verde, Lammo < ' u . Uuihi I lorn I , /• -^ , (ASU) Mirasol, /fandrum 
3822 (MICH, NY, SGC -t in ' I I I ill f . 10,UC);Valparaiso, 

Dec1862,PrY//pp/sn u. n U ' 1 .i < . / ' II Y ,i , / vb34 (K,MICH,NY,PH); 

Valaparaiso Rusty oW UI J Yul til , K i , n Oalpaiaiso, Sch/ege/ 967 

(CONC); Quebrada V( i i - , - f I - n ua & Skottsberg 947 (GB, S); El Salto, 

So/br/g efa/. 3600 ( CYI II. Umll Ii n n - i,' - , > < miNt , F, MO, WIS). Prov. 

San Antonio: Quebud , < . Joh i <" ^ U,i-Y,,l t Y , i-mho 'ammers ef a/. 7795 (B, 

BM,CONC,F);NofSan « h , ti i i I , I l< I , » II i I Ft I I "b8, Santos s.n. (VALD). 

RegionMetropolitana:Ou. hud,, hurmih , m "i ' •? (CONC); Cerro La Cruz, 30 Sep 1932, 
Olathes.n. (CONC); Dun i kl ui I ti ) - u ONC OS); Quebrada La 

Plata, Schlegel 1680 (CONC). Prov. Cardenal Caro: Pi. I, in inii \u^r, • ^ '86 (CONC); 9 km S of 
PichilemuOommtf N [if tol 79/7(CONC F NY) 

4.2 km N of Pin aluiui , ' \< ' i I fklilunu , t >/ ta zai 573 (SGO). Prov. 


]■!,,', Prov. Curico: i 1. \ , , ■ p mavida, Spooner & 

Contreras 4332 (CONt f I ' iint i I r nil F I l <> Ocm n (SCO) 1 km antes de 

; Y . ■ '■■ Prov.Ialca:. v. ..■■; .V ■, .- . ?4 (B);Constituci6n,Feb 1895,P/i/7/pp/ 

s.n. (SGO). 

CULTIVATION. SWITZERLAND; Hen [ ,r L < ,1 Y:' "il 1 558, anonymous s.n. (NY). 

3. Lobelia tupaL t M ' >l ti.iin I nllu ?oo 1834. Rapuntium 

[ r r Tr it i . ( itke Linnaea38 727 

]874. Dortmanna <• u ' i I i h^ n I '1 > r ,< w << < ,'c 

i i i lie ii I Uni i nil II III rill M mi ioCu hi uch i Net n (i i 

MA! [photographs r' GH' il II' ' i ,ni i l 1 ' 1 'I 

Lobelia serrata Meyen.Reise 1:300.183---. I > -"II I I Hacienda Monte Grande, alt. ca. 

600 m, Dec 1924, Werdermann 572 (\u> ,pf here designated: BMI, ,, ^uiypes: E! F! GH! M! MO! 

I i i f I i nn il i i ii il i I ! I i i i i II tion from the same 

general area, which aqie- il (i i in i mated as the neotype. 

Tupaberteioi A DC in L i Ft Ii 1 rim ia DC.)Vatke,Linnaea 

38:727. 1874, bertun f <> > I I mil I i nil i '2 1891 .Lobelia tupa 

var. berteroi (A. DC.) Reicht Analc Un Chih 1 17 ^- inns berterii.' /.obe//fl mucronafo var. 

bertero/ (ADC.) E.Wimm Pflairn,,, | , i l Hit in 1 i HII F.Rancagua,in sylvaticis 

umbrosis.torrentesetnvulu v P i I i>>( LX [nn< mtiLhe'l n h iffl 

Wimm.,PflanzenrlVJ n| -, i 1 < 3REAT BRITAIN. Scotland, Glasgow Botanic Garden, 

Aug 1832 (KjLoivni: Hooker 1833, pi. 32071). 

p hi ii h I M ; ..:-:;: - - -, : -..:", ii h ■ :.,riseb,, 139. 

\S66. [)ot!man!hi[)hiiippiar:ti KLinuv,R>'v\Gcn.R!.\)/2. 1S ( H./ nbei/n /npr/ var, /no/if orw Reii he, 
Anales Univ.Chile 1 1 7:459.1905. IwnCI III |-.Hac[icnda].dcl Principal, }87Q, Philippis.n. (noinivft: 

i i GOm - Ml I i Jph GH!]) 

upo Vi, '/e/ ^ar ni (ll M/ ' , a ,jtl - hi i i "4 1 T BRITAIN. Lowe's Nursery 




-:0-\\\\. Ruiz & Pave 

Robust perennials, 0.5-3 m tall; stems several from the base, normally unb 
iffruticose, densely short or long pubescent; latex v 
7.8 cm wide, ovate, oblong, elliptic, narr I llipti 
I i I i it | ul - r ^r: in - i i Imi i 

mucronate or cuspidal ha < u h ntt at ■ 1 M . : >. n t oi basal iobi > I' utrent on the stem for 
2-30 mm (rarely tun i- I u 1 i t n nt tl i e t a 10-65-flowered 

raceme, densely shot i pub cenpbran 7 1 i 8b) mm long, 2 l r i '0) mm wide, ovate, 
lanceolate, or rarel\ In >i th< if .< ioimn it Mi- in mitiii tin 1 pair of basal lobes 
decurrenton the stun t i inn i t i i i t tt ti h t tn) lately obtuse and 

non-decurrent;pedicHs 8 A) mm lono.obrk leolaie. I Ivpanthium b -10 mm long, 8-14 
mm indiameter,hemisphenc d< pn II i i \ h i ' n i In ndlycampanulate, 
densely short pubes< onto alvx lobes 2 8 mm Ion* j, I -3 mm wide,triangularor narrowly 
triangular, short put i. m i| i imiint non imiint mi k anute.Corolla 31 - 
49 mm long, reel (wi mi I ^ II fit I t ' n h 1 1 pubescent, tube 19-35 

mm long,2-4mm indiameierat tnicb;b\. au. mitt lobe: It) 2 mm long, 1 -2.5 mm wide. 
Filament tube 22-32 mm 'ong, red; ant her tube 6 ( -> mm long, 2 3 mm in diameter, grey, 
the dorsal three pubescent with long white hairs oi t a rely olabmus. Capsule 0.9 cm long, 
1.2 cm in diameter, ovoid. Seeds 0.7 mm long, 0.3 mm wide, broadly ellipsoid, honey- 
brown, minutely fovi lull n ti ul ! <M n n , | h> ti i i vn i hromosome number n 
= 2 1 (Vilmorin & Simonet 1 927; Spooner et al. 1 987; Lammers & Hensold 1 992; Lammers 
1 993a); the voucher (Sanz573, SGO) for Sanz de Cortazar's (1948) report of n = 16 in L 

event (Lammers & Hensold 1 992; Lammers 1 993a). 

/cones— Feuillee (1714), pi. 29 [as"Rapuntium spicatum ...' |; (bovunilles (1801), pi. 516 
[as L mucronato]] Sims (1 825); Sweet (1 827-29); Hooker (1 833) [as L mucronata]; Lindley 
(1833); Loudon (1 844), pi. 66, fig. 1 ; Santa Cruz (1 932), p. 99; Wimmer (1 953), fig. 96; Finnis 
(1966),fig.48;Schultemmi m, p pn,C »M>,g I 2 \ Matthews (1988);Thomas (1 990), pi. II, 
no. 5; Hoffmann (1 997), pg. 21 8, no. 2 [as L. bridgesii). 

i ] isi r il n ituin, I lahif, n,, in, i i'hcnokhjy. -I in lenin to mmtc south-i emtio t. Inie between 
latitude 32°S and 4_ i th i i i t li i I n i u en temperate rain 

forest (cf. Walter 197 m Also naluiali/ed on Masalienu (oLi Robinson Crusoe) in the Juan 
Fernandez Islands sometime prior to 1 824 (Matthei et al. 1 993). Common on roadsides, 
fields, streambanks it i \ I | mil i i m nun it I it n fiom near sea level 
to early April. 

Discussion -I oh, ' - ; it i I i I it th n it o -! t I. if md bract bases, 

variation which Win ii I i Ct i it i i i , illy, the leaf margins 

continue as a pair a bi n i mini niricl fo m lit , below the point of at- 

bracts, as var. tupo oi (if the leaves vvete lam eolate) \<,u. pavonii. Those with very long 
auricles were vai.moniann or (il the auricles of the bracts were ftee from the inflores- 

current leaf bases wen otjreguted i ) ,t , Plants h t re intermediate be- 

lli n n it 1 th ilti it it ninnies were treated 

as /. rnuctonata van/ovaeme the in ten mm I on y o ilns vouetv n niohlighted by the fact 
thatVatke (1874) and Reiche tbn Mm [m ton L ,l u I it to L tupo 

Careful study of i Mti ml i t , In' ihe length of the basal auricles 

varied considerably within populations and even within individual plants. This was ap- 
parent even in herbarium material. In Gay 1468 (SCO), the auricles were only 3 mm long 
in the basal-most leaves, but I S mm i i, 'ilm i inflorescence. As such, the varieties 
of L. tupa recognized h\ Wimn m 1 - l L, mS) do not merit recognition. 

The case of L mui > > m , i m m| b ml i net t i i ed by Wimmer, it 

differs from L tupa not only in n non <le< ummt loaves, but also in its very sparse inflores- 
cence (cf.Hooker 1,sr>i Hm m m i L , Mm before the apical meristem aborts; 
the flowers appear to be largely in bloom all at once and to take on a nodding rather 
than spreading orascending po tu I II ih. i i 'in t the f)lants, including 

the flowers and the e e b n I- it ,1 1 _ r utti - tmore,all specimens referableto 

e been collected \ith n iln i i n phi men I >nal range of L tupa. 

; unable to locate plants in nature that matched this description, and have 
a few specimens besides the types cited above. While it is possible that L. 
'Xtremely rare oi p ill hit >p< mes, perhaps adapted to some 
unusual edaphic nu h otluh f I in I I ' I II I mdful of specimens 

examined may simply represent plant t ip hose apical meristems were dam- 
aged at a critical point of development by some insect or pathogen, resulting in aber- 
rant growth. They may represent ui 1 i i! no . ount | i i haps a simple Men- 
evolutionary divei f these two species. 
Although they are not sympatric today, their ranges do approach one another (see above) 
and may have overlapped in the past. The most recent gathering of plants referable to L. 
mucronata (Montero 736, CONC) was collected in 1928 in Prov.Colchagua,in the zone of 
closest approach between these two species. 

Till u h tin i nti i i 1 i it | mi 1 ii | i i i i i oi of some of these 

hypotheses,! am loathe to recogniz tint [ ies Because it occurs 

within the geographic range of/., tupa and because its flowers and seeds are indistin- 
guishable from those of that species, it is here relegated to synonymy. 

Populations of /_. tupa also shim I i ,tmn i pubescence. The hairs on leaves 

surface of the anther tube becomes increasingly pubescent from south to north. The 
corolla of L. tupa typically is red, though it does not change color (Weiss 1 995) as in L 
excelsaA have seen just one specimen (Reiches.n., Jan I902,SGO) in which the corolla is 
yellow ("flores flavescente"; cf. Reiche 1 905, 1910). 

Representative specimens JUAN FERNANDEZ ISLANDS. Isla Masatierra: in fruticetis apneis 
collium,fiertera/24" i i Mnhm _ )l iC M,OS);Valley Anson, 

Meyer 9580 (MO, RSa I ) ii I m , i i i r t , > \ o291 (B, CONQ.Valle 

Colonial, Spatre 7 (CON( w ,| > <> , , «< / '- , Knf? s HI i between Pangal and La 

Centinela Sfuessy&t M t rl Mi irl ,n 'uan Bautista,Srivessy<S 

Crawford 63 /rj(CONi t m ilun in;i , t etal o200 (CONC,OS). 

CHILE. Prov.Cach,v"dl i n i i I n, , •, , >s,3 Nov ]952,Pfister 

s.n. (CONC), Prov. Colchagua: I i M, , , ^ , F haurrina, Montero 736 

(CONQ.Prov.Curico:! Prov.Talca:Constituci6n, 

Feb 1 895, PMpp/sn i ' n, hi . 1 ' tl i niu mm Nov I891,/?e/c/ies.n. 

(SGO)Prov. Linares: I hil i i i nil una de Catillo, Montero 6276 

(CONC). Prov. Nuble:a Quirihue despues deTiehui ' I Prov. Concepcion: 

Parque Hualpen, Carmsco 333 (CONc l; < p. ion hcio/< ' ' n iOMI.Ui i ( oriLcpaonhhof / // 
(BM,NY);PuntaHualp. n ,-if IIU 1 n F r I <v eta/. 6376 (C, CONC, 

F,MEXU,MU,NY)?Ukm if mhli n / t < I II Ml I h\ i hi Soldadohammm 

et al. 6329 (C, CONC F h MF f J h tfl t h fh60 (B, CONC, F, MU); 

Concepcion,/.^ I m i nt ' - ahlhhl ilcahuano, Dec 1861, 

Philippis.n.{SGO),Ja\ uhu -" ,*>->,. f „,hdi -fuessy ef a/. 668? (OS). 

Prov.Arauco:Contulitu . < ' in N i in i .t i luln i u i K Boeza6510 (CONC, 

F,MU);Laraquete ~>QD I I 1 h M hi 1 i Lebu,Spooner4483(F 

WIS). Prov. Malleco: , • ,„ , ammers&Baeza 

6509 (F,MU);Nahuelhut - l h ' i ' nt ih hi nn > h ' (SGO);entre Puren 

y t onLultno ," ,< . ' II Prov C.niui 

Trovolhue,/WonferoPh hh - Pmv.Valriivia 

Valdivia, Bne/ges 66 / (BM,F R hi mi Nf ot 1 ii i I J GO, UC, WTU); Niebla, 

Garaventa5533(COHQ;Hu\eco\\a,Gardner&Knti ^ ' I I i j * h llm t 

hwy on road to Corral, Lammers & Baeza 6463 (CONC, I, ML' J; hs km I of Corral, Lammers & Baezo 

6464 (F, Mb); 31 km E ohm ml , '' - /, d ' H hi, J >l ' I. nhui ■ I »m 

Baeza 6478 (GOHi F Ml h) hi F >h ' bun t , t a 6496 (F, Mb); Mehuin, Lammers & 

Baeza 6503 (B, F, Mb) id m ' .f s m los. d h hngnr , ■ ", \ r ,, (J 6506 (F, Mb); Corral, 
Rudolph 6097 (VAFDj t i I Ih ' I h bid h/im/Tz 9/ (VALD). 

Prov.Osorno:Alenci| II It l h , < I h) i '" hVM hi < > 

(VAI D); La Barra d I Prov. Chiloe: ■ di '.2493 (F, GH, NY, 

PH),25Dec 1951 Pfi I ' dJ 

CULTIVATION. U.S.A. California: I CFRViAfs'i 

\. Lobelia bridgesn 

Arn.)CPiesl Piod ',, n n bh - > h HI in \ DC, Prodr. 

h 1 i ) II I hut I hi hi I ii hill E. 

d ii id n« nEI t I ii n i i I i I lhh| TfJ.ii- i 
[photographs:A! F! MICH!]). 

TupablandaD. Do ni.h t BiihluH-ho bh ' 1 blandum (D.Don) C. 

1:437. 1842. Dor? 

hni hid M hi hi lib iii it i ii to i i ,n here designated: D. 
Don 1835,|il.308!bAsn(H:ngiiialmatoi!alvv,ihoc ( itih.;h l 'pid'j.'pubi5hedwiththeiiiotok)<uie 
is here designated as the lectotype. 

in It )di ,- I ' lilt n II < 1 18m 1 ii hh 

Sep 183.5, mionymi 
Robust perennials, 0. 
hollow, herbaceous c 
cm wide, lanceolate, glabrous; margin minut 

Oil I III 

5 55-1 lowered Mntm lnnl. Cm eolalo, IS -60 mi it lono, l he bases, 
' basal lobes decurrent on the stem for 6-22 mm; pedicels 14-30 rr 
ousorsparsely pub. i M m | o a lituj hairs Hypanthii 

5 mm long, 6-10 mm in diam< t< t I in lm t I i i hi us Calyx lobes 5- 

olla 25-36 mm long,pink,glabrous;tube 13 10 mm long,2.5-4.5 mm in diameter 

g; anther tube 6-7 mm long, 2 : mm u Imm im i I i ,il three sometim 

with scatterec 

I long 

spreading hairs, and/en all Cm nubescom 

itwithshortappressed hairs. 

Capsule 16-1 

i long, 13-14 mm in diameter, ovoid to 

subglobose. Seeds 0.5 mm 

long, 0.4 mm 

in di, 

ameter, broadly ellipsoid, golden tan, rr 

in I 1 . I • m J in It tl< ill il 



bern = 21( I, 

smmers 1993a). 

I cones — 

D. Don (1 835) [as T. blanda]; Hooker (1 839); b 

emaire(1843) [as T. blanda]; 

Loudon (1844 


.6, fig. 2; Lamms i I" - I 


on, Ht 


immediate vicinity of Bahna 

San Juan in Prov.Valdivia (latitude -imMQ'S), in the evergreen 

tempi i ite rain forest region 



.outh-central Chile, at elevations from ne 

ar sea level up to 200 im Cot 



um >[ i I i n I i t tn mini . nni titan nuom| tn\ 

with L tupa. I 

t is rr 

lost easily found along the road that n 

ins from the main hhihwav 

south ofValdi 

via tc 

> the In he coasta vi aim of Corral. Aiso 

lecten or . (from mtt 

duced plants? 

) in Pr 

> ( ) ' inn I i I i II nl o I I ring mid Decem- 

ber through e 





dimmer (1 953) distinguished L blanda (ir 

him I i i /. m n -ma in /. 

Lammers 1993b). 

Representativespecimens.CHILE.Prov.Cautin: imailh i I ) Prov.ValdiviarCorra 

Brooke 6985 (BM); 24 Cai sF t ml il 1469 (5GO); Amargos 

Gunkel29 (BM, F) Corral ^ - I 1 1 J I , i ,018 (NY);Castillo Sar 

Luis a San Martin, klempui i\AIM a, h i , mN< oO) 31 km E of Corral 

Lammers &Baeza64t <B HJ F C I I d lers&Baeza 6470 (CONC,F,GB,MFXU 

MU); 2.5 km S of Corral, Lammers & Baesa 648 r > a ml I to t, > - km F ot Corral, Lammers el at 

7856 (CONC, F, UC); Anmm< ' IJ i I t , n n (SCO); Amargos 

Corral, West 4880 {G\-\, MO, UO.Prov.Osorno . LlanquihuenVmnv 3obre,l ago Llanquihue, 1939 
Santa Cruz s.n. (BH). 

CULTIVATION.GERMANYrMt • to in ( M' " ' ' M>,30Aug 1867,/Cumme, 

Lobelia excelsa Bonpl.x L.polyphylla hook.& Am.Tupa kingii 

'"' i " l lu I m i I 'I 1 ,l f i- ,i o Poppe's [PoeppigC 
i | i T" ' 1 1 l ' i anh GH!]). 

Presumably a st 

irub;stems woody,glabrc 

ina 4.8-1 


3 2.4 cm wide, 

oblong, glabroi 

js; margin minutely serrul 

ate, part 


cuneate. Flowei 

, 1. 1 ,f . t m 1 1 into a 18-flowered 


ence; bracts 21 

3-38 mm long, 

'1 9 trim wide, < 

Dblong, minutely pubesce 

?nt; pedi 

eels 15-1 

8 mm long,ek 

xacteolate, mi- 

nutely pubost t 

?nt. Hypanthium 5-6 mn 

i long, 8 

;-10 mm 

in diameter, 1 

-lemispheric oi 


lulate, minutely pubescer 


lobes 5 n 

imlongj mm 

wide, na"Owlv 

triangular, mint 

ttely pubescent; apex aci 


Corolla ■' 

40 mm long, a 

pparently dark 


minutely pubescent; tub 

e23 mr 

n long, 2. 

5 mm in diam 

eter at middle, 

curved; lobes 1 

7 mm long, 1 mm wide. Fi 


:ube21 n 

im long, dark i 


anther tube 6 t 

™ long, 2 mm in diam. 

ster, pale 

? straw-c 

olored, the dc 

rsal three with 

scallered long 

white hairs on the surfaci 


I apex. I i 

uit and seeds 

not seen.Chio 

mosorme numbei unknown. 

1 , ;ion 

— Though Tupa kingii wa< 

; treated 

as a synonym of L polyphylla by Reiche 

(1905, 1910) and Wimmer [ I 953), the typ 

»e appears to rep 

eseiii ihybri. 

i between that 

species and Le 

xcelsa. It resembles L exa 

>lsa generally, par 


size and shape 

of the leaves, b 

-ut differs in its well-demarcated 


em e (vs. Iloweis solitaiv and 

axillary) of darkc 

?r flowers on ebracteolate 




it ' .,' w , 

Various quantit 

r Live features of the flowers are in 

termediate in size: pedi 

eels 15-18 mm 

long (vs. 7-1 7 n 

im in L polyphylla and 1 2-45 mm 

in L exce 

w h pel tin 

jm 8-1 Omm in 

diameter (vs. 4- 

8 mm in L polyphylla and 

8-15 m 


iO mm long (vs. 

15-25 mm in L 

polyphylla and 38-65 mr 

n in I ex 


th tube 23 mn 

1 long (vs. 7 17 

rti ti t i - / h . it I 2 42 mm it i t 


id lobes 

17 mm long (v: 

i.4-1 2 mm in L 

"MP'ii mm] 

I 2-33 mm in L exce/5fj);and Flame 


>1 mm long (v: 


"',,'',. ml 

29-47 mm in L excelsa),^ 

her tube 

6 mm long (v 

s.4-7 mm in /. 

■•''.; >\ , in ! ' 

Ml mminLexce/so).Asr 


specimen is th- 

5 sole evidence 

I have seen of hybridization between the 

?se two : 

iym| iatrii 

: species. 

Field work in Chile during I 989 (including lit to CON ,0 ALD, and the Univer- 

banum Fund of Miami I in it t i I n I t n I i i iieaitfelt gratitude to 
my many collaborator at the Utiiversidad do Com epx ion, who accompanied me in the 
field and in general I ) their wonderful nation, espe- 

cially Clodomito M i ti i 1 1 1 ' - u Ch - ' n ' 1 ml i M i> Ouezada,Marcello 
Baeza, and Patricio I t I i t II i u f h I 1 tntions and staffs of 

visits: A, ASC, ASU, B, Bl I, BM, C, CAS, CONG, E, h, GB, GH, ISC, K, M, MA, MEXU, MICH, MO, MU, 
NY, OS, OSH, R PENN, PH, PR, RSA, S, SGOJEX, 1 UB, U, UB, UC, Universidad deTalca, UPS, US, 

VALD,W,WIS, and W1U. Finally, special thanks to my colleague Neil Harriman for his cri- 

Bentham, G. 1 876. Campanulace D. Hooker, Genera 

plantarum,vol.2.Reeveand Co., London. 
BiAcow,N.W.(ed.) 1 972. Martindale.The extra pharmacopeia, ed. 26. Pharmaceutical Press, 

Bonpland, A. 1813. Description des plantes rares cultivees a Malmaison et a Navarre. P. 

Didot, Paris. 
Candoli f, A. de. 1 839. Lobeliaceac. Pp. 33'-) A i 3, 784 787. In: A.P. do Candolle, Prodromus 

systematis naturalis i> urn * i t d il " m ^t .Vuttz, Paris. 

CavaniiifsJ. 1801.lconeset descnf t i |l I nun M (> Regia Typographia, Madrid. 
Chittenden, FJ. (ed.). 1 923. Perennial lobelias at Wisley, 1 92 1 . J. Roy. Hort. Soc. 48:239-240. 
Don, D. 1 834. Lobelia polyphylla. Leafy lobelia. PI. 242. In: R. SweepThe British flower garden 

(ser.2),vol,3 lame h I i ml | ondon. 
1835. Tupa btanda. Bin I i > ltu| C 308 In; R.SweetJhe British flower 

garden (ser. 2) vol. 4. James Ridgway and Sons, London. 
DoN,G.1834.Ageneral h t r it h I I i I , I n I o mi I E.Rivington, London. 

Feuilltf, L. 1 714Journal des obser it n \ I |u n itnematigueset botanigues,vol.2. 

Pierre Giffart, Pans. 

Hill, R.K. 1970. Pyirolidinc, pipoinim f i im h uti.imIi aldloids. Pp. 385-429. In: 

S.W. Pelletier, ed. Chemistp 
Hoffmann J„ A.E. 1997. Flora silvestie de Chile, zona araucana, ed.4. Fundacion Glaudio 

Gay, Santiago. 
Hooker, WJ. 1833 ' tL h up pointed Lobelia. Bot. Mag. 60:pl. 3207. 

1837. Lobelia polyphylla. Many Mmed lobelia. Bot. Mag. 64:pl. 3550. 

1839. Lobelia bridges ' i G I I h E )t Mag. 65:pl. 3671. 

command of Captain Beeche\ I- il I ii.h.Hh botan f Caotain Beechey's voyage. 
Henry G. Bohn, London. 
Huxley, AJ.(ed.) ]992.Lobelia. Pp. 104-1 06. ln:The new Royal Horticultural Society dictio- 

h-'.-NirJ. 1 955. Fanorogamia. Editorial Univet sitaria, Santiago. 

Knox, E.B., S.R. Do tin md i b Pi m ll ° 5 ( hloroplast genome rearrangements and the 
evolution of giant lobelias from herbaceous ancestors. Mol. Biol. Evol. 10:414-430, 

Kuntze.O. 1 891 . Revisio generum plantarum, vol. 2. Arthur Felix, Leipzig. 

Lammers,T.G. 1993a. Chromosome numb' i I i n i I - ,< I Review and integra- 
tion of data for subfamily Lobelioideae. Amer. J. Bot. 80:660-675. 

1 9 ?0-75 + pi. 220. 

1994.Typification of the nam< Campanulaceae) 

published by Wilh i H , n I i I i , M s pc cimens.Taxon 43:545-572. 

Rhvlooenv, hiogeooiaphv, and svstematko of the Wiih.leahagia 
;omplex (Camp in n i i< itn| it il ul i i > I 1 ' 1 1 5. 

- - J from ^ 

■■■ , P'MS.iVlotphoiogiwil vacation r; Wo i oi oov pohgivgia complc- 
) of Chile [abstrac 1]. Amor. J. Bot.SS (6, suppl.): 140. 

andN.HiJ n i 1992. Chiomosumo number ofb impunuLu eae.ll.The lobelia 

upa complex of Chile. Amer. J. Bot. 79:585-588. 

i C 1843 lupa< If i mt H )il Lin ei < I 1 '59 262 \ plate. 

. ,,'VV.I I ill I ' r I I I ,'. '. I ' I -II il t I ill loWl I ii I ■ , N I 1 1 

ji.eyJ. 1826. Lobelia ar una I inc-t - it In Moduli hiwards Bot. Reg. 1 2:pl. 973. 
1 830. Lobelia pi itpaie< i. Purple lobelia.; cbvaoM Bot. Reg. 16:pl. 1 325. 

. 183" w - i I 1 tup | i n | b ii I i I I t hi i 1 g I 

jAtus.C 1753. Species planturum. 1 aurcntius Salvius, Stockholm. 
1762. Species plantaium,m1.2.< aurcntius Salvius, Stockholm. 

■■ ■•\, J.VV. I844.1lic Lures" Howe uauLm of ornamental pewniuuls. WilLm Smith 



le hlpnosis. Audi 


articorena, C and M 


/ada. 1985. Catalc 


and R. RodrIgui . 

*(eds.) 1995.Flor 

a deC 





Fernandez. Gayam 

i, Bot. 50:69-1 02. 

,atthews,V. 1988. Lob 

elia t 

upa. Kew Mag. 5: 


,un'ozR,C. 1960. Last 



ad de C oncopci 
chipielago de Ju 

sidad de Chile, Santiago. 

I960 Sn ; lb mm! ii- 1 i u bll I mil u • 

Iurata, J. 1992. Systematic implications of seed coat morphology in Lobelia 

S hi t ii ml i e i i obwowieae). < bu i b ii b » . l . ; Lot 15:155 I 72. 
1995 An i i v b iiH'u ii i it ti t ' < (Campanulaceae- 

l obelioidoao) with spin uil leicreru e lo seer I coat moiphokwv- I. Lac. Sci. Univ. Tokyo, 

Sect. 3, Bot. 15:349-371. 
Iurillo, A. 1889. Plantes mucin inales du ( hi!i. I /position I wveoello, Seccion Chilienne, 

Iavas B„ L.E 1979(1011 1 b u u i L ti i t I hi I Universidad de Chile, 

i ui i er, L. 1 874. NomencLitor I mtuiw us, vol. 2. b Fischer, Kassel. 

himppi, R.A. 1873. Di u n de la plan! mi v, ii poiada ultimamente en el 
herbario chileno.Anales Univ.Chile 43:479-583. 

1895 PI n r f , e ischkena \nah Urn Chile 90:1 87-191 . 

1. 'In I 'ill K , , | , , 

I - I l°O k h ti ]| I ,t| I i i I r i 1 In i ii i Id i -u I, 

I I 7:206-208, 45 1 -464, 481-482. 

!910.Florade Chile, vol. 5. Cervantes, Santiago. 

Santa Cruz, A. 1932. La trupa o tabaco del diablo. Revista Chilena Hist. Nat. 36:98-1 

Schultfs, R.E., 1 976. Hallucinogenic plants. Golden Press, New York. 

1981. Iconography of New World hallucinogens. Arnoldia 41:79-125. 

1 990.The virgin field in psychoactive plant research. Bol.Mus.Paraense 

Goeldi,Ser. Bot. 6:7-82. 
and AH i I ■ I In m i I h n n ti\ oih illu in u<ni I 

Thomas, Springfield MA. 
Sims, J. 1 8}0.Lobelia gigantea uigdhti i I Hi I r Mdg.32:pl. 1325. 

1825. Lobelia tupa. Mullein-leaved lobelia. Bot. Mag. 52:pl. 2550. 

Spooner, D.M.J.F. Stuessy, D.J. Crawford, and M. Si. va O. 1 987. Chromosome number 

III II it i ol tin linn f in m I I i (i I i I , i r 1 i 

(Campanulaceae).Amer. J. Bot. 83:1 356-1 364. 

S'i '■■■, B A 1987 ^sti-mntk , it I - Ju i i .t I^'Uopogon subgenus Centro f 
(Campanulaceae:l_obelioid( i I I I i ii 1 h i ]U ii i . i i I 

S TF and f M i Orih i ' I i f ith t honoring Bertero 

vascular flora of the Juan Fernandez Islands und continental Chile. Gayana, Bot. - 

Sweet, R. 1 827-29. Lobelia tupa M ilH u I ..til J'Hn PI 284. ln:The British flower g 

vol.3. James Ridgv* ay incl Sons I ond t 
Thomas, G.S.I 990. Perennial garden plants or the modern florilegium, ed. 3. Timber 

Thompson, S.W. and T.G.L e H'Q'PI ir 

Lobelia cardinalis complex (Campanulac 

Vvi..,W. 1874.NotuhH m< air S , I n 1 n n i in I i n usis L mnaea 38:69 

Vilmorin, R. de and M. Simonet. 1 927. Nombre des chromosomes dans les genres / 

linum. etchezguelquesautresespeces /ecntale not.-H:nd Hebd. Seance: 

I.Paul Parey,E 
'ergence. Ame 

;, A. 18G 

)4. Campan 

ulaceao. Pp. 


. In: A. Si( 


terei Beschr 

eibung Kultur 

und Verwendung, 


^ .'i j,M in, 

3n of the earth 


ringi oVet 


395. Floral cc 




Vimmi k, F.E. 1 14.ln:R.Mansfe 

(ed.),DasPf!anzenreich IV.276b.Akademie-Verlag, Berlin. 

1968 i aiiifinm kH' I "ilndi « ||" HMl " Lt Campanulacea 

Cyphioideae. Pp. i-x, 81 5-1 024. In: S. I.Xiner t, ed. PfLinzenreich IV.276c. Akademi 


Harold Robinson Hamilton Beltran 

[/ Mayor de San A 

Trepadonia was established as a m n >, h, i> , rn ihe broad concept of Vernonia 
(Jones 1980) by Robinson (1994) based on the one Peruvian species, Vernonia mexiae 
(Robinson 1981). Tin «i> mi I i - I i i ■ ni h ' it and the 90°-angie 

branching of the primary branches ofthe inflorescence.The species also has distinctive 

? inflot m ^ t I f i t ,aonia oppositifolia is 

) from [* in Inl tin i I i ii i | | n I i n miiotm mil 

d more florets in the heads. The new species occurs in southern 

Peru, and if distributions of cohabu mi < n ih i -imboo Guadua are indicative, the 

new species may be found eventual v in nearby western Brazil. 


9 m high; branch* mi , il t u i es opposite; petioles mostly 
, base dilated and reddish; lamina ovate, 11-13 cm long, 5-7 cm wide, 
largins entire to slightly undulate,apex acuminate, adaxial surface bright 
rib ■ il urhf> pal ip |P nl ni t 1 1 1 j i hans, venation pin- 

airs of widely spip if h i n it i itlu < ( -nee rather thyrsoid- 

i primary and secondary branching mostly spreading at 90°-angles, 
sse. Heads separate, mostly sessile, homogamous; involucre campanu- 
gh, 4-5 mm wide; involucral bracts ca. 28, gradate in 3-4 series; outer 
nm long, 1 mm wide, puberulous outside, brown at apex, inner bracts 


m long, 1 mm wide, glabrous. Florets 
brous, ca. 5 mm long, tube 2 mm long, throat ca. 1 mm Ion 
lanceolate,ca.2 mm I u 1 p . I i 2 mm long, 0.5 mm wide, 1 
appressed setulae; pappus bustles white, ca. 38, 4 mm long, 
series ca. 0.9-1.2 mm long, scabrid. Pollen ca.37 mm in dial 

ouiple, <ilu 

mi. t 1 1 i t lilt 
smbuci, sguamae oi outei 
it. i it II n t tu . I, m it 


'.i ippoMti' ami veilu illale < 


id the Afric. 

I. h !(: /'..' /. 'HI 

nown only in one Jamaican species of Lepu 

• spi - I- el Lt ' '7t"e,, |\,hlli .-ii 'U'lOi p- 

.hegeneia in which it occurs.Only in the An 


\.AIonsooftheSI/i II i i , n il Hoi the Biodiversity 

Wiiiiilmina Flimsilr Jasi-hmski. 1999. A Pompeian Herbal: Ancient and Modern Medici- 
nal Plants. (ISBN ' ~ i - ' 't ' I > I I F ■ "°l " A u tin I X 

7871 3-781 9, U.S.A. $ I /.9b, pbk., $35.00, hbk. 133 pp., 1 5 color and 7 b&w photos, 36 
b&w line drawings. 

Whilee^avatinqi'iin it I m mil i il It in i,' t in n ' ^al workmen collect- 
nq ,i \.)ti( t\ nt pkml t i in tli in I | i J r f < i , lit in ill iiiLiuontinucd to 

collect plants, I beq m t n I i Ml- [Put th - i I 'nil i m dk tne were the same 
ones the ancient Romans had used Ion urns." I Kit quest inn qn initiated mlo'l l'om;h'ion Herbal, J 
collection of 36 plants i mniniit " mp- m la 1 1 ' ■ ■ r --. f m, nedicmal uses with an- 

area.but it alsosei 

to be doing ar- 
chaeological excavation hit i 1 1 1 1 1 . j r i I mollis turn .I ' hoi,ini( ,ii infoimation liom local 

In tin inti idu tiou ' Huh i | I hi li Is I I in I ii - d I ' i in I <n i ipi>n>| i 

it I k [ |i ur i nl m i i i ii i i - I i I i i i id.- in lent literature of Pliny 

td i I I i I ul Id i l it m I ih i i in I i i ti i 1 metv of sources is what 

'I . , ii it in ii and ' ul ii | i i | ii in i ii I what ill 1 1 i I a li it - / oi lisvi pu 

I i li u P v i n ut in i ul ha i I it li i in i I Ih i ' I mi il in l ti i ds in i! i it ill i I 

II ui -ii IIh il it - i i I ill 'I ■ Pi i j i I Ii H I ' H ] i , I In , 

able archaeo-bot 

i i ii I a I I i [In i i I i 1 li t i i I t n it 

' i ink in pu u in 1 1 I | k 1 1 h, in in i i i n ui in I u ,ni but not of genus a 

I i nli . i I n dl i i P also t '. th- mil patterns 1 ,n i nt Pompeians- ti 

it Pompeii over the years have neglected ti 

and urn 

lii lm i I i |. i ih inti) m It tu t I n i I mi i pn | in I i i int. tin it il n o n i 

I ! M III I I I | I t | 

to appreciate. CompP : '■, , !ii iPI i lill itip ' in nt lit > it n unci notes on citat 

1 00 pages and less llian M.'O makes il a baiqam Ins ihuk-iv inadei inleiesled in aliernativo mod 

MP II I I I | II 1 I t| I It t i I I I i I It t II n I kit I 

lot ■ oik 1- ( oui i I ut nil.l |.| 1 i in i h j ink t I u i 1 i nm « ti isitt htiup )kl \\ 
paleoethnobotat . tmii ,mi 


Mark T. Strong M.S.Gonzalez-Elizondo 

Department of Botany, MRC/166 Herbaria CIIDIR, ( 1IDIR - IPN 

Snmhsonion Institution Apdo. Postal 738 

Durango, Dgo., 34000 
Washington, DC 20560-0166 USA MEXICO 

:i:\'n !:nsp:)t(i/<h-uGitipancnvi \A alannq ,-;'■,; i . ; i<\\ a ,■■.■ .■■.'..■ ■■ aa; h'< aiioraa a', ' Aonza.k ■/, from v,a: 

n it' 'I ii <it')> -nte similares. 

Rhynchospora Vahl nom on i 1 i i t n u ,p < les worldwide. It is 

most diverse in the Western Homi , pi im , mini a am tmnporate North America 
and theneotropics.EspejoSema m i L f ^t > i «int^ d 51 taxa (including 

f i > tlu Rma) from Mexico. 

Rhynchospora, a vn\ m im h . Ii n n mi ur pi a its that range from 

small diminutiveannuals less than •> inmllt I i it . i it a,, pmennials up to 3 m 
tall. The leaves are piinitiiil h i ii ^ i J 1 ill i imp m dorsiventrally com- 

pressed, and often have scabrous mamma. The inflorescence ranges from a single soli- 
pound panicles of mam spn < k i Am m| I I i terminal and a seriesof 1 -several 
lateral, remote oi conti'iuou mil 1 i | n i i I it an oi cymose partial 

panicles from the upper sheathing bracts. The flora! morphology is generally uniform 
as follows: spik i scales, with (1-)2- 

pt tm I I- t III ii n I i ,1 ll md j. I ii 

-2 scales are stamina: . nl\ mth an al > t 

1 st vie is R-bram hod ot . mtire;and achenes 

often tmnsvmmAy rugose or rjgu : ose,cach bcarima ,;t 

riangular-lanceolate or discoid base of the style. 

. Tig.H STATKZacualtip^n 

n i pi , iniitn I ] in |(ii mm I In! I r i in i i fi I II u I It' H IK H 

3869s -A KmaR'n isotvaao m i : am! M ' I Rose Am m barium amonyim, amae also amordeR ia 

Fig. 1 . Rhynclwsporazacualtipanensis M. Strong (drawn from the type). A. Habit. B. Sections of leaf blades, proximal end 
(left) to apex (right). C. Section of leaf at sheath orifice showing junction of sheath and blade. D. Detail of terminal 


r I Mi K 'res . i .flection notebook as pro 

However, recent searches made by c ura 

P.anta peienn's caesnOosa glalma rh'zonate b 

alti, 0.5-1. 7( 7) mm laii, obtuse mgon vei set; 

ebmibitam laminae 10 30 cm lonqae, 0.5 r 

i -2,pedunculatae, 

pedunculi ad 6 cm longi. Spicula< ' 1 mm I n I 1 1.6 mm latae, anguste ovoideae 

vel ovoideo-lanceol 5-7, ovatae vel ovato-ellipticae, 

mucronataevel bie iM m t it > m t i , ■ m ml i 5-p 6-3.1 mm longae, 

2- 2.7(-3) mm latar nn'in I i li i mm mi mlh ti l r si mm longis. Stylus 

Caespitose perei ?rect or ascending, 

20-60 cm tail, 0. 5-1 7i mmn id. bin I men n (o subterete, finely ribbed, often 
channelled along on- i 1 li t II ,i i ihous Leaves basal and lower cauline;sheaths 
eligulate,coarselyveinedabaxially,greentopal i ften i I lish biown proximally, 

glabrous, the innm b ,n I Nimnbmtv hi finch m h ' Mm n with concave ori- 

flattened) widely to narrowly v-shaped or crescentiform in cross section, sometimes 
subinvolute or tight ll n i ihnm nm i i e t ,e ly ribbed abaxially, 
n )tl i I t I i ] in en gl il i i tioisely cabrous or 

sometimes essentially smooth proximally. Inflorescence composed of a terminal and 1- 
2, small, lateral cymose-fasciculate partial pun I il lateral | licles on slender pe- 
duncles to 6 cm long, the lowest panicle remote from the two upper subcontiguous 
ones; peduncles flattened, finely ribbed,antrm i I it > i , nut nns, glabrous; bracts 
of the panicle branches setaceous,those subtending the peduncles, particularly the lower, 
leaf-like; spikelet pedicels subtrigonous or plano-convex in cross section, the margins 
often antrorselyscabrous;spikelets narrowly ovoid to ovoid-lanceoloid,3.5-4.2 mm long, 
1 .2-1 ,5(-1.6) mm wide, with 5-7 scales, 3(-4)-flowered, maturing 2 achenes, the scales 
spreading with maturing achenes;scales ovate to ovate-elliptic, indistinctly finely veined, 
glabrous, margins entire, the midcosta prominent, pale brown to brown, prolonged be- 
yond the acute to acuminate apex as a mucro or short awn; fertile scales (2.5-)2.6-3.1 
mm long, 2-2.7(-3) mm wide; sterile scales 2-3 at base of spikelet, smaller than fertile 

1.5-3 mm long,basifixed,thecae parallel, longitudinally dehiscent, apiculate atapex;style 
2-branched from below middle, ca. 2/3 length of style, the branches glabrous, minutely 
scaly;achenes biconvex, obovate or obpyriform,achene body 1.8-2.1 mm long (exclud- 
ing style base), 1 .2-1 .4 mm wide, light brown to yellowish brown, finely transversely ru- 

gulose;style base n m i iimulit i -I nun long ) i^nn ie pale brown, of- 

ten tur eh t at mf ■ , i li ' nit ' ely bar bed, reddish brown, often setose at very 

) equaling or exceed! 

ngthe length oftheachene bodyv 

j the tip of the stylet 


i—Rhynchosporo za< 

uattipancns i known only from 

southern extent of thi 

3 Sierra Madre Oriental mountain mi 

, N,98°39'00"W,ca.4.; 


type collection made- in thes 
at approximately 20° 43' 00"! 
It occurs in pine woods and sphagnum bogs at about 21 00 m. 

Rhynchospoai Zu '■ < t il i- mi ' npora. It keys to 

R. section Stenophylluc Kukentha in Kukonlhal's ( 1 mo) monumental worldwide treat- 
ment of the subfamily Rhynchosporideae s. i i o ei ,'. - included Gale's (1944) 
Rhynchospora section ' u I Mil i ib (in part), and series 

Rariflorae Gale (in pait ' - i t I. in st placed in R. sec- 

tion Glaucoe C. B. Clarke as circumscribed by Guaglianone (1979). The obovate or 
oopvnfomi, yollo bio n i hen- ith i npiiu inaip mat urfacc interrupted by 
transverse conugath i tin n i m hi h > i i tu of species in that 

section. However, the narrow (0.S > mm),stiff,ihickonod,otten v shaped or crescentiform 
leaf blades of R I 5 mm), flattened 

leaf blades typk al of p i in ti u . >' / >> in In li tinoui'hed 

at once by this featuie fiom closely teiated species such as R. brownii ssp. americana 
Ginglnnon< h; i h ' mini a r i I ih il i nil R.zacualtipanensis 

has longer bristles and I m II i him/ i « I /ens The bristles of 

R.zacualtipanensis typically equal or exceed tin m it! > ' tl - m one body with severa 
some tirmo equaling ihe tip ot tin- stvlo uase, while in the iatlm two species, there are 

of the style base Tb t l> t ,,, -i :>,<•> o , | jMll , t-1 mm in length 

while those of R hi it I \ rugosa range from 0.5-0.8 mm 

in length. In a treatment of Mosoamermm Rhvnchospora by Thomas (1992), R. 
zacualtipanensis falls out at couplet 79[b] (/?. rugosa) in the pari of the key that treats 
species of R. sectiot ml lotn unshed from the wide- 

ranging R. rugosa by tin m iiph >i . o >i tin it I I kK 1, | i< oously discussed) and 
further distinguishes 1 1 long) and smooth 

margins of the style lease (ollen anlroiselv s< enrow, proximally in R.rugosa). 

Eleocharis R. Brown is a widely distributed genus of ca. 200 species with 43 species 
recorded by Espejo Serna & Lope,' t-emni (imp/) liom Mexico, the taxonomy of the ge- 
nus is difficult in pat 1 1 hi u n it iii | I i il n uters represented, 

Eleocharis moorei Fig. 2 I XICO .Hidalgo Stail: 

Fig. 2. Ekocharis mooreiM. Strong & S. Gonzj 
summit of leaf sheath with acute, mucronal 
membranous orifice (left). C. Inflorescence (s 

e,9 35 flonferae;squam 

longis. Stylus Trifidus. Achae 

Perennial m i i m ' - mi Ii iMI i li i - k ca. (1.5-)2.5-3.5 

mm thick, coveted by i onspe Lioir,,ek)i u mt«\,M m;\Ul i I ike, da: k | u.i ph d Minus scalps. Culms 
caespitose, or sometime ' htm il i I M tsu n - • « 14 m un lall, (0.2-)0.4-1 mm 
wide,angularto terete, soih fattened vvlu^n < I tiiH:l,linclv mi lcaic,p,-ilc< jreen;sheaths mem- 
branous, purple, 01 tta.mii i ii I | irpl tn I t it i iteen to translucent 

0.1-0.2 mm long mu i th ml, i upmimt n-.m , nl 1 1. d upper sheaths with 
the ventral orifice r dm , I i I il uhth m 'i i il t uncate to somewhat 
convex.Spikelets ovoid to ovoid Hk . ok. id . - "mm lonp 2 . •'. mm wide, usually acute 

to siihuummit at i| i In I < 0.3 0.4 mm wide, 9 ic floweret:; scales polyst 

J 'o mm lout a i) mm \ I in btt i il u li ' |< nl I 

stramineous midvonmh, h m I. id pmp I Mil nil 'Mm mini ind apo^ 
narrowly hyaline, (at most 0. 1 0.2 mm wide); 1 -2 basal stabs shorter, 1-2.3 mm long, 1- 
1.8 mm wide, ovatf t mmo M i in i I th >ame color and ap- 

pearance as the culm, the sides put pie, I du Lot hv. ill no, .mo the margins and apex hya- 
line, 0.3-0.5 mm ' 1mm long, the 
connective shottlv | I i t I it | t I Mil I M t 14 1 b mm long (in- 
cluding the style-base), 0.75-1 mm wide, very obtusely triangular or biconvex with a 
co lula in tin live ,| t i h k t I im ill I Ot I | uloim e II t 
us; |d is; i oiown, line iv .irnl shallowly c el lulu i mi it mate, often i or 
i t -i ml i i nl ]u n ntu in m i i- I III 
doisallv sublluttened, 0.4 0.5 mm lono and .is wide <il base, sometimes slightly widei 

mcilesoltht i In i hinhm 1 i n i it o h in ndqe atjunction with 

the body oftheachene; bristles 4 7,;edoish brown or vellovvish, rottcarsely barbed 1/2 to 
2 1 On i I. ngth •- n ilin T ' oxcot dino tl > at heme 

Pakaiypl.MEXICO.Verao;;u/Ssas oiarrotera al sur ckO luayat ocolla, i / km del horde con Hidalgo,2100 
883 (F,US 2879357) [distributed as Ueod uu, uooOom i , R hi 

r—Eleocharismoorei is known from only two localitii t i it i , i 

ose proximity to c<]< h old, t m ilu nni,' imitotth u i i f 1 u ire Oriental moun- 

lin range in the states of Hidalgo and Veracruz.These lie at approximately 20° 33' 00"N, 
3° 29' 00"W to 20° 43' 00"N, 98° 39' 00"W, the Hidalgo locality near Zacualtipan being 
Dproximately 25-30 km noiib estwib > ■ . ■- r u locality neat I luavacocotla. E.moore/ 
ccurs in sphagnum hi i in i r I , d- to est at about 2100 m. 

The name of the ne \ m [ i t i II. til ml i [ t i i\ Moore, Jr. (1 917- 
980), world renowned Arecace i t | T i i i i a md -in lot of the LH. Bailey 
ortorium at Cornell University, Ithaca, New York from 1 960-1 980. 

' I e| i | t i t- , iio/mes of Svenson 

329, 1932, 1939, 195^i sub ( in " " , t • , . m in the subgenus Eleocharis. How- 
/er, its achene with oom I it> , ml I I t i b - tmguishes it from all 

:her species in the oh u[ G u ,i , m r< m m i nimntly costulate, but 

cross section the\ n tin i a i i I ^ "</ are very obtusely triangular 

earing a much shot u i and n in i I ilut * imthei mlobate nor decurrent, 

id they lack bristles. Furthermor tl h rh mi I + if 7 differ as well in 

?ing lighter-colored,and the e to subrounded 

apex while those of £ moorei are a< ute to subacuminate. I. nioorei differs from £ 

< i II i in in hi im I r | i i t i I t I iii i i i buret achenes with 
different tubercle an I m n il i i i t i I i I i , , I md coarser. From £ 

differs in the t i it l ,i I Inlpuipl md fibrous scales 

I t i nM Hi i I i im m ' i nle the spikelets 

biconvex or veiv ill it uselv liuricjuLu achene with cost u late angles; 

n ii I i I ul i tl lowest spikelet scale being 

:■ scales,; trie a o ] ! ' i , biconve.-. as obtusely tnoonous ,11 henes with 


Je would like to th 

lankDan Nicolson (U .) ml I 1 ml Goetghebeur (GENT) for reviewing 


monymousie ie t , 1, | t 1 1 1 m tl it i i pro od the manu- 


I) for collection and biographical information on H.E.Moore; curators 

t B, G, M, MICH, an 

d UC for seamh'u i tl i II t t I n H E Moore specimens; and 

all 1 , wi ,1 ]i| lie fo: 

the fine drawing of Rhynchospora zacualtipanensis. 


sp F joSERNA,A.andA 

,R. Lopez Ferrari. 1 997. Las Monocotiledbneas Mexicanas. Una Sinopsis 

Florfstica. 1 . Lista 

de Referencia, Parte V. Cyperaceae. Consejo Nacional de la Flora de 

Mexico, A.C., Un 

iversidad Autbnoma Metiopo t,n ,1 1 1 Nacional para el 


Jsode la Biodiversidad. Mexico, D.F. 

Uaii, s. I 1 ) I I hin in in it i I i h United States, and 

the West Indies. Rhodora 46:255-278. 
Gon/an/ Ei i/cMX), M.S. and I'M h i i 1 1 1°" A ^ la . mIm ilr >n . it .m 1 1 e\ tothesiipraspecific 

taxa in Eleocharis (Cyperaceae).Taxon 46:433-449. 
Guagiianone, R 19'" It '* '' ' , • ' iM'Uih K \|k laceae) y algunas 

._ ,|n i. ,lm<- I'M it in 1 ^2 M »1 
Kukeniiial, G. 1950 Voi i ii n i in in it > M i I h t I t ind h bcji hhtl 

Svenson, H.K. 1929 M »n it >phi tu in nth j- hi ' . ' I hodora 31:1 21-1 35. 
1932. Monographic studios in ihc cy tins ' icoci a Rhodora 34:193-203; 


1939 Mm ir if hi li Mill i in i t > i na 41 : 1-1 3; 43-77. 

1957. Eleocharis (Cyperaceae). In: North Amencan I lora 18:509-540. 

Thomas, W.W. 1992 n n f ,i il ' i , | t i in i u oamerica. Bnttonia 


nset, LA 70584, U.S.A. 

AM', 1 1 -: AC! 
■ iHkii Ir i ifi( ihi[ n ,« In, , hi ii i I ,i ' , ' , I lam hon)Comeaux.F 

( umcaux, v se asiunan a la srnr Ot t uimuiic: Muir-onA , () hipo noi tciKx < 

Thomas Volney Munson Munson (1843-19' 
on the indigenous species of North Americ 

of Vitis for nearly fifty years and <tn i h ia _■ -A, >ns he produced a comprehensive 

treatment of the cjenus mi hi hi In I A ■ - riH ( » > ' ,i 

Despite Munsons imnn n i h I i< nu In mi f i I ition of V. blancoi 

(Munson 1909)included two unr^L, In i r, , n A- I lnhutions. The plants from 
northern Mexico and extreme soui,' H nT ■ ■ :n mc] to series Cinerascentes Planchon 
and the more southcilvdi uiAiir- A n h h il A- I » M i t ) < n<> i >< i identales Munson. 

The objectives oft 
ofthe taxa involved; (2) to emend f Inn, i |. t npi ion of V. blancoi; (3) to thoroughly 

(Planchon) Comeaux,a currently unrecognized taxon from northern Mexico and extreme 

Luciana Blanco of u i i ' il .1 m f 1 1 I m I mi i I h n in M f 1 n>< ies Blanco discov- 

tih of I, ill >u 111 1 | it I 1 1 nt f "unison specimens ofthe plants. 

Munson (190^) pu' 1 l< mi hi. mi .1 ii|>lu>i II ilono ithi photo 

graph of a specimen I it I 1 It 1 1 1i1nl.n1 n I ill n 11 ison stated that in 
1895 he received v\ liii 1,1 | met from M truii If 1 I near Montemorelos, 
in the state of Nucvo Leon in northern Mexi 1 i < as placed in series 

Cinerascentes Planch t 1 ' ] m n 1 h lei t o 1. bited to V. coribaea DC 

h / ' 1' 1 I liiinP hi in . null i 1 1 it 1 1 1 U> that st ik 

and classification will mi t'n mil I 1 m> I ul< \ I > ' int< toioted I 7 blancoias syn- 
onymous undei V tn ' , I ut Im it, 1 r uk \ 1 ^ 1 1 e umiii . 1 it a distinct and related 
to series Labruscoideoe Planchon, which included V. labusca L. and other large-fruited 
(15 25 mm dia.) species -tandley (/I9X) 2o) also consideredl blancoias a synonym of 

Uiut. 1// / hi sene 1 ' toloen ,alet. 

Bailey (1934) mad i ul I u ml m I i i i t n I nit n ill one. t stilus 

of V.blancoi. He point I til el 1 h 1 I ^ I n ti 1 iP <> dandien, was based 

on two forms Plant > hi n ibmo I, , , n , lotvpe PH!) from New 

Mexico or Texas wei 1 t 1 1 h t ti d I -n n ' >hndieri, and plants 

referred to var me'. , huh lh h t i tin inn hi t been treated in any 

subsequent publmol i in I il I '< um t 111 t ih i mentose form might 

anolhet n mn 1 i - II ut I 'n i| I > hi i 1 I i th nb i nam' 

1 '[t I I S AND Ml I 1 RUMS 

veuled that the noithern and southern plants t 
"t, two unrelated taxa with sepaiate distributioi 
irphologically from the northern taxon (Table 1) 
i long), which are often obscured by arachnoi 

■\50. 9,:>iT v mnd oOp KPT;, Jalisco if 'am 
ve leaves witfi mlescent pubescences 
1 havim)nly white pubescence.Larpestii 

C/nerascentes, which have short si 

tipules (approximately 1-3 mm long). The fruit si;m m 

13 mm dia.) shows that it is not, 

a rtdarivc ol 1/ labrusca and other large-fruited (15-25 

mm dia.) species of series Labrust 

:o/c/eoe. Instead, the above characters and others, such 

as relatively early flowering time 

.slums lounti in uoss set tion,,md medium si/v iiuii 

■ipen'nq in midseason, suggest th 

^iin -.''itli-uii! ?d to as V. blancoi has clos- 

2St affinities with the western ser 


The northern Mexican vines 

considered by Munson as V. blancoi are, as suggested 

jy Bailey (1934), the same as Ben 

' i hi li i in ,i k Planchon as V. 

yerlandierivar.tomentosa Planchc 

m. In Munson's (1 909) description of V. blancoi most of 

such as small stipule mini II i | i nM i nl ' m I end nnlv in the northern 

grapes (Table 1),and are characteristic of series Onerascentes.lhe northern taxon differs 
from other meml e< t n I i nl t ' I rac lets including: to- 

mentose abaxial leaf surfaces, a general absence of short, straight trichomes on leaves 
and stems, entire leaf margins, U or lyre- shaped basal sinuses, and relatively short fruit 

and the absence of one oi iwo i harac icis in an individual duns not exclude it from the 

Vitis blancoi, as hi t \ h I I I i I M n I i I I mts from southern 

Mexico having large stipules (2 L mm long) ^<.\ telativetv lamed nut (6 13 mm dia.).This 
taxon initially name d hi m n i o te I i g« umen {Munson s.n. 

n ml II ti it I i I i i n inGray)Engeln 


).2 mn oi |) ti light, | ointed 

I i p U h nl t il hm i .h nl u I . i lillui inn > \ 

(including peduncles) S.S 20 < in lone; stems ancmlai in cmss section 3 

I. I V.cinerea 

3. Leaves pubescent on abaxial suifaccm hemes giaucescent V.cinerea var.cinerea 

; blancoi Munson emend. Comeau> merd orests 3 74 I 90;USDA Pom. Bull. 

TI '! Id. I 

»||, . I. it t I IH I ill i I I ill ' 'I In I I H 'I •' '' 

r rufescent arachnoid t 

. I. , I I voung 

Tinate, infrequs 

) lyre-shaped, late 
i long (typically 2 

Dm U-shaped, to V-shaped and 
) nearly entire, with teeth to 3 
/eins frequently extending be- 

pubescent (puberulent c 

rfaces,with the pubescence cot 
Hi i it pie traight, pointed trie 

...<-• 10..) 

) brown, especially a 

inc base, sometimes obscured by dense pub( 
caducous. Tendrils and inflorescences absent e 
furcate, to 18 cm long. Inflorescences 2.2-9.5 en 
der 2.3-3.5 cm long, sometimes a 

diameter, with a pleasant flavor w 

D^tiibation.- VVidely iHstnnmecl in tl 
evergreen oak forests (De Miranda 1989) c 
ond parallel, in the stales of ( olima,Gueneu 
PueblaandSan Lui Polo I p ill i unci 
tions (1,050-2,450 m). 

12 9.5 in I nig |)eduncles 1.9-4.; 
it or replaced by a tendril. Flowers 
m long. Fruit a berry, black, glaua 
fully ripe. Seeds brown, pyriform, : 

vy 55 to Guadalajara, 

i MEXICO. Colima: 2.6 km S of Jalisco stat 

un 1 986, Comeaux 4207 (BRI I !; 25.7 km N jet. Hvvy 55 to Manzanillo vi 
150m, 27 Jul 1991, Co< ^ L F,T) Guerrero: 17.1 kr 

o, 1,550 m, 18 Jul mQ2 ( • . all > km W jet. Hwy 95 a 

\992,Comeaux5190 >19 Hill 6 km t co via Hwy 23, 1,200 rr 
5RIT); Mpio.Chilpancim U) « 5 las kliavu a ka 1 al W da Chilpancingt 
EXU);Mpio.Cutzamala de Pin/on, 1 km al E de Ventarron, 1 7 Mar 197: 
' of Chilpancingo, 21 Oct 1 944, Sharp 441413 (MEXU). Jalisco: Nevad. 
37 m, 20 Nov 1 968, Boutin and Brandt 2378 (MEXU); Mpio. of Autlan d 

2,0 a 

eJilotlan 1 ,"()i , | i I > m H-n, | ,, ,\ 

ie las Coloradas, Mpio.dc Jilotlan, 1,720 m, 5 Apr 1 988, Mendoza 3730 (MEXU); Near Tecalitlan, 
)0 m,2 Aug ]985, Rodriquez and Rosa 167 (MEXU). Mexico: Valle de Bravo, 25 May 1 971, Boege 
5 (MEXU); 9.8 km N of Guerrero state line via Hwy 55, 1,550 m, 18 Jul 1992, Comeaux 5194 (BRIT); 
' km N of Guerrero sta 15 i.H , 5, 18 Jul 1 992, Comeau> > Ell N of Ixtapan on toll rd. 

roll f ID kuM| I t I ( ill i it in, , | hil | 11 );N of Ixtapan on toll 

rd.l.OkmNjct.rd toVild u m 1 i im Mull , v I I II i Nepantla, 2,000 m, 1 7 

May 1953, Matuda 28368 M X\J): Am,nc\ i inia 000 m, 1 1-12 Apr 1954, Matuda 30668 

(MEXU). Michoacan: Rim on I Apt I M i i < .« W (Ml XU); I a ( isi ada, Testarazo, cerca de 
Tacambaro2 100m> , i 1 i 1- ' I i I il ( . JeTzintzuntzan,2,2O0 

m,23Aprl979,Cab,7 1 | J i n 1 M 1 - s W / (MEXU);5 km Eof 

ZitacuaroviaHwy is 2 0i)0in MM M ( L , m< ?, 4-RIM 4 Hm I ot Zttacuaro via Hwy 15, 
2,000 m, 24 Jul 1991 , Cot ueaux 504/ (ERIE); 18 km F of /iMk uaro via Hwy 15, 2,000 m, 24 Jul 
1 99 1, Comeaux 5048 (BRI km b o ila lat nH ,1 ) i m 1 Jul 1991, Comeaux 5049, 

5050 (BRIT); 5.3 kn BRIT); 5.6 km W 

ofZitacuarovia Hwy 15,1 )S0m M Jul I "M ( <>m, n« >lH .0 1 >n /(liRIl ),7 7 kmWofZitacuaro 
via Hwy 15, 1950 m M lul 0' I I II 'u 4 n i u no via Hwy 15,1,850 m, 

24 Jul! 991, Corneous , nO Mil - Mi r nta uato via Hwy 15,2,000 m,24 Jul 1991, 

Comeaux 5058 (BRIT); 10 'km A of i lidalgo via I K y 15,2 1 SO m, 25 Jul 1991, Comeaux 5059, 5060 
(BRIT); 10.8 km W of Hidalgo la Hwy I ' ',150 in lul 1 ' M >meau> 061 (BRIT); 1 1.0 km W of 
Hidalgo via Hwy 15 Jul M 25.9 km W of Hidalgo via Hwy 1 5, 

2,300 m,25 Jul 1901 t ,,. , MM *, h t H 1 ilgo via Hwy 1 5,2,450 m,25 Jul 1991, 

Comeaux 5064 and 5065 (Mil I ); 61.1 km W of I lidhgo via I Iwy 1 5, 2,450 m, 25 Jul 1992, Comeaux 
5066and 5067 (BRIT ) 20 t k nl tH I mil 1 ivnn ,4n M' > m 25 Jul 1991, Comeaux 
5068 (BRIT); 31.7 km E jet. Hwy 15 and 126/43 neat Morelia, 2,300 m, 2S Jul 1991, Comeaux 5069 
(BRIT);31.0km E jet. Hwy IS and 126/43,2,300 rn, 2S Jul H)0| , Comeaux '.(I/O (BRIE); 13.1 km E jet. 

Moreha 2 200 m 26 Jul I I ^ FM HI n i Hv v 1 5 2,000 m,26 Jul 

1991, Comeaux 5071 illhl I I it H > i t |u nm V' > Zitacuaro, 2,410 m, 1 1 
Oct 1980 Contrera ' 4 is Mn t f t 1 "l Hi It 1 kh I In t 

27May 1979,Sofo/Vum.r MMMJ loh iM.I tua| n u I <S o Nunez 1445 (MEXU); 

MkmEdeCoalcomm i I i rn A n i i . I mundeo 6 km al NWde 

Tuxpan,carr.Mexko Mi tMi i I I n Y M i . o,mi i m Jo.e Purua, 

Jungapeo, 10 Mat P- 1 4 ' m I i r Mdera, 2,240 m, 29 Apr 1979,Zara/o 

206 (MEXU). Morelos: 1.1 km N jet. Hwy 95 and IMn. n I put hi i in 2 lul M0 

5033 and 5034 (BRIT); Base of mountain with temple at fnpo/tlan, i,SO0 in, 23 Jul 1991, Comeaux 
5035,5036,5037 50^ > ml , FFO i I im ll| zlhu 41 1 1- m, lul 

1 99 ], Comeaux 5040 (BRIJ fl t>, in 1 lm n itt i ,l_ nil I omeaux 504 / (BRIT) 
1.3 km E of square il f | 14 11 1 041 BRIT );Cuernavaca, 

Nov 1941,/W/rancf 1906 (Ml XI J >; Mountainside abovo CTicrnavaca, 3 Feb 1899, Pringle s.n. (MEXU); 
Sierra de Tepoxtlan ~> ^ i i ,, F n 1 | ti tl hi 1 eari Mextco-Cauutla, Cauutla 10 i\Mt i , ^n'1 -ismi. ,| i t , M ma I Apr 1970, Vazquez 

24/5 (MEXU);Barranca Santa Clara.N.O.d! Mratlipa Apr IS) I I uqut 00 (MEXU).Oaxaca: 19.5 
km N ofjet. Hwy MO an f 1 at Oaxac a, 1 )0 rn lul 199 180 (BRIT); 10 km NE 

Oaxaca caminoa Y^tlat M I . , iF< t i tM mi ta de Juarez, Ruta 175 

al kmalNdelxtlan ie n. ,t 1 |tl ' t 1 i , 1 , ,1 , 

Ttaxiaco-Putla, Dto de In n . I -i in MtM> " r m I s 4 km al W del Vado, 

caminoa San Sebi ti ,n , i , ,t , t | | (1 , m " (MEXU). Puebla: 

Zacatlan, Feb 1943 8r ,r ' ME' 1 1 M i.MJ 4 I nut intntdtiill ta Hwy 190 S of Atlixco, 
Comeaux 5172, 5171 5 T4 0, - m=M San Luis Potosi: In RioVerde,50 m Sof 

'Mini |. t H\\\ n > ,nd ii 1 one M nl I < , ,. MRU 


an growth tomentose to pubesoent,rarely with 
y terete to faintly angular; internodes 3-10 cm 
long; nodes faintly banded to ,■ itl ,i i i , 1 i tjtm | nl un tnipted at the nodes 
bya diaphragm 2-3 mm thick lenti I I i t it n i iipstomentose,with white,arach- 
noid trichomes, not envelope. I i it i - I - i es cordiform to long-cordiform, 

generally without lol) ot M n gi < m lobed, with lateral lobes acute,apex acute to 

shaped, to lyre-shaped and also V-shaped, lateral run 1 ute; margins mostly entire, 

pairs of prominent veins;lamina glabrous to puberulant on adaxial surfaces, tomentose 
to pubescent on abaxial surfaces, with the pubescence consisting of mostly arachnoid 
trichomes, sometimi-s iik ludin< i impl immIiIi i ii Mi- home 5 5-1 1.2 cm wide, 
6.2-12.8 cm long; petioles tomentose to puberulent, 1.4-4.1 cm long; stipules brown, 
0.5-1 mm wide, 1-3 mm long c 1 lu n I It Im i absent every third 

node, tendrils bifun ah nt mint im t 1 -nl u Ml t - id 1 1.0 cm long, pe- 

duncles 1.0-4.2 cm long, should-M ' ' ml m n Mimes absent or replaced by a 

tendril. Flowers of th- functional pisiiTi [ I i ii Hi mil I mm long. Fruit a berry, 

brown, pyriform, 3.5-5.5 mm long. 

Commonly found along streams am! othei moist sites in the semiarid scrub and 
grasslands (De Miranda 1989) east of the Sierra Madre Oriental along the Rio Grande 
from the Del Rio,TX, area south to the Tropic of Cancer (1 20-1 050 m. elev.) Individuals 
from the Del Rio area have chat h U t in huh Im I t < /, a var. helleri, prima- 

rily from the Edwards Plateau region of Texas, and the more southerly distributed var. 
tomentoso. These inn m lin it i I n hi man I i i u 

1EXICO. Coahuila: S edge of Morelos via 
km N ol Morelos via Flwy 57, 1 6 Jun 1 986, 

4038 (BRIT); S of Acuna at 3.0 km S jet. H v i km 1 > i< 4039 (BRIT); S of Acuna 

at3.5kmSjct.Hwy2and29,17 Jun 1 ' i 12 41 (BRIT);Sof Acuna at 23.0 km Sjct. 

Hwy 2 and 29, 1 7 Jun 1 986, Comeaux 4042, 4043 and 4044 (BRIT); 27.2 km S jet. Hwy 2 and 29 in S. 
JuandeSabma, 17 lun 1 II T hi i |ui h Hwy 53, 1 7 Jun 1986, 

Nupvo I eon: s I ildago via Hwy 34, 16 Jun 1986, 

Comeaux 4026,4027ai TFIh i i i IM i i - \ 16 Jun 1986, Comeaux 

4029(BRIT);21.1 kr imeaux 4030 (BRIT);22.1 kmW 

of Sabinas Hidalgo via II CUmunn, , , [|| -th u \ »5 km N jet. Hwy 85 

and Santiago 1 Mi in n\ S n » , ; ,] [if - v m,-v at 2 2 km N jct.Hwy 

Corneaux 4057,40^ m I [III I in i M it i i I 1 1 Jun 1986, Corneaux 

4060 (BRIT); 37.3 km S of Montemorelos via Hwy 85, 1 8 Jun 1 986, Corneaux 4061 (BRIT); 3.5 km N of 
Linares via Hwy 85, 1 It 1 < > - I III ill I in ipId \uHn\n 'i 

Sep l'VS.'.t wmooo /my i BRIT ); TD.5 km N ot Monteiimmlos via Ikvv 35, /n Sep IMm, uwowm 
4947 (BRIT); 1 9.4 kin N of Monteinnielos via i Iwy 35,29 Sep 1 987, Corneaux 4948 (BRIT); 24.2 km N of 
Linares via Hwy 85, 29 Sep 1987, Corneaux 4949 (BRIT |. flturbid ia Hwy 58,5 Oct 1990, 

Corneaux 4975 (BRIT); 2.4 km E flturbid. ia hi > I Oct 1 HI mcciux 4976 (BRIT);5.4 km E of Horsetail Fail mH ^-n,,' f.Ul'M .m, ."> -HHI 1 ),8 8 km Wof jct.rd. 

to Horsetail Fall and H« - i i m G, -1 . ,. 002 and 5005 (BRIT);5.6 km E of jet. of Casacade de Chipitin and id. to I lorsetail Falls (torn I Iwy 85, 850 m, 22 May 1991, Corneaux 
5004 (BRIT);9.4 km N of hidie Mo. met I nH 2 n 1 I lul 1 ^2 t omeaux5154 (BRIT); 

1 .0 km S of bridge at Genet al Teian via I Iwy 85, 7/5 m, I 1 Jul I 222, t. utneaux 5755 (BRIT); 1 3.0 km N 
of bridge at Linnares via Hwy 85, 275 m, 1 1 Jul 1 992, Corneaux 5156 (BRIT); 4.8 km NW of General 
Teran, Hacienda S Pi Tamauhpas lof Rio Corona near 

Hwy 85, 19 Jun l c )8n . - >< ml I I IT II mil tl i una via Hwy 85, 19 Jun 

of Tropic of Cancet nil I I n > MM I m N of Rio Purificacion via 

Hwy 85, 29 Sep 1 986, Corneaux 4950 (BRIT); 0.3 km S of Rio Punfi .mm , I Km . < m " 

Corneaux 4951 (BRIF); 10') km N ol Iropii of Cancer via I Iwy 180, 170 in, 5 May 1991, Corneaux 4980 

a S of Quemado via Hwy 2/7,7 75 m, 2 I May 1 986, Corneaux 


FIFI 1 01 N(. I 7 

..H.I 895-97. Wfoc 

eae. In: Robinson, B.L,Ed.Gra) 

lew York. Pp.429-432 

■ o nop;o 

a ! m ot Nut; 

_.H. 1934. The sp< 

?cies of grapes peculiar to r> 


I' I I h[M|i t li|IIO I. I t III. On I I |IM 111. U. lit i 

I r 1 1 f ti i . i - ' i it m | v | ,, ' ' i || 

i Mi - I ' . . ' 'Mm i ill i | mi i.j. I i I | ul iii ii in. t I ilium il 

S.A., Mexico, D.F. 

I i Mm | i ih ih iti. mi in. I I. in ill. I tut ilh ii ii in I 

lutwM, I.V. 1890a. Ac luvol'i. at ion of Anion, on propesw ,nrd. I oiesls. 2.47 A 4/5. 
NONA Pom. Bull. 3:1 14. 

iunson, IV. 1896. Series VIII- Onerascent 

es.Rev.Vitic. 6:421 -425. 

iunsonJ.V. 1909. Foundations of Amerk 



anchon, J.E. 1887. Monographie c 

Jes Ampelidees Vrais. In: DC f 


Phanaerogamarum. 5:305-368. 

nfro, R.E. 1 983. The Thomas Volney M 

unson Memorial Vineyard. Graysc 

>n County Col- 

lege, DenisonJX. 

andley, P.C. 1 920-26.Trees and shrubs 

of Mexico. Contr. U.S. Natl. Herb. 2: 

[in Bi P K >2.Ethnobiological Classification: Principle 

and Animals in Traditional Societies. (ISBN 691 09469 1 

emityPros 11 .iIImm t ihu t nfK v Jersey 08540, U 
Mil M-' iltawings. 

As more and mok Till lit - i 1 1 I i I i I- [ mm m I in ill mm Hi and ethnobiology 

becomes a bona fidt pi j im ird to irmag - m not being an es- 

sentia text. It is hm i - I tin I I md tin In t | nf I th it I I'- i i l ■< i ,1 t i m no 

ning millennium. It is fitting thai or 

m of the nation's leadmoetln 

major [tarts, "Plan" and "Process, 

"Pc-dm dismisses the foecm 

?s involved in the functioning and f 

..".■illation of etlinoPioPmf a ■ 

ueneial, I nti ill P i i I i i m i it i n i I mill ihn mtli i ) 

with >t n n Bit i'ill i it I t i Pawn t th It sixties 1 ,h .it I he continued 1 publish 

to chime in on the i ut I mmn lunn . ii Pi n i i, r,i nam. i tew either agreed, 

disagreed, or agreed t I n ithlmiu J l t n il t I i u< it deal to the overall 

Inowlediii nfmPnot Ii n il i i i in hi i iii.lih. i Ii mi hu ut h topics become thor- 
oughly researched n i t nil h i t l it th ubiect is revealed and 
worthy of inclusion in a college progiam. 

Part one is divided into loan . hapten, drmt r, ,no im m.i ot a e ompaiativo kthnohiomavf 
"the Primacy of Gem mo laxa in I tin I i ' i '. lasb'k at mo,"" Ui< Naluie of Specific la -a and 
"Natural and Not llmPi-P'tdm aiefmum, Pmliutacr 

ti iti in d' I it it il i i I i lit i i | i 1 'i i I ii i i i I il i | 

tint turn iii mil mum i th i | m \ too lit ,, ,mn ml t miimu ituml pi 
i i i I t I m. i ii mm lo name a t I ,it two i m ! ; i| ' i 
n i llmolnolouii al Knnwli'doe."" Im Nonan ntiaiinesx oi i tlmmPmli igu a Non 
ailot in i it II Ii n ' I d I I n .I .a i ii In i Pt" tl t- I 

Ih m in I i b I m it hi n il i m I i ik I Put i k lurthm 

lengos those amhropotogbts who see wmxiy as a "set of culturally construe te 

ti i i in ifn p m< t t te i I ,i ii i i i f Pi I i il i am tam in. ; i i I in i i \ i i I i u 
scientiti. ititf mho I I , i , i t il ul |e t in fe 

I ronia siudent's mem sf k m. tivetlm Pest ihirm i < an say about the boof 

the il saw and taught me an rnvrntia lesson m mseatx lythat oftm' the n 


; monograph c 

l ' I hi l "It: 

Santhosh Nampy 

Department of Botany 
St. Joseph's College 

-> ' I -673 008, INDIA 


oecies of fern from Kerala, Indi 

a, is described. 

st> .- -i7' 'H 7 n N |,,dia for 


R[ a, Ah FN 


echo de Kerala, India. 



(1 977) recognized 44 specie 
escribed, including two fro 
1 993). During recent florist 

■ I 1 ill ms in the Western Ghats of Ke 

plani iirovving on exposed wet bouldtt n I - I ii " n i n m I- n 

Since this material does not ana > itl ' \\\ imns of any of the known s 

t i i u [i in I It , I i ii ni I ii i 'i i nl i "iti i 

i ni'l !f I >l' nihil i III i i A h iin in, ml , 

,10-13 cm tall. Rhizome 1.5-2 mm thick, short -creeping, sclerenchyma strands 2- 

peringtoagland iht i| I t i ,n II I I i h i m nn pale, glandular. 

I usti lid imotphii i ititi.t il i'i | in I < i n i t 

? fronds 6-13 cm long, 2.2-3 cm broad, pinnae 4 6 pairs; terminal pinna conform 
he lateral pinnae Stifu 1 I'mn tl in < i 1 mm wide.Pinnae sub-opposite to 

wrowly cuneate, herbaceous, apex hunt to acute, margin shallowly lobed with 
setae; lacking foliai bulbiK m i tomosmg to form a row of costal areoles, 
costal free veins present. Fertile fronds to 10.5 cm long, pinnae acrostichoid, a 7 
3ng. Sporangium 261 x 226|im.Annulus 1 3 ot 1-4 celled. Spores monolete, bilateral, 

B. Venation; C. Sporangium; D. Rhizome Scale. 

Fig. 2. SEM photograph of the spore of Bolbitis i 

Etymology.— The species : em reality Thomtr 

fall at an elevation of 400 m in wet evergreen forests of Kerala. 

'ui • vn< hob ' it — r i t e y ft i he I lili i it 
Kerala, usually found growing on hut 'I I- i m I t u I in n m 
■ hie ,nj|,|.. unien,-ar,in.'d INDIA. KERALA. Idukki Dt: 1 hommankuthu, 
75 (CALI). Kozhikode Dt.: VellarimaLi, 1 i Auu I'M 1 ) 1 '. Nampy S 74 (CALI). I 
1 I 'nu i j_| i i 'II |i- , n n (CALI). 

Notes— Bolbitis thommankuthlana is closely allied to B. semicordata but can readily t 

distinguished by Is stria Ict s'ze, t :; eel- v. in n : n I ■'■-., the wnqs extend in.; to tie r ; 
zomeand complete absence of foliar bulbils. In li semicordata the plants are 15-70c 


of Scietu i ,n I I In n.i' 

I tor 

ce in SEM photography; P. Bas, HP. Nooteboom (Rijksnetb.ii iun 
s College, Kozhikod' + | t i if lit s in! A.Radcliffe-Smitf 
sand A.K.Pradei p t ,t >\ oh hi for various help. 

Hennipman, E.I 977. A monograph fib in'' 1 I i mi lopsidaceae). Leiden, Pnsv 
M 'tin \V.S. n i, 1 997. Pteridophytit flora of the Western Ghats-South It 

Bishen Singh & Mahendrapal Singh, New Delhi. 

New Delhi. 

i 'i I I DA 
.a Ai /Vi i \B i and I lib i 1' hi Applied Population Ecology. 

' PlumtrecRoad, 

in .1. 1 1 mil i\1 \i)i II i/.' ' < ) .41 95.280 pages/64 illustra- 

tions, text with CD-ROM). 

i, ,4 li- i ,h- i i hi ii i'il i 't ■' t m t ii ' it nlu it' in. I iii hi i, .' il Ihh it n ifr (I 
it in in.|t im-ait . i ii iiuiiii . ii' i urn It ii t wit: mo I LcoLab < <i 

et || to I. i I | 1 III III IH I II I | it I III it n I | il | i «- tl ut 

h- hook is ii. .Ii 1 1 nt i S ; Ii 1- i in h .in , A' | j in i .tl mati n I i ul 

tion Regulation.4) Age Structure, !:,) Stage Sir ucti 

ii" A) Motanopulations and Spatial 1 


Population Viability Analyst 

s, and 8) Decision-Mi 

iking and Natural Resource Manage 

ment. E 

chapter includes the introdi 

in i jun 1 he exercises, u 

;ing the RAMAS Ecolal 

j, provide students with the opportur 


manipulation and analysis^ 

,nd will be useful whei 

iitant with its CD-ROIV 

1, provides a poweilu pa. kage that w 

tate mastery of the cc 

niise maleiial.but that will also serve 

work upon which students 

can formulate their o 

wn hypoethical models and testthei 

ti using 

with the chaptei en rnetapopulation 

precisely the most commor 

i phenomenon botan 

sts face when designing a\v\ Aghtinc 

1 Integra 

Conservation and Develop 

1 ii o,n lies loronsoivationol nfiaspecitic genetic 

/here we fully expect 

m fi'-ai mat ad.aiSena eleme ,1 t.Othe i tha^ t 

Ate he comple'asAcailv presented and eeilainlv a haiuai 

tware provided on th 

e CD ROM 1 unhesitati-gly recomrr,. 

who must teach a populatic 

,n biology course. -A 

"■■ i . , , ' '■•, .,,",,,,, 1, 1, ., • 



Alexander Krings 1 

ation for a taxon tt 1 i uill,|li I arc ostemma R. Br. (Correll & Jor 


artesz 1 994) is proposed. Recent cladistic analyses by Liede (1 996) failed t< 


^ monophyly of Sarcostemma sensu Holm (1 950), instead showing suppi 

Drt for 

aophyly ofgeneiu i mi im til M 1,1 Mi m M14) Based on her an. 


996) suggested, among othei n n riptions, lhat Sarcostemma be rest 


ampanulate Old \AMi Mi ■ M , t r , p r ll|lh p , M j jff , 

panulateNewWond I i h m i n j , lemma cynanchoides 

'artwegii (Vail) Shinm i i m m ill I- scribed as Sarcostemma lineare 
40. However, the e-i M t i i i I i ,n Sarcostemma lineare 

^, (based on a Venezuelan ,11. nnn t- ym> i the u- e of another name for 
taxon. Although application, Holm 

that the name Sail i' ,, ' , Tu h, iihb toi th< ul 

a of Sarcostemma c - - . , h t f ,- , ' New Mexico, USA) 

spurn Benth. Holm (1 950) decided to employ the epithet "hartwegii" 

a new name 

ssp. hartwegii iVail I 1 YhMIr mm I mbmation: 

Funastrum cynanchoides (Decne.)Schltr.var. hartwegii (Vail) Krings,comb.nov./ 3 /i/7/fcierfe//o 

'* • ul I HI T i [i ul ^ 1 'i iPM I ' M euM/7, 1839 (NY). 

- ■ n Mi t ., . ,il > I, In -m i | (, Mm | l p 4 ,/ oste, u \a 

vnci'K lu}idei\\x'ne.\.;\!.!nir!'.\r,i:>i\/,)''< ' ■■'■ -m.M, ; '■ 'M. I'M !M L ;-, , ■■■.'rnvn.ii ynanch<>it1es 

sandy,or gravelly soil in Trans Per < \ ' i- f 1- . , Mizona,and Utah, as well as the 

Mexican states of Baj.i J Hit im t In i , n I ,u u oahuila,Durango, 

Guanajuato, Jalisco, Querela ro, Sinaloa, Soiioimmik I Zacatecas. 


)rrell, D.S and M.C.Johnston. 1979. Manual of the n 

/ascular plar 

its of lexas. Uni 

versity of 

l< >d ,it L ' ilia m I it , i 

\Rif sz,J. 1 994. A synonymized checklist of the vasci 

ilarl ira of tl 

ic United State' 

;, Canada, 

and Greenland. Timber Press, Portland. 

dlm, R.W. 1950. The American species of Sarcos 

temma IT Bi 

■. (Asclepiadaceae). Ann. 

Missouri Bot.Gard. 37:477-560. 

de, S. 1 996. Sarcostemma (Asclepiadaceae)-a cc 


generic circum 


reconsidered: Morphological evidence. Syst. Bot. 


:hlechifr,R. ]9]4. Philibertia Kth.und Funas 

tmm rourn. 

Repert. Spec. N 

ov. Room 

Veg. 13:279-287. 


Bruce A.Sorrie 

f , - , . / , ' ,- ■ , ; : 

11' /." , -L f'i-1 


Based or ar,iiy. 
iiuimIh',' of mensural. imiqu, ml, i 
:-"l.7/.^-- , ( smaBasoo or the rooif'vi 

■ ease of finding ft/eprocarpa in suit 

Ml eUMI N 

r- :josi? a analis's mnrio : 6amn 

.■spec in diforente. Antique clatamet 

Rhynchosporaleptocarpa{Cha\nM * inn n ill i lit I < .nun t d as a distinct entity 
by A.W.Chapman lun itit i mil it M -I I i J mil m mihor of Flora of the 
Southern United States (Chapman 1860, 188 >, 1, ' r > unou i, c hnpman did not pub- 
lish the name, although he had prepared a manuscript (fide Britton 1 892). In the three 
editions of his Flora, Chapman does not mention this entity.ln 1892 Britton published it 
as R.glomerota (L.)Vahl var. leptocarpo Chapm. and ascribed to it a mnge of South Caro- 
lina, Florida, and Alabama. Later, Blake (1918) um i r Itim , no tangle by split- 
ting off the generally more northern R.capitellata (Michx.) Vahl from the more southern 
R. glomemta. Blake treated the entity as R. capitellata var. leptocarpo (Chapm. ex Britt.) 
Blake and provided a brief Latin diagnosis. Blake gave its range as Virginia to Florida and 
Mississippi. In the second edition of his Flora, Small (1933) raised it to full species status 
(without explanataion or discussion) and provicf I tin mil >m| f te English descrip- 
tion available. Small gave its range as South Carolina to Florida and Mississippi. In her 
monograph of the section Eurhynchospora, Gale (1944) reduced it not to varietal status, 
but to complete synonymy under R. capitellata. Although she treated two other botani- 
cal forms of R.capitellata, Gale omitted any discussion oftaxon leptocarpa, possibly due 

notably ClewelK 

1 985), Godfrey and Wooten ( 

1 979), Jc 

, Radford etal.O 

968),andWunderlin (1998), 

do not 

.None ofthe sta- 

te Natural Herilacie hnqtum 

s within 

bler lists it as i 

cept foi North Caroli 


ognlze R.leptoca i ;h i ,:\\ anv lev 

tangi ot R.k nh> aipa ii coqn 
references to Rh\" • < :' ' < nil" , 1 t lot - itui at those of Anderson 

(1995) and Some iM il * ' ' Mil i II I I i in n in tin Florida panhandle 
andtreated/e/ it n. n > k > i j u , i tlutiA , apitellata complex 

needed furthei I that R. leptocarpa 

may wananl renewed status, 

Myattenlion i ti t I mi |i it n il | u ml hiImIiiN Ium ollected plants 
in 1992 on Fort Ri 1 , i km > i it uM tl n | nt if a lare species sur- 

vey.Although thes« pi if I II ml im ' lit ^ n I i i n '> n) jnd Gale (1944), 
her plants did not do - I, i< >m,ibk pkmi . I An ttom New England as R.capitellata. 

■VIA IF Sl/V '-■ A"A i iVl'-'i I IA' v, 
Specimens of R/n " i H i I m ih I Id dutinq the course of 

botanical survey v . n il u I I i n I N tl uokna Iheo 1 1 iw 

A- -n ufpkment i | i i is it hi it h ' it ill I ma DUKE, FLAS,FSU, 
GA, GH, IBE, NCSA, NC 1 1, SVVM , LIS, USA, VI )U. From ac toss the tange of each species, five 

lowing achene ( le i t i I i ' i ' i I i I I t I t qth n In ith I | w 

anih bristles relative to tubercle (e.g., the length that tin 1 bristles exceed the tip of the 
tubercle). Measurements were also made olth. i 111 * |i I i i ters (n = 50). Counts 
were made of the number of spikeks clusters nereulm and tlm- number of glomerules 
pet nlloreseence node (n = 50). 

Notes on habitat were made at ovet SO sues in trie fieid, supplemented by her- 
barium label data K,n iii i ' | it It in i ii il I In in ji atla r es,herbarium 

Herbarium actot \ m t II It t hi t I i | i nil il mum 

Distinguishing characters 

M< n ut k 1 1 it i m t t is i it i 1 1 I i t it i u i i mi i nospora leptocarpa 

differs ttom R.capiieiiairi n ,t east tour it i| it mt ways. 

hero! I- < ' >r> net ■ Rhvnd > <• p < sses 5 H In 

.si oAoemaA" nA.Ao. I acu spike or , lustei A composed of i S neaddiko glomet 

assume a turbinate shape, rather than the broadly ovoid to hemispherical shape of R. 

3) Achene body anrft<^< nV . > / Mitv >ugh the combined length of body and 
tubercle is similar in the two speu< tf .- f t n utter Body length of R.leptocarpa 
averages lone j. m than , ,,; ,",, i bui ; Ut f |, n , ith averages shorter. 

4) Bristle length— A striking remit _ . ■/ o> ;, , ,-,■; ( that the bristles always exceed 
the tubercle (in rare instances nn Ttl ' bristle may be egual or slightly shorter). 
Moreover, the longest bristle um i >\ , . I i .. 'ubercle by at least 0.5 mm. In R. 
capitellata, bristles normally are slightly shorter than the tubercle, but the longest bristle 

There are qualitative e haia let In. h i i < , I iinou I , the two species. 

The first three ate t i 1 mil i tM it I ' , n it . M I ntification. 

1)— Plants of/?, leptocarpa found' r < n h in hi. h >a li it. _> many lazy culms 

pendent of flooding or fire event q I I mi , i ^ ' iM although more- 

strongly ascending. Anderson (1 995) reports 30-40 culms for R.leptocarpa in Florida, and 

is the norm. Herbarium specimens almost invariably consist of a lesser number of culms, 
because whole plants are rarely collected. 

2)— Foliage and culms of R.leptocarpa are light green, often glaucescent;those of/?. 
/ f ^ .' ; it.- medium to dark green. These differences are retained in dried specimens 
unri sunn to he independent of light levels. For exampr , , , , mowing in clear- 

ings created by logging or military activity show the same pale coloration as those grow- 
ing in shade. Similarly, R.capitellata retains the same dark coloration in dense alder thick- 

3) — Spikelet clu t i t < ' n | il hi hit ' * n d ni 

A) -Numbei >t il>m ink | t pil i iust< in < jlomerules occur 

singly or in pans it it t t I i i tl ill n mles are paired or 

tripled-there clearly are 2-3"heads';arising from separate or branched stalks. Combined 
with greater gloim m i tliii ml il n I > t ty much broader 

5) Mature acholic color is pale bmwn with a pain i cntial bulge (umbo) in R. 
leptocarpa; dark brown with a more contrasting pale bulge in R.capitellata. 

6)— The base of the achene narrows into a shorter stipe in R.lep to carpa than in/?. 

: uw'e/T'uWis depicted bv IGike ( IhiW 


The widespread Mn , , t . uf u d mad rangeof moistto wet,mostly 

open habitats, from n mm if e tin n mek banks, pond and lake shores, 
beaver ponds, peat h n 1 1 > i I i I lit I I i 'n i itpo is very limited in 

its habitat prefeieiin o! o,w 10 ilm e. n ,phagnous streamhead commu- 

nities within longleaf pine < eceive water from seepage 

which discharges from adjac ent uplands and form the head ends and upper portions of 
drainage creeks. In tie h in i ill n t« imed"streamhead pocosins/'dueto theden- 
sity of shrubs in them, larther south and on the Gulf ConstalGaiiethey are called"baygalls," 
due to the presence of "bay" species: sweet b i << , to L; red bay, Persea 

po/ustn's (Raf.)Sarg.; loblolly bay, Gordonia lasiantl is 1 Flic Nh n embedded in a mesic 
ravine, as occurs frequently on the Gulf Coastal Plain, the baygal community is called a 
"steephead." All of tin < m gn G Ion t I but tin narrowness of such 
streamheads normally allows for fill t mkjht to t in horn the sides. There, R. 
leptocarpa occurs in the semi-shade of trees and tall shrubs.ancl especially in small open- 
ings created by blovwli 1 1 tin iiit i i ti i i lieut li tuil it c I downstream, where 
flooding becomes too frequent or of long duration '■ e; to arpa is absent. Table 2 lists 

, i i ' i i Gjyer/eeworo / ' W , i u u i iGt \ h in u inn t 
beaver ponds and hum it mi unli t | nil I i ti ibntnies enter.One speci- 
men from southern Got nl ' , .unr eel mum, i Ic\ t. d tiom a 'pitcher plant bog," 

presumably from tli G h u I ' i ill n G it t ti amheads. Ander- 

son (1995) repot t ^ ' li null i[ i i lo I iti mIui it h in panhandle Florida; 

it is the only record from a near-maritime habitat. 


Missouri, and Arkansas. Howevent ihu: tl I m mi- miti^ an i ind ib >\ i li m 
the rest of the southern Atlantic and Las; Gulf Coastal Plains. 

In southern Aikans / t i I 'ml mm u Smith 1 988), but I 

question whether all spcimen-, jrt - m tl hi nttn I in - thm< .pecimens have re- 
cently been annotated to R. glomerata: Bradley Co., Demaree 24539 (NCU); Hempstead 
Co., Demaree 54102 (NCU); Pulaski Co., Merrill 183 (GH). 

In Louisiana, Thomas and Allmi 1 1" < m n a '< /' , ir ii e parishes, but two 

cited specimens are actuall nas 72521 (NLU); 

Rapides Parish, Thomas 40834 (Nl U). In addition, specimens from Grant, Morehouse, 
Ouachita, Rapides, Sabine, and Vernon parishes (FSU, NCU, NLU) prove not to be R. 
capitellata. /\(Jn\or\ Parish specimen is equivocal thea< henevi/c is closer to R.glomerata, 
but shape is closer to R.capiteHa. < Ml II i - t - pecimens from Red 

River and Winn parishes that were cited by Thomas and Allen. 

Two east Texas spteumm mi I Until i ' ' > r pora capitellata: 

Henderson Co., Correll 26688 (GH); Smith Co., Moore, Jr. 888 (GH). Both were collected in 
hillside seepage boqs.A Leon County collection, also from a bog, proves to be R.glomerata 
vaf.angusta Gale, an overlooked taxon of the WeM uuii ( oastal Plain:6or/c/ey 13505 (GH,US). 

Rhynchospou icr' i m n I i i i I nti indLi ' 

Coastal Plains (Fig. 1).lt occurs from the vicinity of New Orleans, Louisiana, eastward to 
Tallahassee, Florida and adjacent Georgia, then reappears in the Sandhills region of the 

i ii almas i uuent I i I I i i i i I i i ii mi the outer por 

tion of the Atlantic Coastal ICiuc [his distribution nathan a Gull Coastal Plain compo- 

cies(5orrieetaMQ^ m i t tn hit. M >< - - - , imooc? (C.B.Clarke) Small, which 

mi ha I ut open oo| 1 i< 1 01 uliacent (o tn mihe ids and baygalls. 

tlr lit | lp I 1 -I ,-f II ll 'l III P 111 1 I 

i I i glaciation cycles forced the ancestral entity 
tes) deep onto the coastal plain. Once 
there, this entity survived in open wet haL'it i mil < i. i i > Ik i farther north, and 
also by adapting to localized cool-water environments in shady streamheads. Retreat of 
glaciers allowed the ancestral emit , r i inn i nth. .aid, while the newly adapted en- 
tity of southern stu imh i I i. n hi I n il il I mi have been involved 
to complete speciatkJii -It. i iivj tn ,,i. n J tnity may have occupied a southern 
range;during glacial * I | tun mI I hi m . hi Opt I to a broad suite of 
cool-water habitats as they migrated northward. 

KiMjrcv-nntiv: ■.peamens.l he Ollov'/", i ;v- ■ ■:'■■•■ "■■■..iOv ,| >eci:-ii "'s from tOouohout :■■■■■■■;■,■■ i 
of Rhynchospora lephx uipa nuiinJu.l 1I1 t .I II nt> ALABAMA: Baldwin Co.:freguent 
m wet wooded stream ith \\ mm i i !• i Hinge, 26 Jun 1993, L.C. 

Anderson 14393 (FSU). Geneva Co.: ,".' nut -^ . m h Sa n ,un sphagnous headwaters of 
tributary of Pea River,3 Aug 1996.AA Some 8960 (bas,NCU).Mobile Co.: pine woods nearthe coast, 
Jul 1879, CMohrsn (U 1 i i | I <[l it! i I i h I 2 Nov 1980, M Powers 

s.n (USA). FLORIDA:Gadsden Co.: Cum.; u , , (GH). Liberty Co.: hammock on road 
12 near Bristol, 22 Jul 1940, Western* \ > < Fl Santa Rosa Co.: 5 mi. N of Milton, local in 
dense sweet bay-C//fnn h mm t t ^ il 1 .7 t O Fi I jlin Air Force Base, bay 

swamp near head of steephead ravin. \\'< i . t u. O " . p m^n i upland Bridges /2567(NCU). 

55250 (GA,GH); swamp near DeFuniak Springs, 6 Jul 1897,/\HG;n 'n,i ^ I Hh^niin 
US) Washington Co.: i i i - ■ \ t i , n II i I | . nn pond, 1 2 Sep 1 989, 

5. Orzell and E. Bridges 1 1883 (NCU). GEORGIA: Decatur Co.: wet woods near Whigham, 10 Aug 
1901, R.M.Harper 1185 (US). Liberty Co.: - >■ i m ,n CO ! ally common in full 

shade, stems lax, 9 Jul 1 ' < < r Thomas Co.: low wet woods along 

small stream, nearThoinasMlh 14^ r uDUKE). Worth Co.: densely shaded 

sphagnum pocket in s.nnp .1 . t • < • - Hi. LOUISIANA: Orleans Parish: New 

Orleans, Ingalls S' MISSISSIPP Simps 

Tracy8616 (GH, US). Stone Co.: Univei it t ,' ' -h i I >i i if Wiie Road and E of route 

1 5, mossy headwater ot Link I hi il in i 1 1 NCU). NORTH CARO- 

LINA: Hoke Co.: Foil I t i t i M ' h i In i i i [ ul' in In inous mucky soil of 

streamhead pocosin,25 Oct 1991, B.A '» <rr ■ ■ H bas, "'. ; rt Bragg n troop trail through branch 
ofNicholson Creek gio him pi tri tin m | i ,i un | " I .orr/e 7002 (bas, NCU). 

Moore Co.: Sandhills ( j urn m F t h O 1 i I II- I through tributary of 

Deep Creek,4 Sep Ow - J I Richmond Co.: sandhills Game Fand.WofSR 

1003,tributaryofNahdi h - I h. i- rn I I, n, I ,r i- f „ in n Sep 1997, B.A. Some 
9490 (bas, FSU,GA,GH, NCU, VDB).Scotland Co.: Ca i| I I .11 U \rm i, boggy depression bor- 
deringfloodplainofFt.ownin.ii ml mo 1-2 ■ >. OJ, it,. Hulls Game Land, head- 

ary 1994,1 7 Au<i I •' - bOUI M CAROLINA: Aiken Co.: Aiken, 

Batsc r.lWBtc , Chesterfield Co I nils Refuge, 4.2 mi,S 

of jet. of SC 109 on SC 145,76 Jul 1 99-1, R. Krai 83865 (VDB). Darlington Co.: Hartsville, flats across 
from paper mill, 5 Jul 1909, 147 CCokers.n. (NCU). 

I , itt t i I it i in r t t ti I t ii t i - I i ' ' h isporaleptocarpa 
is not rare but mcml mhnl. I In L hi -u m r« •,. i! , ,|h* tmns, I have encoun- 
tered it at 40 sites from Notth i atoltnu m iui ,,t|pi j p >ia leptocarpa is not 
difficult to find in | t [ i Ii t t i I n( il i i i 111 i h.- hi me A/hich is widely 
distributed over mu I I tl it I f I 1111 I tl - ru imh ads are usually very 
shrubby and tangled in >■ i ilm inn nit i I I mi t h In pi iickKvLk n 
it is most easily detected- July to mid September e our' ol formidable heat and hu 
midity.Addinsect^ i m i i - i onous plants [Toxicodendron vernix (L.) Kuntzc), 
and resemblance lo > ml ih i ' ' - i nil • , , vtphala), and there 

Lindera subcoriact i i nn Im n i mill ml i I Mjut subsequently 

has been documented liom ovei fiftv site, in eioht Mate 1 ,. I he two species co-occur at 
several sites in th >n and conver 

sionoflongleafeuo, mo . jk i i n n t h lit r n f l\ to vield many more 

I ul it i h ' 

Field work in North ( i I Ii ihmt II Ih 'Cm >n mev the North Caro- 
lina Natural Heritage I mourn I mi I i um / iilit.n i > i un t it, I the state Wildlife Re- 
sources Commission. Field woif [[i i i.Jii 1 1 ■- I Im Albritton of the 

UMISS Forest Landsjulie Moot, n e Leonard, and Rot ' i< ' ind In Alabama, Bill Finch 
guided metoseveiul Im hi quoiuv koojtomt Mteo.iiiehem i hilou.i contibuted many pleas 
ant hours afield and ( i I | t ^| nun t t t ii inh It genusd thank staff 

of the following herbaria for access to collections: DUKE, FLAS, FSU, GA, GH, IBE, NCSC, 
NCU, NLU, SWSL, US, USA, USCFI.VDB. I am indebted to R. and an anonymous re- 

st I t il Ml I e 
nth, | I ii t to t n th I Ii in' i I tda 16:581-587. 
in Clayton herbarium. Rhodora S0:d 1 -28. 
species of the o m. t i > , m I ' h^nJiospora occ 
ew York Acad. So. 11:74-93. 

1883. Flora of 

Company, New York, N x 

1897. Flora oft 

pany, New York, NY. 

iewell,A.F. 1985. Guide to 
of Florida, lallahassee. 


Godfrey, R.K. and J slants of southeastern United 

States. Vol. 1. Univ. of Georgia Press, Athens. 
H S L K N G i in and l E b ■ I h I h ,t "t the vascular plants ofTexas. Texas 

Agric. Exp. Sta., Texas A& M Urn . i it <\\< l tation 
Holmgren, P.K., N.H.Holmgren, and L.C. Barneii (eds.). 1990. Index Herbariorum. Part 1:The 

Herbaria of the World, Hn - i,ti nN , il i >ta> u al Garden, Bronx, NY. 
Joni s, S.B., Jr. and N.P.P >r • 1 ,l II I i ' i . i ih r' iht 11 in of Georgia. Dept. of 

Botany, University of Georgia, Athens. 
KARTEsz,J.T.1994.Asynonymized checklist of the vascular flora of the United States, Canada, 

and Greenland. Second ed t 1 i 1 iPi- Pottland, Oregon. 

Radi ord, A.E., H.E. Am ls, and C.R. Bell. 1 968. Manual of the vascular flora of the Carolinas. 

University of North Carolina Press,Chapel Hill. 
Smith, E.B. 1 988. An atlas and annotated list of the vascular plants of Arkansas. 2nd ed. 

Sorril,B.A.,B.vanEeri nandMJP 1 < i t| Pm* from Fort Bragg and Camp 


Dept. Wildlife & r Field Office, Baton 

Ri. i v ( IIP: m v k. 2000 A Primer of Conservation Biology, Second Edition. (ISBN 87 

732 3 pbk i ^ it I H i ti r I it I Hand MA01375-C 

U.S.A if ubh imunauer.ium) s <\ b - phi ^ illustmtions 

lions in he 

ild print, <incl itulude numbemd tab 

les and figures. Ann 

II i 

iPb,' ',!,■ 

■ b i| n t i M 1 i 1 in* 

-hi , r 1 u t i,i h i 

oxes to further explain conce 


il 1 ni| 1- it , -tth- tt- ■' u t 

xes its purpose as a 

nasiantioduaow underpin 

,lud' ill- Inn b,r i bl -I -i| 

& oli ig> i 

5 prerequisites so that time can be> 

spent on suppleme 

ntaty articles or perhaps ash 

ternr pap< 

?r.The summary points listed at the 


' 1 ' 1 U'l 1 t, bi i |t„| it 

it I mint ihVi.iti ill hi ml r I In' tn it i * u i\f bibliography, and 

A suqaestion lor iinpiovemenl wouU be- to update tin- two boxes of Chapter 
minimi lamhm, | i m in t f f i i (jtesented, respecth 

int. nds t K .pi mi th. hit i u I "IihI - h m u' nut -pui itch mpht Hm nil t mi m I I n I | ,. iz( d evolutionary re 
aid mcornmend explaining the hmtuiC-y in teims ol nenealogy, so that it would 
n what students will learn about plul u * in l - |uent systematics cou 

and the profound effort 1 , o' habitat Ma. m mutation bcttoi appiee 
iienbm-.'nmtubutnm thai nioiet n;ai OudioninCiLm up: CNA finoni ;. 
ilu <|i nt tu in ill u tfbi i ' u t in . I. in n . t itt'in 

quality paqesntsfipuK 
pli ii urn ire in i , I n 

- ' dl h oh - t mtm i f it f 'i 




Brett E.Serviss 

Department c ' 

Sidney T.McDaniel 

CharlesT. Bryson 

I ulii spC( it-, 'i four . |. r ,, | i ,4 V,, , ', , , „ , 

li h I ii Mi' , 'ijt||, a -i m l tilth i i i' 1 

The Araceae (including LemnaLfMi Mu t i 1 t n im nmily is represented world- 

wide by 1 05 genera and about 3,200 species (Croat 1 998;Mayo et al. 1 997). At present 1 6 
genera (excluding Acorus that it is now placed in the Acoraceae) and approximately 31 

nentsofourfloia mi , - ml i n i tin non native natu 

ralized (01 potential i in ml m i [i mo- I i n | . cies in the United 

States. The origination ol ihosogeneia I torn vat kins i topical mjions and the characteris- 
tics of the naturalm I - ii Hi mi mil i nt pi isticity for adapta- 
tion to different clirnatiL conditions and habitat types. In the southeastern United States, 
differences in season il '- iup> ntun tani- I H it* »< mi HI o-tn species composi- 
tion all < umbnu to | t I . in ni iu i i i il I i. mm fiom the conditions in which 

these species evol I h , t th > Hit | m , | i i me degree of range 

expansion since introduction, and three varieties ol bo/ouom? esculenta {Colocasia 
esculenta " , - ' . n , , I'lhs, and Co locasio esculenta var. 

nymphaeifotia) and Xanthosoma ujitt n« rapui I ming invasive. Intraspe- 

cific morphological m ■ m ill mm , ml i • < >\ nil mty in appearance 

between the seveial uiMn T v , n ,, , and certain species of /Vocas/'a and 
Xanthosoma allow foi ntm il im i rt lit ( . in In Id idoi tifii sti n > 4 thisgroup.Ambigu- 
ity in terms of spec no lonlilu at ion ,md ]>■ smitM toi n asi ni<m ol these introduced 
Araceae necessity il n ui in tmin Ml n i |l il md ecological vari- 

naturalized ranges t mil ni i'i u i Ii i tuft m while Colocasia, 

in addition to occuniu i il . m ih t H t h n In ' >f ■ i imu lexas, also occurs in 
scattered locations tl i mol mini i milt i t n i noi , ,< alentavat.aquatilis, 

Colocasia escul luorum are estab- 

lished as far north as east centra i< ■ i i ithmt on iana md throughout the lower 
in Inll d did inn - im Milf 1 i hi II. i hiee varieties of C esculenta are 
invasive weeds of son mac iu,it ic and aquatii condil n ihroimnout the Gulf coastal re- 
gion. Xanthosoma s 11 mi Mil n i i il Ii In 1 i > n m weed, is becoming 
more prevalent in i < ninsul n U rid m 1 mth. rn Tt <a 

/was mil iu I n I Hi- diluted i ite is a food rop lot it i dible tuben 

ornamentals with minima! i onsideiation as d food souoe(Neal 1991;Vaughan &Geissler 

tubers are a source of food and pmpauulos lot ornamental us< > We would like to clarify 
and delineateamon i n i r i t m mm i H iitterences between 

llu p i ii I l I W i / 

Aroideae (Mayo et al. 1 I n I ui n i . o i i <ph I. tu as once thought 

when formerly plain I in tl i. Hd ul t in n i < ilm mleotropica! Alocasia 

and Co/ocas/oiem-iiu u ri , , i i I nl I i tin n. >ti pi a 

Caladiinae (French et al. 1 995; Mayo ei ,il. 1 9')/). Subfamily Aroideae (based primarily on 
Croat 1 988, 1 998; Grayum 1994; Mayo 1997) are perennial, predominately terrestrial, rarely 

hemiepiphytic or epilithic herbs hi< h pi In nte or yellow to orange latex sap. 

Plants caulescent to acaulescent, hypogeal, stems sometimes aerial, erect to decum- 
bent, often rhizomatous and tuberous, lea^e i-h"'it_ . . a^, are-lanceolate, ovate- 
elliptic, or triangulai o a- n u | n 1 I i ill . utt ,tt hastate, or cordate; 
petioles elongate mill h nh f i i i i ti ' h >u it 1 1 newest unemergent 

the apical portion of the stem and emeigi f m i i n ithm the terminal petiole. 
Laticifers present, simple, articulated oi an. t mi ing 1 h ho . h leids absent. Plants 
monoecious (occasionally polygamomonoe i >us) in Irog nou | erianth parts absent; 

atanthesisthe spade i in, I [ , i i h t 1 e iimiimI ippendix and a region 
between the stammitr- ,n I pi i 1 H m i.d if i m liodes (sterile stami- 

nateflowers);stamensareconnat. ml n mi i u iudiid,tiuit a I numerously seeded 
berry, endosperm copious or no! pr I -i a i I u ect pollinated by various taxa of 
beetles, flies,and bet )« p I i a- ti n ii u tranean tubers and 

or rhizomes. Production of these f> intn i in ill f a efficient and prolific 
e-g>a iii .e '-'production. 

Many Aroideae form ieui ' i I n if i i li nil iduals.This is the 

typical method of leptoo mioi lip I n- i u nm he I ) l > and one of the pri- 

mary factors allowing for natuialr a t o> let T Hon also occurs if environmental 
conditions are conducive, but this is :am in the US. 1 hroughout the plant, but concen- 
trated in the tubers,are deposits .P p - a i ul , of pointed calcium oxylate crys- 
tals. Plants are poisonous unless cooked or prepared in such a way as to remove or neu- 
tralize the crystals. II = skin and mucus 

KEY TO AL0CA5IA, < a, . . , i , \ \ Apra XANTHOSOMA 


tially green, sometimes glaucous or giauioscent o 
lobes of leaves ,i pit ally ,k ute; leaves not pelt. He _ 

s ol iif 

i i i I I in i Mini ml t ill lul eis ( t( 

9. Basal 2 lobes of leaves rounded; rhizomes usuall 

for most of then length abovi or hHoss umuno 

NOTE '-\-luu /-J! in In I tin tin fev! in .1 r oi| h -I nc il umlauts and | 
sion with the other genera. 



vnopitva 'st hot! in n.ivUis.i i i i I i I it ivum I 124 186 . 
Ii >tt- i n il i in I r A > ' IPS 1880-1881. 

Perennial caulescent or <k aulescciu ninths usually from a creeping tuberous-rhizome 
caudex, with or with it nni tin t t ft lu i, ill In econdary tubers, 

tuber offshoots at i I nh n -M n m I t n til umbent;leaves ovate 

to triangular-ovate, bns.i I lv sac nil ate or ratelv sliqlulv ! xhion n s|o,k nx pedunculate, arising 

separated by a sects ii ' ,.- n It \> t iN nor h I <t i appendix present- 
comprised of mi in il t n. ii f i till a i II 1 4 carpellate, 1-4- 
locular, stigma weakh 3 lobed, ovules 1 numei u pin nt tioo basal; fruit a 1 -numer- 
ous seeded berry som< lime nh cot lions of the .tnnuu pi t si stent; ovules 6-10, 
orthotropous, honn iti i | i iniiuht i i .i t f is funicles short; seed 

il " ih 'I ii o to ellipsoid, tes 

;ta smooth o 

, scahtnm 

>, endosperm . 


About 70 species nath 

/eto Indomalaysia and 

11 ' Mi' :l ' J s 


.Three : 

cies occur in the US.AIocas 

to macrorrhiz 

" an.; i, 

Kasia odora a 

re both estat 


Etymology— colocasia 


.1 a loci hi c 

j prefix mcani 

ng bedfellow 

• Ii | n Mi- 1 the morphologic 

,il similarity! 

Mocasia and Colocasia. 

KEY lOSPIUESOl ,\j(x A:>iA 

i. macrorrhizos 

1 . Alocasia macrorrhizos (L.) G. Don var. macrorrhizos in Sweet, Hort. Brit., ed. 3. 63 1 . 

H k.f.,FI.Rr.lnd.i S26 1./ , I , ,i m I i iHrad niya 1 1 

KrauseinEnglei I'tl - i . m ' E 1 ^ - 1 < ■ Fig. 1 

1 , >n <rorhizonR.Br Prod 336 lHi - L I nttm K tt 

Endl., Melet. Bot. 1 8. 1 832.Alocasia macrorrhizon (L.) G. Don in Sweet, Hort. Brit., ed. 3.63 1 . 1 839. 
Coiocusia macronhi.iu lv,n:h, ! num. PL 3:39. 1 8-1'' . 
! | PI '66 l 7 53. 

1 2. 1 786. 
Arum indicum Lour., Fl. Cochinch. 536. 1790 (. olot , i ' I um I mih Enum. PI. 3:39. 1840. 

Alocasia mdica (Lour.) Spach, Hist. Nat.Veg. Phan. 1 2:47. 1 846. 

Caladium giganteum Blume, Cat.Gew. Buitenz. 103. 1823. 
i '' pot ' a • L e I ' it H >it Pdi,ed.3.385. 1829. 

'■ 'iincVl:!!! (P;>< ', '.'.'»."/:' i ! !■ ' MV'I III HO' )H Bot. MiSC. IH'P). '8 "I!). 

t El nk o Fl Filip 658. 1837. 

Calla maxima Blanco, Fl. Filip. 658. 1 837. 

■\'um gni.njiloiiian BL'.iu o, 0|\ LP., ed. 2. 1845. 

' < " '' n H - Ho t El I H' i i i irteriz 136. 1857. 

Caulescent herbs up to 5 m in height from a tuberous caudex; stems to 1.5 m tall and 20 
cm in diameter, becoming thickened and caudex-like, erect or often falling over with 
age and resting on the ground, basally covered with dry,chartaceou:,,nersisiontcataphvlls, 

able in length, elongate, white to whitish-green; leaves 35-160 cm long and 30-90 cm 
wide (often smaller in dimensions if on young, damaged, or spring emergent plants) 
ovate to ovate-tnaiNu' it i| i P ul unit > to short acuminata ! i illv sagittate, young 
leaves sometimes pel p- margin lungly undulate, primary lateral veins 4-8 per side, 
midvein and lateral veins prominently raised above the lamina; petioles 70-1 30 cm in 
length, lamina and petioles green; inflorescence of 1 -numerous spad ices, peduncles 20- 
50 cm long,spathe 1 1-35 cm long, 3.5-6.2 cm wide, chartaceous, a pically withering with 
aui' intc to > iie^i . , j ,i _>ju\ Pu muni] p> a I > , i it orange with age, 

' ■ , * .. 

spadix 11-35 cm in length, yellowish! u mi ,,-mli-; 7] cm long,foi tile slammato 
region 2-7 cm, sterile staminate region 0.3-2 cm long, fei 

long, pistillate flowers emit the odor of ether prior to lortili/ation; berries ovoid, orange 

Some species of Alocasia, mo tim| ii nil i , m it d 4 7 n in f >i 

tant food crops in tropical regions, particularly Asia, the south Pacific, and to a lesser 
extent, Africa (Bailey 1 997; Neal 1 991 ).The actual crop and source of nutrition is the large, 
edible,subterraneantubers.ThetuberspioM h I • t i t u staich (approx.25% 

by weight), ascorbic acid, low amounts of vitamin A, B complex vitamins, and small 
amounts of protein (Bailey 1 997; F 

cultivated in the southern US, it is only truly established in central and southern penin- 

to no enforced dormancy. Its ability to reprodui - - i> iti - I hi aided naturalization. 
Its rate of expansion is reduced in the absence of disturbance and where soils are with- 
out adeguate moisture and drainage. Once established, Alocasia macrorrhizos appears 
totoleratea wide range of condition variability (di ought, periodic flooding, variable light 
intensity, and occasional hard freezes) and interspecific interference. The rate of range 
lacrorrhizos may at least in part be impeded because it usually colo- 
s vegetatively by secondary or daughter tuber formation. This method of establish- 
t is not normally as conducive to invasion as is rhizome production in Colocasia and 
xThe apparent inabi'it t " , r m mu h fruit (in the United 

States) for dispersal and potential long-range ml m iti n ' t I iblv contributes to 

In the United ,t Hi fl , ~ t ltn d ftutt nioduct i i pically occur from late summer 
into winter,though flowering idii i oit u i, mt,< t - h - o . oral inflorescences 
are produced per plant at any one time. Aloui n ', i ' - i , r\f compatible but 
probably reguires out-crossing between different genotypes for full fertility (obviously 

single genotype, which may explain the apparent lack of fecundity in our plants.The lack 
of appropriate insect pollinators is yet another consideration. When fruit is produced the 
number of berries are usually few (10-15 per spadix). 

Alocasia macrorrhizos prefers moderate to well-drained but continuously moist, 
moderately to highly acid soil. It also seems to reguire at least moderate disturbance for 
establishment. It can be found along wet woodland edges,thickets,roadsides,old fields, 

established itspro U i t i II n hi tul e hi h|i . many small plant- 

lets or to a limited extent along tuberous rhizomes. These plantlets originate and grow 

limited exposure t ti n i i i| mi tin in it ' n t i aids A form with 
purple foliage very similar in color to Aloca * i n tun cultivated and oc- 

Comrnon names.— pai, taro, cunjevoi, giant taro, ape giant, elephant ear 

2.Alocasia odora (Lodd.) Spach.l 1 2:46. 1 84n;(Roxb.) K.Koch, 1 .c. 1854. 
Arum odorum I > I llotf m I 111 I I'M IM I Md Bot Cab 5 t 41b 
I820; Wicjht, l« M.lnd.Oi !(6): I '97. I 8AA. (. 'oitnlium odomtum \ odd hot Lab. t. 41 6. 1 820. 

Nouv.Ann.Mus Par. 3. 145.1834. 

AMo/oeom/outmo Schett in ( )es1i. IkoAAOx hernbl.400. 18V1. 

Caladium odoratissimum I ioti.ox t . Koc h, Ber I. Allg. Gartenz. 20. 1 857. 
Extremely similar to Alocasia n\< > hn lit' i i h ' i uoi peltate leaves and 
loliaue often glaucoscenl. Also simil 
, 1 1 1 ■ ,i pica I Iv i oi 1 1 ided, whereas t hose 

uv i ii I | t il it ipe mnl 
3. Alocasia plumbea Van Houtte in Fl. Des. Serres, Ser. 2. 6. 93: t. 2206. 1 875. 

Akh ooo nit'Uiiinj.) etiott io Oosti\Kot.Woenonbler 0. IsOO 
Caulescent herbs 1 3 m in height front a tuberous caudex;stem to 1 m tall, basally cov- 
ered with dry,chartaceous,persistentcataphylls ( ihi7omcs present oi absent, when present 
variable in length, elonoale, while to white preen, oilon with pinkor purple cob 

to short acuminati ! 1 ill n .I t i | M n > it I ,t pr unary lateral veins 

od above 1 the lamina, petioles 
jo ,11 ,,,1, d'l( puipk puipl, 
lerous spadices, peduncles 25- 
50 cm long,spathe 10 20 em long, 3 7.5 t in wide, char tacec his, a pically withering with 

\ \L [)Uiple pnl t ) piui I | | I 'I if .id , | .Hi I oil, n \ l"let t it , 

spadix 10.5-19cm it, unh -II it it M nli-M 13 cm long,fertile stami- 

1 .5 2,5 cm long, pistillate flowers emit a slight odor of ether prior to fertilization; ovary 

: to purple-green foliage (petioles often pink or pinkish-v 
when on young | I, -tit it it it t under shade a 

/persists subsegucnt 


Caladium Vent, Mag. Enc. 4(1 6):463. 1800-1 801; Vent., Descr. Pl.Nouv.Jard. 
1801;Vent,Arch Got U ei|m<p ' i I 1 ii I I ,i I , I , 1 
Engler, Pflanzenr.4. 23E. 71:23 N20 I 

Cnt>>:;pa:1i.\ i Kcrh, nd. Sr-n Hot !. Bemi 1 ,-',.-,6. 

Aphyllarum 5. Moore in Trans. Linn. Soc. London, Bot. Ser. 2, 4:50 1 . 1 895. 
Perennial acaulescent herbs from a creeping tuberous rhizome, reproduci 
by daughter tubers. Above ground stems erect; leaves ovate to triangular-c 
sagittate, peltate; spadix pedunculate, arising axillary and subtended by 1 
nate bracts,staminate and pistillate flowers separated by a section of synan. 
ile staminate flowers), appendix absent; spathe present, longer than spadix 
basally to form a tube which encloses the spadix until anthesis;the basa 
mainin i ti jhl closed i mi n ii il > m ortiiii d f ,-m mi n n mdn. 
mens; pistillate flowers 1 -4 carpellate, 1-2 or incompletely 3-locular,stigmadi 
1 -numerous, placentation subbasal, pseudoaxile, or parietal; fruit a 1-nume 

0:348. 1 801 ;Vent.,J.Bot.(Scht,i im did 
). (Fig. 2). Aran 

U'xammntly none establish 
Etymology. — Latin:/ce/o 
and Caladioid genera. 


1 801. Engler, Pflanzen 


Caladium surinamense Miq., 

Delect. Sem.Hort./ 

5 from 0.2-1 m in height from a tuberc 
well-developed, plant without elongate stoloniferous rh 

basally sagittate, peltate, margins subundulate, angular, c 

spathe 7-25 cm I te to white-green 

on the inner surface;spadix 9.5-14 cm long, white to yellowish, appendix absent or much 
reduced, region of fertile staminate flowers 3.5 5.5 cm long, region of sterile staminate 
flowers 1-2 cm long, region of fertile pistillate flowers 3.5-5 cm long, pistillate flowers 
essentially without odor, ovaries with 1 -numerous ovules; berries white. 

A highly variable species comprised of numerous varieties separated primarily on 
the basis of foliage coloration. Commonly cultivated in the southeastern US for orna- 
ment but not currently documented as natm ih ' or is a prolific seed 
producer and is spontaneous in new growth forests and moist disturbed areas through- 

outthetropics.ltshould be expect- 1 - i li , imbed sites, woodland edges,ham- 

mocks,and fields in southern Florida. 

' ' ' ■' '< il i li ii - I- i I it eai, angel-wings, mother-in-law plant. 

Colocasia Schott in sdion and lim I i> l< [, n is |,: ;> nom cons; Krause in Engler, 

Leucocasia Schott in Oestr. Bot. Wochenbl. 7:34. 1 857. 
Perennial caulescent to acaulesceni herbs from a creeping tuberous-rhizome; tubers 
and rhizomes usualb f t -in i i i |i I i i means of secondary or daughter 

tubers,and or rhizome \l a ii 1 i i| i - iMfi,ubi ii ' 

sallycordate,cordate-sagittate,ois f iOiSit t | o ill ( itaie, often glaucous to glaucescent 
abaxially; spadix pedunculate.ansino i-4 a » 'I tended by 1 or 2 acute to acumi- 
nate bracts, staminate and pistillate IS . i ( ,, h b i ,. n it ot >ynandiodes (ster- 
ile staminate flowers), appends pr< <tt \ itl - en, 2-6 times longer than spadix, 
constricted basally to form a tube which end >• • o uv ■ spadix until anthesis; the basal 
portion remaining tightly closed s 1 - mnri it- ' i n| if mncate synandria of 

numerous,placentation f nitult jt I nue u I l'-i it i persistent stigma; 

ovules numerous, hemiorthotropou m hi I i ,h, I , , I to « I h pisoid, testa costate, 
endosperm copious. 

Seven or eight species primarily of tropical / ia nm n lrica.1 'o species estab- 
lished in the US but only certain varieties of Colocasia esculenta widely naturalized. 

Etymology.— Greek: kolokasio, term used for the toot of Nt'luniko nucifera Gaertn. 
ae), thus applied to Colocasia because of its edible tubers. 


I , mi 1 1 nil" i I 1, I.C.esculenta var. aquatilis 

ly at very base of tubers 8. C.esculenta var. nymphaeifolia 

it - i imiM' ' numerous, o'l en n tit i In. r r ■ i iim i i i 

!'V (III III' i|n h '" Mil I Hi I t. I II III I lup Nit |i | ii mil u Mt . 

I h t i ] iti ii i m i i t it f k lit 2. C.esculenta va 

. I I ,nl 4UI I mt ' i ill ^11 Ii i n ' ui i'ti n . mill 1 m I f • I 

in] „, M| ill le, I it , hi Ii Uii tit i Dill an , J i UIIIImI ■ LJ T auP ! MS 

ally whitish; spathe white to cream 9. C. gic 

t i hit I i i i i hi' i n l il in i n i II n it 

m with white coloration; spathe yellow-green to orange, or pink. 

i iin, n mi npi m uit i in. mm ill ui | nple) petiolesand 
ivver min veil", pntpp oi puiple-blacu leaf iiimli'im, usually 
, Nil > I it in n lm I p[ i inn Ml - i I'll mh| l 1 i ltd, ii>| 

veins on tne iowei suiPui epwith oi without purple .■■laipiiu; plants Luge, 
often 1.5-2.0 m tall 5. C.esculenta var. fonl 

ikidii t R Bi I i i II I II II 1-1 lit'' n .Fn tl ' ' I. t L ' 

IN" Pu/wj it l t i i in Mil Prodi Syst.Aroid. 140. 

1860; Engl in A Dt M u . ,i mmu , P P^ • tu i m ■. - uu . I nil rtlanzenr. 4. 23E, 71:68. 
1 920— differs from the above varieties by the absence of an appendix. 

Colocasiaesculenta I hottm Itott FndL.MHet.Bot. 1 1 >Kunth,Enum.3:57. 


(I i Pi m in t, N n II i I t I t 1 I i u u P it till I I in I h 

r - U P h I pi P t mstMi PI iim - J JPC 65 1920. 

rily on the basis of vegetative differences (Engler&Krause 1920; Hill 1 939; Neal 1991 )The 

duced, texture and chemical composition of tul I n i il romp forma jt j i I 

foliage morphology and coloration. Flower and fruit characters are also used. Varieties 
are used as ornamentals or as food crops for the edible, high carbohydrate tuberous 
rhizomes; some varieties are also invasive weeds. Spadix morphology and color can vary 
depending on variety. Flesh of tubers ranges in color from white, yellow, lilac-purple, or 
pink to reddish. Varieties of Colocasia esculenta are separated into two groups based on 
differential affinities for available soil or subst Ni m, .: , M or wetland varieties 
prefer aquatic or semi-aquatic condition-, of continual soil saturation, whileupland forms 
prefer well-drained but continually moist so'l , f I . i| -.ill i . r.e under either set 
of conditions. Vegetative preparation - ut ! Ii inn if 'In . ludex, new secondary 
ordaughtertubers,rhizomefranim ii n ihul r .| i ,ne third of tuberand 12-20cm of 
stem; Neal 1 991 ). Propagation by seed is also effective. 

-.-<■'. u, ajz.inespec irs , jhoii ii t i t >y Linnaeus as 

'My-. '■ ; 33. Our varieties of 
attributes are discussed below. 

Varieties aquatilis am , < • m ithr only slightly in tf 

i I i Viiieh nil ,i )i ii i n il [Ml Hit in I ii ini| I 

n •:■■■•• i; I v' green in color, usualK h it i il I pi tit ,i ,d loss developed r 

whereas both aquatilis and nymt ^ m i petioles usually 
dish o- purple coloration, often ink t n-i il ml - I. elopment, and long, well- 
developed rhizomes. Varieties aa - ' n I ' i//r/ essentially differ from each 
o:'ir only in area of origination of rhizomes from trie tuber or caudex (see key to spe- 
cies of Colocasia). The i Hit- hi i ir I n i| | i d I i . im and ecology and 
probably encompass only a single, highly variable taxon. 

1. Colocasia esculenta (L) Schott. va r. aquatilis Hassk., PI. Jav.Rar. 150. 1 848. (Fig. 3). Cala- 

i f ii jl I rl mooi 4 f 1 bf- 1920. 

' oOLmoo-o'ooio'fTiSLhotow.os^m/o^oo m „ , | i,B„ Nom-i'.t uu.l' India, -Neo<-i! ooci [-',l-,i.-.:- 

Essentially acaulescent hetlo 'o 1 in mi I >li> ' i i itl . f il etous caudex, tuber n m, ih.nt veil-developed rhizomes presen- 
ts 2.5 m in length, elongate,brittle,and cove I it i i ,miil ataphylls, distal por 

tion greenish-white and usually larking in . uaphyll il ives 7-40 cm long anc 

5-30.5 cm wide, ovate, ovate tii, ii ii i i i Hi i i| i n unded to acute, ba- 
sally cordate, cordate-sagittate oi iiitnt mi m in I undulate to entire 

primarily lateral veins 3 mi -i o in i : i o it u meon to yellow.or purple, midveir 

\ k,,1 Lolotasin psculenta{\ .} Schotf 

glaucous to glaum , r i il i- ill i 1 n |m M , ( , n to red or purple, 

often green and splotched with pi ii 1 1|. omnium pink basall inflorescence of 1-nu- 
merousspadices, peduncles 15-50 cm long,spathe 10-40 cm iong,chartaceous,broadly 
acuminate, yellow-green, yellow, yellow-orange, or orange; spadix 6- 14 cm long, yellow- 
cream and often tinged with orange or pink, appendix 0.5-3 cm long, region of fertile 
staminate flowers 2-4 cm long, n mm i > ' i i mimmite flowers 1.5-5 cm long, region 
thout odor; ber- 
ries red, ellipsoid to oblong and 3-7 mm in diameter. 

]uorum and 
nymphaeifona have become pantropical weeds of aguatic habitats, particularly rice fields, 
fresh water marshes, lake and waterway margins, and drainage is truly aguatic 
and cannot tolerate prolonged conditions of drought without the onset of dormancy. 
Extended drought over several in uii u ill In n I I ii 1 1\ It was introduced 

into the United States as an ornamental and is completely naturalized as a member of 
our flora. Colocasia mo/t > v , n ,. m ',- ■ m an invasive weed in the south- 
eastern U S. It ranges from eastern i i n i 1 u um; pi ltd Alabama, through- 
out Florida and north to Georgia V m ' i th< most established of the varieties 
of Colocasia esa under any condi- 
tions with adeguate moisture th- h< ' \, i n if rub nitiatihs in the south- 
east US are continually wet or flooded roadside ditches;freshwater marshes;open areas 
of floodplams; fresh water beacln ml I ill m i ilmm th. - dqes of ponds, lakes, 

continually wet soil i a p< u nil il n I hi i , , ' m< mm , little to no distur- 

bance for hasth p o i n i i i tdv invade and quickly establish 

vegetative colonies regardless of ilii'tlon ,i inpom til oi degree of disturbance.Small 

colonies and isolated juvenile plain - i i i it Mnt pi a i n I ^cause of vegeta- 
tive propagule (rhizome fragmen ts) frag im -it, Hum, translocation, a nd subsequent plant 
regeneration. This scenarm i turn , n r- h ^ ttl h f ^ >f fruit production 

sagittifolium, vanet\ 1 1 <> < i m I i >i I i i i .11 n i moie vigorous rhizome produc- 

tion.Thesestoloniferous th nu n i i i ' i i i i I m along theirlength 

(up to one per no at the rhizome 

terminus. Each plant can produce 2 1 mm i th m h i m time. Rhizomes are 

rhizome production. Rhizomes severed befor< 

i i mots have emerged fr 

om the nodes 

can root and produce new plants on moist < 

soil or in standing water. Extensf 

jc colonies of 

variety aquatilis occur throughout Florida, bi 

jt are otherwise currently restric 

led to a zone 

approximately 100 miles inland from the Gulf 

: of Mexico.Some isolated and lo 


lations occur slightly farther inland and nortl 

iwan 1. 1 unci development is us 

ually reduced 

or almost absent, the subterranean portion i 

jsually only consisting of many 

i tibious 

roots and rhizom* Li i n n m h m | i nnccd tuber develop- 

ment at the crown region. Plants with la i ( jer more developed tuhotmooe Colocasiaesculenta 
var.onr/quorum) are not uncommon. Once fin. eer p. . 1 1 , t, I i inie.they often emerge 
from the soil and U i t mi i in I i i u i i i i producing new plant- 
lets.At various lengihs.the rhi/omes son letinies repcMieiiate ihe qiwn substrate.Newcolo- 
niesare nearlyexelu ,i >l i u i I t i I i h | i hi tahlishment of nodal 
rhizome fragments or small plantlets. 

v hi, t\ uquatili onlvounsionnllvpiodim dlowei m In Puled Stales, Flowering usu- 
ally occurs in mid hi 1,1 muni hit i im- I i 'ii i -p ml i \ toher), during which 
time conditions are dry throughout much of tie tin oei I || the site occupied be- 
comes moderately dry, flowers are often produced Usually I- > inlloiescences are produced. 
Plants will sometime- i 'I t n t i In i hunt i in nnalK produced in the 

Colocasiaesculenta var.aquuti i , , i t I i , li I | I i it i livision from rhi- 
zomes. Plantlets can be removed immediately alter emergence from the rhizome so 
long as the node from which the phiiulet originated is left att.n heel, even leafy stem 
sections with a functioning apical men , ten ill mi atei or wet soil. It can 

be grown with or without soil as long as standing water is provided, but establishment 
and growth is enhanced with some type of rooting medium. Plants thrive on excess 
nitrogen and phosphorous. Mature plants transplant readily. 

A variegated fc 
on leaves and pelioh in mtn 'gin nth tint Id type populations. 

Common names, agnatic elephant ear, elephant s-eamodd mi 

2. Colocasia esculenta (L) Schott var. antiquorum (Schott) Hubb. & Rehd. in Bot. Mus. 
Leafl. 1:1.5.1 932 Colocasuuo ,, ,, ton m lm iv n.JI ' V lot Mot 1 8. 1 832; Schott, 
Syn.Aroid.41 18 L t I I F li i - i i 1 i ' t H Em r t h ^ 1M1 Lnql in A. 
])( ., Meinour. Pha-mMS'l. 1879; Hook. f. in I'ot.Mna l>() t./nel. IS'Mp n ,,|.& Kiause in hnqt, 

Vight in Contr. U.S. Natl. Herb. 9:208. 1 905. 

eties aquatilis and nymphaeifolia .variety 

eahy gmen | letmles, l.iigei tubets (es| >e 
dally those at the u a el miah I il fid -mutes though these 

can approach tin I t it I ill p i i r I m ' n ti n imtn 1 1 nt 

and nymphaeifolia. 

than variety aquatili An iy ant i pi > m luiins of ponds and lakes, but will 

also rapidly colonize and invade lawns, flower beds,and other disturbed sites so long as 
adequatesoil moisture is present.Establishmentand complete invasion of various lawns 

inn inr it-n 1 b f <- in, [ iiwii ! on i ui i n i ' U I in m <\> , 

and nymphaeifolia. Plants prefer moist to saturated soil or shallow water and usually do 

not occur in water over 30 cm deep. 

The primary mode of reproduction is vegetative and occurs from secondary tu- 
bers, or plantlets produced from rhizomes. Variety nntiquorum requires similar condi- 
tions for growth, uTjmo tun n I - i n i mill n i n i"ty aquatilis. Flower 
morphology is similar to variety m.A n ij i m i i:,,; ttuit is not normally 

variety aquatilis. It occurs from east-central fexas, east 
Georgia southward throughout Honda. 

Common names— Egyptian taro, culgas, qolqas, t 


1 1879; Engl. &Krause in Engl.,Pflanzenr.4.23E./ 

1. 1 40.18i ii);lnql, ' 

nA.I i i it 

< tw'uKis/iJ escui 

< ' ' > 1 tin ' t it >: < f mil Vt i , Fill u 1 lu 

igsqescHlx train 

=,F,o re „ g eb.8 

terized by its large tubers (up to 22 cm long and 1 8 cm in diameter), leaves, and size (to 
2 m or more tall). Usually only a few tubers are produced per plant at any one time. 
Colocasia esculenta is the most important species of Araceae in terms of food produc- 
tion worldwide (Bailey 1 997;Vaughan & Geissler 1 997; Neal 1 991 ).The tubers have a high 
starch content (approx.25%),some protein,and 1 3 mg/100 g vitamin CSmall secondary 
tubers, also known as cormels, are equally consumed (Vaughan & Geissler 1 997). Sec- 
ondary tubers develop directly from the primary caudex or at the nodes or terminus of 
rhizomes.The foliage is also eaten and normally contains 7 mg/1 00 g carotene, and 52 
mg/100 g vitamin C; all parts of the plant are used as food in certain regions of Africa 
and Asia. Removal of calcium oxylate from the tubers is accomplished through boiling 
(Vaughan & Geissler 1997; Neal 1 991). Chemical composition of the tubers produced by 
Colocasia is similar to those of Alocasia and Xanthosoma (Vaughan & Geissler 1997). 
Colocasia is grown as a food source in the United States and Mexico, throughout the 
neotropics, Africa, and Asia. It is a staple food crop in many areas of Africa where it is 
known ascocoyam (Vaughan & Geissler 1997;Neal 1991). The amount of tuber produc- 

plant (Young 1946). 

> , 1 I' 1 1 I pi III, . il III 111 pill 1 

4.Colocasiaesculenta(L)Schottvar.euchlora I iJnnl ell- ■> ^ F Hill,Bot.Mus.Leafl. 

77 1^39 ( ' , | (mmI II mini ,P I ill il i I App4 1854.Co/ocas/a 

-inliijuori. I '1 it ' o hott 'i mi I - I • I i li 't i i 1 i | -- 

mil ii \,K fl n ) |i I in 111 I n il hi in In il llhi i I BE. 71 :67. 1 920. 

Plant to 92 cm tall, leaves with lamina bright green, margins usually purple, petioles and 
veins red to purple. Rhi/omes to those ot varieties aquatilis or nymphaeifolia but 
red or purple in color and usually shoOm. Mmilai to variety fontanesii in coloration but 
smaller. Vegetative i« | i I t nl i i o I uidaty tubers. Grown in extreme 
southern US but no cunvnl recoid of establishment outside cultivation. 
Common name, -purple stemmed lato. 

5.Colocasiaesculenta I I'-h m , fontanesii I i ' ( i Mus.Leafl. 7:7. 1939. 

Colocastafontu 1 on ' ti I i- m- 4 i antiquorum Schott 

ii / . )M it - h )U n A, ,i | l 1 i h Hi I i It I mi I li 1 mi I nil n A. DC, 

Moikxii Ph in ' ! 'I 'H >o I mi, l i 1 , , I i I O M , , i | ,, , I , ,i , ml n , I'll hm hi 

„ ij , " , m • ~ it ■ Pnql.m "^ , , , , I I, ^ l < 1 - I - t Lai. Hort. Pan n I 3. /. 

Caladium colocaooides Hon. Pan. ex Rionmun Nouv. Ann, iVuis. Par. 3. ISO. 1 834;Kunth, Enum.PI. 

Colocosia violacca I lot t. nx I iook.t. in Pol. VU). 1 26. t. 7732. 1 900. 
Nearly identical in too i i Put I mi. a In tall, and more robust 

with the petioles and veins of leal usually black oi oatk put pie; Un una dark green, often 
tinged with purple coloration Pin mo in, hi i, a t< ' s and nymphaeifolia, 

but purple to put pi PI i Mini t I m Hit io| f no pie to purple-black 

foliage is also cultivated in the US. I amina, petioles, peduncles, and spathe all purple, 
undersurfaceof leif vi iu i n I t i lf i nl t t f tuple eoloiation. heaves 
on plants grown undei shade mudi'iions and newly emi'ignit oaves are blue green 
with purple coloration along veins and petioles. Newly emergent leaves on juvenile plants 
\. 1 his foim actually appears to I >e an n it ei variola cross between variet 
'ont mi < ami ' '- b it t pi n i , PI it t i li It. t>n the two. Both 
as are vegetatively aggressiu imiln m t t I ' <t il 

?d in extreme south- in Us mid >Psu d p, ilum - ;> t iii Uv at these sites but 
it i it I Pin nl ill iln a -I It i hi il i | ' ilP ueedy variety. 

i i i ' I m li nllU i t | n| I t ,i Pi I tin it tm 

.& Kiause) Young in U.S.Dept. 

Common name, aashcon. 

7. Colocasia esculenta(L)SchoUvn illustris(BulL) A.I.I lill, Bot.Mus.Leafl. 7:7. 1939.Alocasia 

i " Bull < it 4 1 Bull in I m mi 1 ri ^1 hull ii Wf 

Engl, in A. DC Monogr.Phan. 2:497.1 
Plants to 1 m tall and resembhn i m t ml 7 in form but less 

aggressive and usually with shorter, more tuberous rhizomes. Newly formed rhizomes 
are more slender than those ot i i r , , i ' ' < - Leaves with lamina 

green and spotted to complett !\ pu'pl I tween primal 'it primary veins green, 
lamina glaucousabaxi ill, p^tuA |i n i im I m md hiown coloration orcompletely 
red. Variety /7/ustr/b hculti ^>,< nil i me southern is not currently naturalized 

but spreads local l\ L ' uli it i nil n i I it i.c under favorable 

Common names. — Imperial taro, black caladium. 

n tlM I I sub t 30 1800Willd„ 

Cobcaiii.irryninhucifoii-jKutrr.l nun, PI, p-nt -47. P,;;;),-^.;,; nm, yuM.;-" vhot; ■.,:'.■ .■■■: " ,• ' ■ .. 
Schott, Syi.Aroitl.47. ' 8L6; Schc:ir, : 7: )4\ v./s:. Amid. ]■■:<). } ;s„0; hyil. r A. i X ., Monoqr. Pnan. 
2:492. 1 879; Engl.&Krause in hnc]l.,l 1 !! < in/enr.4 23E. 71:67. 1920. 

Colocasia esculentn) h m lilt t p n n nl i i < h i Ay from the base of 
ers. Similar range an i ^ I i i n mingly less common. 

u ju hi i' pi nit "it, taro. 

9. Colocasia gigantea (Bun H I 

( aulcscent.eiec pmttostt ial ot opilithic herbs to 2 m 

caudex. Leaves 20-1 20 cm long and 1 5-90 cm wic 

basally cordate, cordate-sagitt it- pi m 

spices short-acuminate, margins 

undulate; primarily lateral veins 4 8 per side, lamina gre 

en, petiole 20-1 50 cm in length, 

light green, white-pruinose; inflorescence of 1 -numeroi 

js spadices,peduncles 20-55 cm 

long.spathe 25-50 cm long,chartaceous, short-acumir 

late, white; spadix 8-23 cm long, 

yellowish,appendix 3-6 mm long, region of fertile stan 

linate flowers 2-9.5 cm long, re- 

gion of sterile staminate flowers 2-4.5 cm long, region c 

'I I'M tif [ i I Ate flowers 4-8 cm 

long; berries oblong to 1 cm in diameter. 

Colocasia gigantea is similar to Colocasia esculenta 

and Alocasiamacrorrhizos, and is 

included here to alleviate any confusion in identificat 

ion of plants encountered. This 

\. Colocasia esculenta (L.) Schott v; 


nd t< tn if hi in t i i 1 1 icli | i fjiiin nl msing axillary anc 

inate flowers), sterile appendix reducer 
5;mc,mcons::k u>d 1 :, js.'d'V io form a mm 

h encloses the spadix prior to anthesis, the basal portion remaining tightly closec 

linate flowers com inn i- n i < pi till ii flowers 1 -carpellate 

locular, stigma discoid, ovuf 

indrical, furrowed berry; ovule in i ' m nui nnus,anatropous o 

ianatropous,funicle Ioihi . di n i nu>nt< uiclosperm copious. 

About 58 species oi uiirnilnu i in ih. m impu mimnt mil „/ n < mm mm 

?d in the United States. 


sagittifolium (L.) Schott in Schott & Endl., Melet. Bot. 1 9. 1 832; Engler, 
mzenr.4.23E m I i I hig S I , M 

maefoiium I I . i _ 1369. ^ >- lT ;/,• Jr/:,: ;,-r; iGquntoitun) [\ • - i ' 1,i , [ i„ 4i r , 4^ 
i1;Veric,Arch.Bot.;lo:p/i(;)2li!:^S1.180';Vert.J.Bot.iSc."ic)f'; , pr)--:iD!:319. 1 801 . Conn;;'- 
;':'in-r/o/. ; .0'.'^;LJ'VVi ia.N\ 1 1.4:48''. imvSjv^viV^nVeu.v ' '1 84m. \,;j.'i/!. no? -H-..1 <og:Hueh:;r: 


I ll 111(1 [ U I I I I I t ' I I 1 

Xanthosoma zanthorrhi/on C. Koch, Bonplandia 4:4. 1 856. 

Xanthosoma wendb" n Mn ' it i O i , t hr 1 5 J . KHmS 

(\iiadium utile Hort.ex Fng ; .'n A DC. iV'.onogr. Mmrs 7:469. 1 H79. 
Caulescent herbs to 3.5 m in height tmm a tn .1 m> n n his and tuberous cau- 

dex poorly developed or absent it u r Dp ,1 . I developed rhizomes present to 
1 m or more in lei - iut scaly covering of cataphylls, often tu- 

berous at various places along the length, white and often tinged with light green; leaves 
20-1 20 cm long and 1 5-90 cm wide, ovate to triangular-ovate, apically acute to acumi- 
nate, basally hastate-each lobe di |i , it. [iKMin ill I 11 broadly undulate or 
entire,abaxia! surh< mm I hi i 1 In nl t side whitish or light 
green, lighter than the surrounding blade, midvein and lateral veins sunken below the 
blade on adaxia! side 4 I 4 1 I imm to glaucous; inflorescence of 1 -numerous 
spadices, peduncles 20-30 cm long, glaucoi 1 laucescent pathe 18-22 cm long 
and 4 cm wide, chart 1 > | D , - n r 4, nt tun n team to peach with 
age; spadix 6-13 cm Ion 1 in nt - n i t h i| 1 nl 3-4 cm long, fertile 
staminate region 5-7 cm long, sterile staminate region 3 4 cm long, fertile pistillate m 
gion 3.5-5cm long,f)iMilLimlf. • 1 1 • 1 hi id I 1 >l turn D urns ovoid, yellow,7- 10 

Native to the neotiopi * ml nt M 1 1 1 Central and South America and the West 
Indies since pre-Columbian times (Vaugha itroduced into the 

United States as an of urn nt 4 n m - 4ti t tl ers and fruit (Bailey 

1997;Vaughan&Ge . " ' ed throughout cen- 

ongate, oblong, subterranean tubers 

tead they possess a network of fleshy 

through vegetative propagation by rhizomes 1 dim p. t im s,zpd colonies of imma- 
ture plants at the ihmmu 1 1 1 n tt 1 1 1 f I m 1 r likely arise through 
vegetative fragmentation and subsequent rein ti u t 1 tin m. fmgments.Dispersal 
and establishment from seed is also possible bin Do-, b I liD u ih fruit production is 
not often observed, plants are self compatible, wit h.i sum In mdividual or colony produc- 
ing fruit and viable seed. It can bo difficult to determine whether new populations origi- 
nated from seedlings or from vegetative plant propagules. Vegetative expansion seems 
a more plausible e am the similar X. 
atrovirens C. Koch and Bouche. 

content. It is generalK i m I it 1 u ^ 1 i 1 , 4 t 4 c ts;along roadside 

ditches, pond and lake margins fit iin i it in i It I it t ilml it n In o > I < i 
sonably wet sites, woodland edges,old fields, vacant city lots, roadsides,and home sites. 
Although X. sagittifolium prefers soil condition , th ii nernl- it« y hydrated to highly 
saturated, it will grow on mesic slopes and othet Int'tii th ll-dtained soils, as long 
as adeguate moisten i pn wit t i hi Imhnh it md mesic sites in Ocala 
andBellview,Flonda ii 1 ' " jtnl ' ; : "' ' has minimal cold toler- 

ing Juvenile, unestablished plants ate most si iscep I bio to dee/tug because older plants 
can regenerate from : ul lh tul it u I it I Ut uqhoutthe Ameri- 

can tropics and to a l< m mmi t I mil in it wl'd ' nle\ 1 997;Vaughan & 
Geissler 1997; Neal 1991). The tubers are similar in chemical composition to those of 
Colocasia orAlocasia, but the starch is mote dilb uh i . I ], i mahan & Geissler 1997). 
In West Aftica the tul . t n pmt- i - I id -t u the production of the 

loud substam e known as fufu (Vaughan & Geissler 1 997). 

Xanthosomasaqitt i i il mi i I inn i | n twomploht I 

through tuber divisi it tin n i i f - I i it ml > in itumi plantlet removal, 
and crown division m ill pi miln h I in, - mil , h it plant at the 2-3 leaf 

stage so long as th i i t II >t \anthosoma are 

I itmvmlv Mil itetmnoaml von ma into individual', tta implant ten lilv. 

) ^ 'it > b i' I | m a it I , h nt i m 1 malanga, tannta, 


2.Xanthosoma violaceum Schott, liul. Sent. I b >t t . Herol. 370. 1853; Schott in Oest.Bot. 

1 1 


150 en 

a in height, from i 



ale or yellow 

green, tube, • 



am lex p. -orly developed o 

r absent in juvenil< 


i- , i ,(., 1 il 


t to 1 m or more 

in length, 

elongate, slender,' 

without scaly 

covering of c 


t tuberous at vari. 

hi' places 

along the length/ 

white and of- 

ten tinged v 

oth light qmen 

or pink coloratic 

)ti; leaves : 

10-100 cm long ai 

ad 15-75 cm 

wide, ova let 






i Im II m 

mate to hast 


h lob« 

? distally obtuse 1 

:o acute ar 

id unequal, margir 

i! 1, bul 

broadly undulate ore 


baxial surface usl 


ius;primary lateral 

veins 4-9 per 

side, purple - 

or rarely 

a pale 

yellow green, mi 

dvem and 

lateral veins sunk* 

n beio\ tin 



nt Ii u ml | 


times yellow-greet 

i.oldei leaves 

dark green, y 




/ith purple or yellowish primary 

it wcondai 


cenceof 1-nume 

sous spadt 

ces, peduncles 30 

SO cm lotiq, 

cm long, initially purple but with age becoming grey or yellowish, appendix absent, fer- 
tile staminate region 10-15 cm long, sterile staminate region 3-5 cm long, fertile pistil- 
late region 3-5 cm long, pistillate flowers with slight odor of ether;berries ovoid, yellow, 

Cultivated in southern Florida but not strongly naturalized. Plants have been ob- 
served spreading vegetatively at sites of culti. ii n ' h! . ntm 1 ,hibhshment outside 
in i I mi i p I I \antn ill it I (htoughout the trop- 

ics for its edible tubers similar to X. sagittifolium.Jhe exact area of origin (possibly the 

similar to that of X. sagittifolium. 

I would like to sincerely thankThomas B.Croat al In i lis ;i iuri Bi >1 inical Garden for his 
critical review and insightful suggestions on the content of this work. A high degree of 
gratitude is also in order for John R.MacDonald at Mississippi State University in Starkville, 
MS and Richard J. Carter at Valdosta State University in Valdosta, GA, for their editorial 


oTricia K. Blak 

;e,Terri Ballinger,a 







1988 (1990). The ecology and life lotn i >t A t ^ ,, t n.icana 1 1(3-4):4- 

ngler, A. and K. Krause. 1 920. Araceae In: A. Engler: Das Pflanzenreich, 4:23, A-F, Leic 

renchJ.C, M.G.Chung, and Y.K.Hur.l 995. Chloroplast DNA phylogeny of the Ariflorae 

P.J. Rudall, P.J. Cribb, D.F. Cutler, and C.J. Humphries, eds. Monocotyledons: Systema 
Royal Botanical Gardens, Kew. Pp. 277-286. 
Srayi m, M.H. 1984. Palynology and phylogeny of the Araceae. Ph.D. dissertation. Univ 

In i , AT. 1 939. The nomenclature of the taro and its varieties. Bot. Mus. Leafl. 7(7). Ce 

4AYO,SJ.,J.BoGNER,and P.C.B >yce. 1 997. Genera of Araceae. Royal Botanical Gardens, Ke 
1 995. The Arales. In: P.J. Rudall, P.J. Cribb, D.F. Cutler, and C.J. Humphries, e 

Monocotyledons: Systematics and evolution. Royal Botanical Gardens, Kew. Pp. 2/ 

Ieal,M.C. 1991. In gardens of Hawaii. Bernice Pauahi Bishop Museum, Honolulu, HI. 

h n Ju andC Ad i I" ' II - till III I Moi I Urn < i «U 

Press, Inc., New York, NY. 
'oung, R.A. 1946. I he I,Im , , -nil, , t , , ,t i .f. fur home use and market. USDA 


Jose Guadalupe Martinez-Avalos Humberto Suzan-Azpiri 

Institute) de EcologiayAlimentos Escuela de Biologia 

'i'"> i<l !'!•', ieTamaulipas Universidad Autonoma de Queretaro 

I 3 Blvd. I opez Mateos 928 Cerro de 1 1 < 

-■'CO 87040 Querelato" ■ 

[ he geographic di tnt i n n t i h t i i I h hi \\\k, Mexico was deter 

mined based on litu. nun huldniin i I I mi i \ ml. ti nul . ial iif li tin inN" 
and 1992.Three specie mdlw< ul pi i u I i led \notarpusagavoi 

T t i u ' | ipub agavoldes is extr 

geographicallyoccupyinq jnarea c lh n 1 n tin main factor responsible for 
soil type, which has c lav iH-aur^ mih nit i i-nt and a slightly alkalir 

retusus h,i th In ih i t r i th I amh. iniiii I poiti 

probably because c I, i such as vegetatio 


geografica del genero Ariocarpus e 

ti - 1 ' t n il l-_ Toii ml | i ' " ■ 

en revisiones bibliograficas, consultas de herbanos, 

de campo realizados en 1991 y 1992.Se encontraror 

; agavoldes, A. kotschoubeyanus, A. retusus subsp. re, 

lyor a 2 km- ;el factor responsable de su distribucion es e 

• ■! 

retusus subsp. trigonus presenta el mayor rango de di: 

ti 1 u Mm ^ upandolasregiones 



^"— u - 

The genus Ariocarpus described by S< hoidwoiler in 1838 i 

Mitich&Bruhn 1977),is widely 


■d in the Chihuahuan Desert (Anderson 1 960).Th 

is desert is located in the Mexi- 

can highl 

ands that includes portions of the states of Texa 

s (U.S.A.) and Coahuila, Nuevo 

Leon, San 

Luis Potosi, Hidalgo, Queretaro, Zacatecas, and 

Tamaulipas in Mexico (Shreve 

1951; Jaeger 19' II < dn si i I , , , „ , , i,, ,, this region is k 

theTamaulipan Arid Zone (Bravo & Sanchez-Mejorada 1992) including the mi 
hes (municipios) of I ula, Palmillas, Miquihuana, Bustamante and Jaumave (Fig. 1 
The genus Ariocarpus includes seven species: Ariocarpus agavoldes (Castar 
Anderson, A. bravoanus H. Hernandez & E.F. Andetson ' | 

uratus (Engelmann) K. Schumann, A. retusus Scheidweiler, Ariocarpus 
ind A. scaphirostris Boedeker (Hunt 1 992; Hernandez & Godinez 1 994). 
i recent revision of the genus Anderson and Fitz Maurice (1998) added 
two new subspecies, A bravoanus H.Hernandez & E.F.Anderson subsp./i/nton//' E.F.Ander- 
son & Fitz Maurice,and A.retusus K.Schumann (Weber) Scheidweiler based 

i m 1 1 Inl ii ,1 mit , in l i ! - li i i , t i,i j j, [ i i i. I In ,ii 

or endangered (Anonymous 1991; CITES 1992; UICN 1985;Vovides 1988). The biggest 

of natural vegetation in large areas, overgrazing, highway construction, development of 
urban areas and over-collection of wild populations by commercial and amateur collec- 
tors for sale as ornamental plants i an hi : Mi jorada 1982, 1987; Hernandez & Godinez 

The literature published on ihe Menus includes the works by Anderson (1958, 1960, 
1 961 , 1 962, 1 963, 1 964); Bravo (1 978), Bravo and Sanchez-Mejorada (1 992) and Sanchez- 

Only one species of the genus have been studied with an ecological perspective 
{Ariocarpus trigonus) by Suzan et al. (1 989) and Martinez et al. (1993). 

InTamaulipas I ' a ih ji n i' im ,,- ,i L . ' of these species 

because no floristic inventories and demographic studies necessaries for the definition 
of the real status of each species exists to date (Vazquez-Yanes 1979). The main objec- 

Ariocarpus in the state of Tamaulipas, Mexico. 

The state of Tamaulipas is located al the nuithoastern portion of Mexico, bordered by 
U.S.A. to the north, the States of m 1 1 ui I i ml oim t tl ,< -nth, Gulf of Mexico 
totheeast,andStateof Nue c) l« mi th t n ^ t i Ii i , regions according 

north region with semi-arid,set 

ni-hot elur 

dth little annual r, 

ainfall (BS1 hw);2)the 

southeast region with hotsub- 

-humid or r 


climates with sur 

nmer rainfall (Awo); 3) 

the south-west region located 

in the Sien 

'a Madre Oriental with c 

:limates ranging from 

semi-hot sub-humid climates (, 



emi-hot humid cl 


an altitude gradient on the east 

:crnslepe, t 


e semi-hot subhu 


semi-hot Bsohwjn a\"\ altitude qiadioni en I 

;he we 

stern slope (Secre 

taria de Programacidn 

y Presupuesto (SPP) 1985). 

The vegetation is domina 

ted by the 


-forest (north cen 

tral region), the xero- 

phytic shrubland (south-. e, in ml 


al deciduous forests (southeast region). 

In the highlands of the Sierra N 

la in 'ii • 


te-Oak and Cloud forests are the domi- 

nant vegetation types i Table 1 

(M M .111 


. 1964; Rzedowski 


Data were gathered from v 



herbaria: DS,GH,IBUG, 

BSohw 900-1 25( 

BSIhw.Bsohw 600-900 

(A)C(wo) 1100-1300 


)SOi!;/q - Soionc-sk gley'c; 3sorw 

Living specimens 

of Ariocarpus 

were studied in t 

Botanico de la Univer: 

iidad Nacionc 

)l Autonoma de M 

Cactaceas del Institute 

de Ecologia 

y Alimentos (UAT); 

Tecnoldgico de Ciudad 

Victoria, Tama 

ulipas (ITCV). Field \ 

the state, in order to vis 

it reported site 


InTamaulipas only three 

species and tv 

vo subspecies of the 

agavoides,A.kotscl mi,i /anus i < ib | . uJSdndA.retusussubsp. trigone 

tribution of the species is expressed in Fig. 1 .The geographic distribution and chare 

1 . Ariocarpus agavoides (Castaheda) E.F. Anderson 

Common name. — "Magueyito" 

Geographic di i tnhi i >> n I u t- t i w lTimmh| \ \\ , species is end 

to the valley of Tula, municipalit luhl mh 1-, The first population reportec 
located in the north slopes of the . ity of tula, i lowever, a new locality was discov 
within the valley recently. Anot i>i , i<< ;- inhabn -in ill hills with medium si 
and easily erodable rocky soils i u Inn I Mtom 900 m to 1200 m.The as 

ated vegetation in the populiti local iorihofTula ere dominated by Pro 
glandulosa, Koeberlinia spine > 1 , \< n 1 I >i the n I 1I 

dominant species were Hechi i '< < heguilla and Agave striata. 

I h I ' 'h-,-1 n in n I pi! 1 1 1 1 1 1 n I - t > run e [ I nit 1 1 in 
and pollination dependent on bees. The type locality is extremely perturbed by a v 
disposal and overgrazing. Signs oi plant extra, in in were detected several times. 
Cited specimens: MEXICO. Tamaulipas: Tula, 1 200 m, Castaneda s.n. (DS);Tula, 1 200 m, Cowpei 
(POM); Tula, 1200 m 1 H 1 MO NY, POM) 

Specimens examined: MEXICO. Tamaulipas: Ma, 1200 m, Bravo s.n. (MEXU); Tula, 11! 
^ l'hh , 10I 2054 (MEXU);Tula, 1240m Sent '^ ,. ', Ml n 1 ,h 1 

,•<>< I 1 tlil I J I 11 in -, J 1 I II \l) Tula POOm, Mar 

Avalos & Jimenez 0441 (UAT);Tula " it , 0146 (UAT) 

2. Ariocarpus kotschoubeyanus (Lemaire) K Schumann 

Common names. "Pczuna be venado,""Pata de venado" 

Geographic distriln,' > I im n ilio r If i .I m > .m. 5S San Luis Potosi 
Queretaro. This species is repoitn- It. )i T-u ml: i t ; . mMs hi Tula, located north 
ofTula City (1 200 m).The specie, oie im i ith poor organic material (xero 

and inhabit medium slopes in a desert shrubland (Rzedowski 
lecheguilla, A. striata, I Icchiio glome: > in rind Dasylirion longissit 

We detected a well-preserved locality (location conce c 
poses), with populations represented in different size classes. 

Cited specimens (And. i nl 1 I' MEXICO.Tamaulipas: Tula 1200 m.Albert /959(POM.US); 
Ju\a, Anderson /6/ 7 (POM, US) 

MEXICO.Tamaulipas: f uki. 1 ' 18 {MO);Ju\a, Arreola 

736 (IBUG);Tula Sche t iU M l n il > i (UAT);Tula, 1 127 m, 

/VlarAne/ /Wo.s 0437 (UAT). 

3. Ariocarpus retusus K. 5chumann subsp. retusus (Scheidweiler) E.F. Anderson & Fitz 

Common names. — "Chaute,""Chautle" 

Geographic distribution.— Tamaulipas, Coahuila ui> < Leon, San Luis Potosi y 
Zacatecas.lnTamaulipas il is common in the nuinu ipaliiipsA iVAjiiihuana, Bustamante, 
Tula and Palmillas.This species grows over hills and rocky slopes with clustered popula- 
tions at altitudes from 00 to 100 m, in i cl« <o nil I, mi h Awski 1978), domi- 
nated by Agave lea In >i </A > •/' >ta A, '"> >n »>w "oc P fo'o/wm and Kucca 

Populations oAAe/uoo subsp.retuuis inhabit also peiturbiA areas dominated by 
L\)doneuu io i in n ill in n lib li in i in i itb o io subsp.trigonus 

were found in eastern slopes of I I ' i I I |l i I > r f ulations were also de- 

tected in pinyon pun On i AmnnmdA ' < lh , ] rinus cembroides in 

(il. |.ii,.-. MEXICO.T.inwuhfJ i \owper 1889, 1958 

(POM, US);Tula, 1 200 m .Anderson 1964 (POM, US). 

Specimens examined:MEXICO. Tamaulipas: Gi, tin ■■ 1 ■ ■■ ;< z etal. 2029 (MEXU); 

Bustamante 1590 m < o < ' I U i i t , i u i 100 in i/menez 0018 (UAT); 

Miquihuana 1524 m <i - ■ m ' H lu m 11 in c n Mvo/os 0/5/ (UAT); 

lula 1 m " ,' -i - i (/ >i 0664 (UAT). 

4. Ariocarpus retusus KO>( homann uAp trigonus (Webet) I AAndersonbl it/Maunco 

Common name.— "Chaute,""Chautle" 

Geographic distiit I niiuh| n i i i I i | i i li nl ut i i 
the municipalities of Jaumave, Palmillas, San Cailos, Villa do ("asas and Llera de Canales. 
The populations inhal it i'h i Inn Sum* Anoi tt< i nt I types The centre of 

I lb A ■ i in n ii b j|i in il ilu n hi i in p - i dandeioded sob A up i \ 
in the Jaumave ValA n li o I i I ub | \ u ,ti i A \ , m i i u« I. 35 000 individuals. 
The plants exhibited «i non-exclusive oh tus-nuisp plant iplaiionship.The must impor- 
tant nurses worn Po - indit' ' I I it A iv ith healthy nurses 
exhibited better conditions than those with damaged nurses (and with smaller cano- 
pies). A.lriqontr, plants arc able to Ihoinio ipgululein open oin but individuals benealh 
the shade of tun ibited better t in rogulatioi . i i | il li hi I data 
Cited specimens (Anderson 1964): MEXICO.Tamaulipas: I lun i I 60 m, Anderson 1580 
(POM, US, NY, MO, GH I in b n \h l i WO I IS), E of San Vicente, 
Jaumave Valley 760 m 1 , - m o s" , , m ,n Ol Aitmn 00 m, Anderson 1 153, 

.','EXICO. Tamaulipas: 

ofTula, Sdnchez-Mejorada 2070 (ME 4 mi i n h rWezefa/.2038(MEXU);Jaumave,75O 

m, Hernandez 2047 (MEXU); Valle de Jaumave, 61 m, Martinez-Avalos 068 (UAT);Vi!la de Casas, 304 
m, Martfnez-Avalos 0383 (UAT); San i j I ' h , \ ilos 541 (UAT); San Carlos, 400 m, 

Martinez-Avalos 0745 (UAT). 

It's difficult to determine the ecological requital >j t t h ■ . 1 1 ,ifL>< t the geographic distri- 
bution of the genus Ariocarpus with the analysis of herbarium specimens, and even with 
field studies in the Chihuahuan Desert in Mexico (Anderson 1 958, 1 961 ). In Tamaulipas 
this genus is restricted to the Chihuahuan Desert region, the Tamaulipan arid zone 
(Tamaulipas biotic province) and some specific tropical neighboring areas with simitar 
climatic parameters (i.e. LLera Valley) (Fig. l).The area in Tamaulipas where Ariocarpus 
grows is similar lubii ii i h n< jl ring states of Nuevo Leon and San Luis Potosi. 
The genus is distributed in altitudinal gradients from 200 to 2100 m (Table 1 ). Of 

central and southern parts of the state. Ariocarpus agavoides has the most restricted dis- 

direct and indirect factors such a I il it t I ti i I i u\ i incineration, marked 

1 992; Sanchez-Mejorada 1987). Ariocarpus kotschoubeyanus is a difficult species to find 
in the field due to its size and form; therefore, the true size of wild populations is rela- 
tively unknown Am i , n I i i n ui ve j stern arid zones of 

l H I III III I I II "Li I l I I' I I Urill I ' 'I II 


:=^ F i[ , 

nonimous, 1 991 . Diario oficial De La Federaoon, (1 7 de Mayo). 1 991 . Listado de esp 

raras, amenazadas en peligro de extim 

endemismos en la Republica Mexicana.SEDESOL.Mexico. 
ndi rsun, E.F. 1 958. En busca de Ariocarpus.. Cact. Sue. Mex. 3;79-84. 
1960 A t in t in taceae). I. The status of the proposed c 

/?05eo-cacfu5. Amer. J. Bot. 47:582-587. 

Neogomesia. Amer. J. Bot. 49:61 5-622. 

. I he status of the proposed genus 
III. The status of the proposed ge- 

nus Roseocactus. Ainei. J. Bot. 50:72-1 / •! I . 

1964. A revision of Ariocaipus (( at tat eae).IV. I ormal taxonomy c 

n us Ariocarpus. Amer. J. Bot. 5 1:1 44-1 SO. 

Bravo, H 197S , , , 

. 5anchi/-Mi iokada. 1992. Las Cataceas de Mexio 
a de Mexico. Mexico. 2:252-263. 
Cuts. 1992. Convention on International Irade in Endangered Sp> 

Flora. (11 ofJuneof!992),CITES. 
G i [ in,, i M ' i Ik k i m< al a ,t< m i de lasili a ion an 

adaptarlo a las condiciones de la Republica mexicana). Cartes 
Herman , H and H. Godinf/. 1 994. Contribuoon al conocimi 

mexicanas amenazadas.Acta Bot. Mex.26:33-52. 
Hernandez,! 1992. Asp-" to I nl if i I- \nocarpus agovoides 

son. Cact. Sue. Mex. 37:40-45. 


MARROQUlN,J.S.,G.BoniA,R.VA.-'oino,and I a Cia a. 1964. [studio ecoldgico 

delaszonasanda ,d 1 1 it t ' i > i i tin i M i n ilL In i tuj k ioi 

Publicacidn especial. No. 2 Mexico. 
MartInfz-AvalosJ.GJH.Si an, and ( Saoa \i 1 993.Aspectos ecolbgicosy der 

'■ i ', , a" a ;afi'ai iWobei) '.( humann.C an t. in . Mi • A M >0 
MnioH.L.and J.Bruiim. IS//. I he genus A/a a o/ . Succ. J. 49:1 22-127. 
RzedowskiJ. 1978. La vegetacidn de Mexico. Ld.Limusa, Mexico., 1 1. PAAAinlorma sol no la Reunion de Tucson paraanafc 

de Cactaceas. Cact. Sue. Mex. 27:90-95 
1987. Observation sobre ei Lstaolo de conservacidn de d 

amenazadas del notesto de Mexico. ( a( t. Sue. Mex. 32:61 -71 . 
Secrfiaria de Procramauon y Prisuruisio (SPP). 1 985. Sfntesis geografica c 

Tamaulipas.lnstituto Naeional de I sludistic a,( leoqrafi'a e Informatica. 
Shrlve, F. 1951. Vegetal ion ol the Sonoian beset t. Carnegie Inst, of Washir 


ovides, A. 1988. Relacion de pi 
Conservacion en Mexico: Si'nt 

d Marty Casstevens 1 996. Wildflowers of the Southern Appalachians: How 

i and Identify Them. (ISBN 0-89587-143-2,pbk.).John F.Blair Pub- 

h ,hi I iiu PLi aDnve,Winston-salem,NC:nn',uM, , llS i 4 < 1)0 768-9194 fax). 
$26.95.257 pp. Color photos. 


.val leaturesthat make ' wbl v 

vorth the money. 1 

r ■ 1 r, lliM us .s- ' 'nipr. is nsi.' 

guide to herb 

i 1 -tl -i .hi] ,| .i 

ifk (leotiMohic region. The southern App; 


•oin, ties , ul ill t 1 ii i .l|i 1 tth< 

:es, with over 2500 spec 


induces sectionsolVrginiaAVi. 

;y, Tennessee, North Car 

(Aiolma, Akit 

.ama, and Georgia. However, sir 

ice many of the 11 

30 species in this book 

have wide 

ge< 'graphic d 

d important feature of the boc 

A ,s the high quai, 

iv of the photographs, 

I | 1 1 it i| h 1 ill 1 Ii i i in Ii i 1 u id hit- 

■ i r i a i iiour i 

Ph ' . ' >nh 

-up lens. The 28-page 


aken from the main se> 

st ion of tiie 

also included photogr. 

iphsol the 

..) hi in- m it. 

rn « i >, tiie author, 

using the equipm 

ent), since many amate 

sir piiotog 

en-page chapter < 

i additional 

.. .. (1 |S1 P 

1 [ 1 llMtl) ||f ,h i All III' ' ' ■ 

hhotouraphv kdt< 

?gories," which corresc 

mnd to the 

useful photo 

1 1 idk! ." 1 1. . M, lllb H pb, 

A third important feature is ihe empha 1 

i.The authors strive tot 

educate th<> 

pi.l i .1 nit i ill i ih i i i r ,i ii pi m is tii i> 

st environmental polio 

dike book 

eis on "The ,ottir 

ip iotanual Kploration ot lh 

e Southern Appal; 


sental Con- 


iois enciiuracic pooplo to phots 

ograph rare flower 

s, uthei ''sin ps k them iheir pho 

urbance to flowers and their habitats, ats 

,r ,. Ti ,,, II. 

ivveis should not even be photoqiaphod if intius 

ion into the habitat is t 



Billie L.Turner 

the sole member of section Plateilema. The latter was maintained as a genus by subse- 
quent workers (eg.Rydberg 1914), and I intend to accept its generic status in my forth- 
coming treatment of the Helenieae for Mexico (Turner 1 999, in prep.). 

The single species concerned, Plateilema palmeri (A. Gray) Cockerel!, acaulescent 
perennial herb superficially resembling Gaillardia comosa A.Gray has been largely ig- 
nored by most phyletic workers (eg. Bierner and Jansen 1 998), presumably because it is 
represented in herbaria by relatively few collections, mostly type material, the latter ob- 
tained from near 5altillo,Coahuila, Mexico. Indeed, prior to the present report, in addition 
to the type, the species was known by only a few skimpy collections from the Ocampo 
region of central Coahuila. 

States, 200 kilometers or more from the previously known collections (Fig. 1 ), as follows: 
TEXAS. Brewster Co.: "Infrequent at Scnulcr mail box 44 mi S of Alpine, ierlingua Road; altitude 
3,600 ft," 10 Apr 1 - • I'm, , t ,n ,, 1M d | ,| Chalk Draw; Schuler 

.( ' it h i i ,|tit , | t ^(SRSC) 

Henry T Fletcher (1884-1955) was a remarkable man in the Brewster County region, 
serving his community in many ways:Vice President of the State National bank in Alpine, 
RotaryClubGovernor,ownerand manager of the Fletcher Cattle Company,to name but 
a few of his enterprises (Casey 1976). He was also an excellent collector and keen ob- 
server of range land plants,as attested to by his rarely cited but excellent publication for 
the times, Vegetation of the Green Valley Region (Fletcher 1 928), a 40 km ; of rolling plains 

the collections on file at SRSC, I calculate that he assembled and placed on deposit at 
that institution five thousand or more plant specimens. And his are not fragmentary 
collections as often collected by non botanists, rather they are neatly pressed, solidly 
mounted and in th> I t t I I' nn j uij fiuitincf ■ C i n short, Fletcher 
collections served as the founding core of the SRSC herbarium, this subsequently built 
upon by the late Barton H.Warnock (Turner 1998). 

Both of the above specimens were said to have been collected on the same date, 
but possess very different collection numbers (2/9and 884),suggesting that the collec- 

profusion of collectable s 

eleven heavy early rains i 

s magnitude.) 

No eollection l)ook 01 tecoids survive lo sort out the pe< numbers assigned 
Fletcher's specimens, but mosi likely he assembled his plnnm over several months and 
then assigned his phi t un nl i | n - i t I r lit tion.simibi uixnna 

lies as that found with Ik numbers alloc! It nth. d ivean ilso encountered with 
othei assemblages of this collector. 

; sRut 

I leprosentod I 

;e if the species might still exist. My hunch is that 
opuhtions aloi gion ofTexas. 

nfortunately, most ranchers in the nans Pectus at the present tone are reluctant to al- 
iw botanists on their lant hlant Is, fear lul I hat an nudum mi ed I axon might be observed 
ad as a result their use of their rangelands placed in some sort of jeopardy. 

een made in the .tatt >H\li h mi , ) ho > > , p , M , i i. , p ih< I lintonfumilv [Hintmi 

3 be a very localised | i u - ' I i ni i i r i ilb t rehtiveK ram 

d judge from the collections at hand. 

RillRINi. LS 

/ lym en oxys and Tetro/ia/m.LuncliTici 1:17 ,'!,, 
:asey,CB.1976.SuI Ross State University,the cultural center ofTrans-Pecos Texas. Pioneer 
Book Publishers, Seagraves, Texas. 

;ockerell, T.D.A. 1904. The North American species of Hynenoxys. Buil.Torrey Bot. Club 

letcher, H.T. 1 928. Vegetation of the Green Valley region. West Texas Hist. Sci. Soc. Publ. 

Iydblru, P.A.I 914. Plateilema, in N. Amer. Fl. 34:1 01. 1915. 
URNER, B.L 1 998. Barton H. Warnock (191 1 -1 998). PI. Sci. Bull.44:78-80. 
1 999. Helenieae, in Comps of Mexico. Vol. 5. Phytologia Memoirs (in prep.) 

Susyn Andrews, Ai an L pand( ,v, jA.i . n (eds) 1999 Taxonomyof Cultivated Plants 
(ISBN 1 900347 89, hbk.). Royal Botanic Gardens, KcwJ 27,553 Pp. 

Cultivated Plants spm i I mtl i il i I m n I I in ugli, the Royal Botanic 

Gardens, Kew, and tl I ti ul ui u hi ith icintptn ft 

foi I lor t n iiliui il l i n 'h it i in i pi I I II' I I I Mi Inn il I i-ottmm i it >i jf >) 

jinih. I I itttu h t ti ul 111 lr ii « r 1 flmhu |h ti >m i d Jul> 1998. 

papers or abstracts up i> h nPnlmh. |u|.i dt i mi I ui h Ii inbuted pnong ten 


. in II ti 



IpLnn :ii„ji 

■not plant i 



: m I, n 

techniques in I: 




Ii ' i ( uti il ill ih iii i i II ul i in ' t m lii i Mh I ilh n ii 

h >it nil ih Pit i it il I ui i ih | i| I 1 iptintniL] ul . ul1 

their origin traced to I ut ■ ' in i In i I if mi | mm m d u iii s impli d ll 

nil i | i iti i n ti I i il | i i I in I i | ul ti ltd | Ih i | ipc i in sum 

guide thu e du i lit i iti i I i i i I | m i i iu inq wild species int 

neirntttuilhui study of each uiea, making urn book 
nake the boot' well poith the prit i I unhesiPtingh 
ists.— n • i , p 

,, s , , )gp, an : Ft Worth,TX '6126 3019, USA. 


■ 1 1 - 

adequate genera! descri 




Plants are usu 

ally included in flc 

)ristic accounts of the North America, 

a flora on the bash 

of their status 

> either as native 

or as non-native but integrated into 

the flora to some 

degree. For nc 

jn-native plants, a 

in assessment of their integration is I 

aased primarily or 


status and dispers 

>ive success. The more summary florii 

itic accounts com- 

monly do noi 

: provide informa 

tion necessary for a clear assessmen 

t of the degree oi 

fioristic integi 

ationof non-nati\ 

/e plants. 

A selection of genca! staM 

?ments of intent for major fioristic sti. 

idies (from numer- 

ous examined) indu dies that a 

broad range of non-native taxa is tt 

-eated. Naturalizea 

plants are inva 

riably treated, but 

the definition ofnaturalized'is not coi 

isistent, and plants 

recognized as wait in-n nm | i i a w m i n -p i n >i b it luri I 

Radford et al. (1 968) for the Carolinasfspecies of vascular plant known to 

Correll and Johnston (1970) forTexas: "all nativeand naturalized llowtnir. i 

Voss (1972) for Michigan: "all species ... known to grow (or to have grown) 
cultivation in Michigan, whether ouginjlly native or not." See Voss (Prefao 

Great Plains Flora Association (1986) for the Great Plains: all va sculai pla 

Welsh et al. (1 993) for Utah: "All indigenous plant species known to occur in Utah are 

included in the floiu liiimiia Ipitii > < il u uuiulimttnt ,u intensively as are 

the native ones.""This flora attempts to present coverage of all established species, all 

common adventivi taxa nd man if ih mm n n i ultivated species." 

Gleason and Cronquist (1 993) for northeastern United States and adjacent Canada: 

plants "growing wild'" (including ■- ftom the text -"often escaped from cultivation," 

"adventive"and nu mud i u ml | il f -I I h 

Hickman (1993) for California: | I it i it mi ill u Ml aliens that have 

become an integral oait of the (mlilorniun flora am included. I he general policy was 
not to include (or to n u n in j i mi .nt in i i |i liu mu but loin] pi isist 

(e.g., "native to Europe") and a brie! description of it 1 , habitat, qeographic range, and 
relative abundance is piovidcd, without the use of any further terminology. 

Cooperrider (1995) for Ohio: "the native members of these familiesghe alien plants of 
these families that n . i i m t .' h if t , t h , aally in the state's 

tl - i in I t i i i ' m th< f a nil no i lit it i | I n I 

Flora of North America Editorial Committee (1996) for North America north of 
Mexico: to be include! I in lull < ire "all native plant s"and "in i;odu.:ed taxa that are natu- 
ralized or found frequ in ut if t iti ii t. . ii,ei,i ii I m the discussion are 

important or extensi I ultivat I plant thai tend not to i ipe cultivation (e.g 

Diggsetal. (1999) for north central Texas: "al II o .. na t ind naturalized vascular 
plant species. Ate i if ' it < r i ' - lit i imon fig), but appar- 

ently non-reproductive taxa have been im ludeil bee-vase of the likelihood of them 
being encountered." 

Kartesz(1999) for North America north of Mexico: "a I known native, naturalized or 

plicitly, are non-natm ta i mcognoed i | i iiit i" (as described below). 
Despite the demonstration in the recent lepson Manual (I lickman 1 993) that the floris- 
tic integration of non native plants cm be pieciseiy iiesculxm without specialized ter- 
minology, a minimal set ol descriptive teims is useful if consistently applied. The four 
terms below pios id il nt i i c il ' i<\ ' n of the major categories of non- 

should be distinguished from naturally occurring plants.Anotherdistinctioncan be drawn 
by recognizing a cultigen (a cultivated plant of unknown or obscure origin) as different 
from a cultivar (a cultivated plant of known origin). 
2. Persisting 

Perennial plants (woody, suffrutescent, and lot I sn . ,u , I lor ornament or in- 

m presence is det « mh hi jone. Such plants are not repro- 

ducing or at least not spreading beyond the original planting. The difference between 
"cultivated"and pei isting prim,) i i ne,telonin notion of exist 

ence,and persisting plants usually are not in .'udcd in floristic accounts with the same 
status of native and naturalized m e n i in i en i they grow outside 

of cultivation, they may appear i n i ii ion no i h tin y may be explicitly 

i flons 


Plants ot non-nativ* spM it ] i i i I in u ion but not maintaining a viable 
population for more than one oi O i -i ml t i it O in the North Ameri- 

can flora are known only as single hisioni n^ ords; other essentially non-reproductive 
plants as waifs apparently may be recurrent. Some species reported with new distribu- 
tion records may become known more precisely as waifs as the site of their reported 
occurrence is reinvestigated.More ihan ,i single season is reguired toobservethe repro- 
ductive status of such a plant or population, and some of those initially suspected of 
being waifs may become natuialm I 'I n i o nnato from various kinds of natural 
and human-mediated long-distatH - J j i h II,- , i nav originate from seeds washed 
out from garden plantings and > > n i ul w fool at sites where seeds are mixed 
with other exotic materials (e.g., ballast dumps, wool mills). Tomato seeds, which pass 
unharmed through I i n i h I 1 n i m ii |i i I in sludge-based 

fertilizer and may spawn waif plants in repeated cycles. The usage here for "waif" is 
a good mate h it i it mi e - i [ i | ^ in 

I. Naturalized 

dants of non-native species accidentally cji ot hi , i iio t,i m iihtotnetL 
r I f i n i it n i ii ii hi ] -not lot i up f ' • ii r ,. at 'more than 

nr a few seasons), and dispersing without deliberate human assistance beyond th 
ation or populations of original establishment. The degree of naturalization n 
rom widespread and abundant occurrence to local establishment (from rece 
Juftion or sluggish reproduction and dispersal). Perhaps the mostoii > o 1 ' O in: 
ion are cultivated plants spreading slightly beyond their original planting — thes 
)e best characterized as waifs, if their reproductive ability is slight [<'.<.}., Iivpatic 

or they may be "incipiently naturalized" if reprod 

uction and spreac 

i appear to be vigor- 

us,evon though testru ted in atea.Speci rati to N )ith m i 

a may become natu- 

ralized in areas of the continent other than where 

recorded as native 

in the past or present. 

Some species apparently have a mixture of native 

! and non-native N 

orth American popu- 

lations {e.g., Achillea millefolium, Galium aparine, i 

and Prunella vulga 


Associated Terms 

in addition to tin 1 lour mine, suggested above to covet the major i 


tion of species and other taxa in the North Amc- 

mean flora, vanou 

s other related terms 

have commonly been used in floristic works to 

characterize thee 

rigin and integration 

of such plants. The usage of these terms varies 

considerably, and 

without other refer- 

ences that provide comparative definitions in a 

floristic context,! 

t is hoped that those 

stent application.' 

'Native species,""alien 

species," and "introduction" are formally defined 

in the recent Exe 

cutive Order (U.S. Ex- 

ecutive Order 1311 2). Because of the difficulty in 

of floristic criteria, they are listed here alphabetic 


Adventive— non-native, becoming naturalized 


rate plantings or acci- 

dental introduction. 

Alien— non-native,commonly considered to ha* 

ve originated from 

a different continent, 

but not necessarily exclusive of an origin from 

an ecologically d 

ifferent region of the 

same continent. 

Escaped— non nati - be oming nututaliz. 1 I'-cmii dehbetah nl mtings, usually only in 

a local area or represented by relatively few inch 

yiduals over the ra 

nge of naturalization. 

Established- non-native, similar to "adventive" 

and "introduced" 

but with the implica- 

tion of being more securely naturalized. 

Exotic— non-native, essentially similar to the tci 

-m "alien," but with 

i stronger implication 

of an extra-continental origin. In the recently issue 

i n iii Mi, l h M i il i ml 

p.,' | i "il , . Ml ,| i i I Hi tin . IMII ' Hi il M 'I i il Mil I l d I 

r continents or from Mexico or Central America, 
lative, released i i h I Mil e I n t th tl n I i 

on, at least forone denotation. Similai to"advonhve"but with dightlv 
,[ t, in | oidin il d tin >ugh i I mt u Hoi hot tic oi piac 

latmg ttom out ou the geogiaj In wgiori of cow.em,cithei (torn a 
nfiotii no >lo ii ill ft t nt ii i itth in ontinent.Among 
ten used interch i i y II I it dn u < ■ m uti > I - 1 non 
is the most general, at least in the definitions suggested here. 


[ ] Won id w I d I i it i I i t i 'h u d| I htii I 

Mou.-on I end ot the Univetsitv ot North i aionn.i i letbjtium I.NK Ji. 

i Order 13112. 1999 [3 February]. Executive Order on invasive alien species, 
on, The White House. [ 
'1 999/2/3/ 14.text.2] 
O iopf rridi r,T.S. 1 995.The Dicotyledoneae of Ohio. Part 2: Linaceae through Campanulaceae. 

Ohio State Univ. Press, Columbus. 
Correll, D.S. and M.C.Johnston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner. 
Dices, G.M., Jr., B.L. Lipscomb, and R.J. OKennon. 1 999. Shinners & Mahler's illustrated flora of 

North Central Texas. Sida, Bot. Misc. No. 1 6. 
FioRAor North Avtii Ernn-C in, 1 996. Flora of North America North of Mexico: 

Guide for contributors (August 1 996 version). FNANM Organizational Center, Missouri 

Botanical Garden, St. Louis. 
Gleason, H.A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United 

States and adjacent Canada (ed. 2). The New York Botanical Garden, Bronx. 
Grfat Plains Flora Association. 1 986. Flora of the Great Plains. Univ. Press of Kansas, Lawrence. 
Hickman, J.C(ed.).1993.TheJepson manual: Higher plants of California. Univ. of California 

Press, Berkeley. 
KarteszJ.T 1 999. A synonymized checklist and atlas with biological attributes for the vas- 

cularfloraofthe United States, Canada,and Greenland. First Edition. ln:Kartesz,J.T,and 

Botanical Garden, Chapel Hill. 
Radi ord, A.E., H.E. Am fs, and OR. Bell. 1 968. Manual of the vascular flora of the Carolinas. 

University of North Carolina Press, Chapel Hill. 
Voss, E.G. 1 972. Michigan flora. Part I, Gymnosperms and Monocots. Cranbrook Inst. Sci 

ence Bull. 55 and Univ. of Michigan Herbarium, Ann Arbor. 
Voss, E.G. 1 996. Michigan flora. Part III, Dicots (Pyrolaceae-Compositae). Cranbrook Inst. 

4W.Simmonds and J.Smarh (chapter^ Mill im m. Spoor) 1999 Principles of Crop 

Improvement. (ISBNO iO' nn >' o |,[ | ||,| ,i" , , , | td ^50 Main Street, 
'148 5018 U.n u l i ta< wwwblackwell- 

51 04.95, hbk.41 2 pp. Illustrated. 

hi It pi. enl 1 n -i li ii t I 'ii i i nil mil i mini ll tudu 
tion with recent and puss liie latum trends in breeding, the text nas 

is a new chaptei on! in I i mi h 1 i mil n the use a I t onservation of genetic 

resources.Throughout the book examples are duu n tt m id i in If nul iHk .i d 
flow diagrams are used to supplem it i I m in i i i hi te-t lin hoof is 

aimed primarily at senim undemraduate and uiadualr students <>l mini ullure, horticulture and 
applied biologv ontentsO) 2) t breeding; 3) Objec- 

tives of plant breedin i 4) ti i | ' F | il it i i I n i 5) heeding plans;6) Trials 

and Multiplication; 7) Disease msislance; 8) Spec ial tm (ungues; 9) Kmtec hnology and crop im- 
I . 1 i 10) . . 11) ■ b ography, Index of 

Epwaki> A.Wiiss (Fore <wui< I hv Sii t I miles iVreita,f Ks). 000 Oilseed Crops, Second Edi- 
tion. ' eet,Malden,MA 

i i ] - 1 1 1 1 \ - i i ii| 1 - In " ). 
$156.95, hbk. 364 pp. Illustrated. 

d stinflnweu lambcMiniet and joioba A final c haptet 
sand the gteat variety of prodm is and usesfoi oilsee 
written to include the latest agricultural research, th 


David M. Brandenburg John W.Thieret 

£ awes Arboretum Department of Biological Sciences 

iiuihinnt!!leicjhi\KY-1!',)9^ f/.\A 
A decade after Clayton and Renvoize (1986) suggested that Limnodea L.H. Dewey is "a 
prairie version of Cinna, "Tucker (1996) redu< < it,, I t i n ..jn I gtass taxon to syn- 
onymy under Cinna L. As the only iu ,tiA iimn to> !.h Diancji h> noted that Limnodea 

i length of lemma awn, a minor 
5 being unsupportable. A sunn, it 
uction of Limnodea to synonymy occurred in 1 841 when Trinius in Steudel (Steudel 
1) transferred it to / ' i i l i it i nus of two species from which Limnodea 
=rs in several fcatun s BmiiDm I - > )n the basis of the lemma awn of I imnodea 
ie, the u 'in. could just as well be ill - A 

Our purpose in this note is to point out that, in addition to features of the awn, 
'e are indeed other compelling differences between Limnodt /and • '<; and thai 
sidering these two taxa to be < ao mm it' nil- i t' ttuuately the new com- 



dy been , 

.ed in t, ) t. mi II. a i ' und< dm I 1 ) i it 1 1 




-1836) recogni 


, he had in hand constituted a distinct 




h he c 

enia I 

Mutt. Unfortun, 

ately, this was a preoccupied name, as 


?re t\ 

her generic nai 

ater applied to 

tli imim klerachneJoney in Trinius 



; 184' 

I) and 


' Bentham (Bentharr 

i 1 It htnll D< - M u t ul h h 1 


1 gcnwic 


:h has hitherto 

n i ill H i 1 i in 1 jll |i ji A , 1 M - I ii 





md a half Z./mn< 

oT m has been recognized as dishw I bv 




other be 

ts (except liter 

cer and the two authors of floras cited 

above) who have dealt with it; we list some of these here (e.g., Allen 1980; Beal 1896; 
Beetle 1977;Bentham 1881;Benth m I I -.1 i ' D w, 1929;Campbell 1985;Clayton 
& Renvoize 1 986; Correl & Johnston 1970; Dewey 1894; Diggs et al. 1999; Dore 1956; 
Featherly 1 946; Gould 1 968, 1 975, 1 979; Gould & Box 1 965; Gould & Shaw 1 983; Great 
Plains Flora Association 1 977, 1 986; Hackel 1 887, 1 890; Hatch et al. 1 999; Hitchcock 1 920, 
1 935, 1 937, 1 951 Johnston 1 990;Kartesz 1 994; Kartesz & Meacham 1 999;Lamson-Scribner 
1900a, 1900b; Lonard 1993; Powell 1994, Dunn i 197? ilyeu 19 E B Smith 1994; J.P. 
Smith 1981 ;Stebbins&Crampton 1961 ;Terrell 1971 /asey18: l£ i;Watson & Dallwitz 
1 992). The genus Limnodea is als> i i I t in E t mical Garden's New 

World Grass Checklist (<> Ac- 
cessed 1999 September 30) and in the Fe> 1 , Am' 1 m . i o ,m , Images (<http:// 

mieiil, re^tiveiv 'umi 

1-1,.. .n. J I 

Pohl&Davidse 1971) 

Cinna was recently revised by Brandenburg et al. (1991). Differences between its 
four species and Limnodea are summarized in Table 1 . 

True it is, of course, that Limnodea and Cinna share certain features. Among these 
are 1 -flowered spikelets, articulation below the glumes, extended rachilla, and awned 
spikelets (sometimes awnless in Cinna). Each of these features is found in other genera 
of Agrostideae sensu Hitchcock (1951). The combination of the four may be unigue to 
Cinna and Limnodea — it is among U.S. grasses — and may indicate a possible relation- 
ship between the two genera. Bentham and Hooker (1883) suggested a resemblance 
between Limnodea and Polypogon; Nuttall (1835-1836) considered Limnodea to be "al- 
lied to Oryzopsis." However, resemblance and possible relationship are not necessarily 
the same. Combining the two genera because of a feature of the awn is, we suggest, 
most emphatically negated by the many other features separating them. 

Distinguishing among the spikelets of the species of Cinna takes some experience, 
but distinguishing between spikelets of Cinna and those of Limnodea takes hardly more 
than a glance. As an experiment we removed the awns of several spikelets of Limnodea 
and then mixed them with a number of Cinna spikelets representing all four species of 
that genus. Next we invited several colleagues to examine the mixture under a dissec- 
tion scope to determine which spikelets'did not belong. "No one had any trouble point- 
ing out the Limnodea spikelets— even without their conspicuous awns— as the "differ- 

The genera Cinna and Limnodea are easily separable by the following synoptic key. 
1. Spikelets laterally compressed;gluni f ' , ti ' i-il i i, , h uplv keeled; lemmas awn- 
less or with a straight awn much shun, i mm i In | ill i t, | >alea well developed, 

usion of Limnodea in Cinna introduces a markedly discordant element into the 
lall and well-circumscribed genus.The four currently recognized species of Cinna 
e similar to each other and all differ consistently from the monotypic 
al significant features, especially in the strikingly different spikelet morphology, 
enera are not congeneric. 

M. 1 980. Grasses of Louisiana. Univ. Southwestern Louisiana, Lafayette. 
. 1 896. Grasses of North America. Henry Holt and Company, New York. 
.A. 1 977. Noteworthy grasses from Mexico V. Phytologia 37:31 7-407. 

BiNiHAM,G.and J.D.Hooker. 1883. Gen. Plant. 111(2). 

Biws, J.W 1929 The \* 'I n i f in i i- < u m I Company, London. 
Bowdln,W.M. 1 960. Chromosome numbers ,hk1 

Twenty-five genera. Canad. J. Bot. 38:541-557. 
Brandenburg, D.M.,W.H. Blackwell, and J.W. Thieret. 1991. Revision of the genus Cmna 

(Poaceae).Sida 14:581-586. 
Brown, W.V. 1955. A species of grass with liquid endosperm. Bull. Torrey Bot. Club 82: 

Campbfi i , C.S. 1 985. The subfamilies and tribes of Gramineae (Poaceae) in the southeast- 

CiAYTON,W.D.andSARiNvoi7F. 1986. Genera graminum. Grasses of 'the world. Her Majesty's 

Stationery Office, London. 
Correll, D.5. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas 

Research Foundation, Renner. 
Davidse, G. and R.W. Pom. 1978. Chromosome numbers of tropical American grasses 

(Gramineae). Ann. Missouri Bot. Gard. 65:637-649. 
Di wey, L.H. 1 894. Gramineae. In: J.M. Coulter. 1 891 1 894. Botany of western Texas. Contr. 

Dicgs,G.M.,Jr.,BL L vilFI O I i i mi ' ihlei s illustrated flora 

of north central I i n ill i Onmi t Texas, Fort Worth. 

Dore, W.G. 1956. Some grass genera wit h liquid endosperm. Bull, lor toy Bot. Club 83:335- 

Feathfrly, H.I. 1946. Manual of the grasses of Oklahoma. Bull. Oklahoma Agric. Mechan. 

'kansas flora." Rhodora 46:1 56-1 57. 

3k Company, New York. 
liv. Press, College Station. 

0< im oastai nd F u\ A&M Univ. Press, 
ratios. 2nd ed. Texas VO i Uni In )ll< ) 

College 43(21). 
Foster, R.C1944.The pub 


Gould, F.W. 1968. Grass sy: 

hematics. McG 

Gould, F.W. 1975.Thegras 

Gould, F.W. 1 979. A key to 

Iron Station. 
Gould, F.W. and T.W. Box. 1 

ses of Texas. Te 
965. Grasses o 

College Station. 
Gould, F.W. and R.B. Si iaw. 

,983.G,as S s yS 

Great Plains Flora Associate 

on. 1986. Florae 

I Iai km, F. I 88/. Ciiamineae (echto Gia 
Pflanzenfamilien ll(2). 

ii Plains. Univ. Press o 

Hatch,S.L,J.L.Sch ii miMD 1- ' u ?nhe Texas gulf prairies and marshes. 

Texas A&M Univ. Press, College Station. 

Hitchcock, A.S. 1 935. Manual of the grasses ofthe United States. U.S.D.A. Misc. Publ. 200. 

Hitchcock, A.S. 1 937. Limnodea. N. Am. Flora 1 7(7):538-539. 

Hitchcock, A.S. 1951. Manual oftht in, Mi- Uitcd States. 2nd ed., revised by Agnes 


Johnston, M.C.I 990.The vascular plants ofTexas.A list, up-dating the Manual ofthe vascu- 
lar plants ofTexas. Published by the author, Austin?, Texas. 

KARTEsz,J.T.1994.Asynonymizedch. Mr ,th - > uht tloia ofthe United States,Canada, 
it I it I i i hint m t -tl i I )tegon. 

CD-ROM. ISBN 1-889065-05-6. North Carolina Botanic il Garden University of North 

Carolina, Chapel Hill. 
Lamson-Scribner, F. 1 900a. American grasses- I. U.S.D.A. Div. Agrostol. Bull. 7 (3rd ed.). 
L i-S mini , F 1900b. American grasses -III. U.S.D.A. Div. Agrostol. Bull. 20 (revised). 

Lonard, R.I. 1993. Guide to grasses ofthe lower Rio Grande Valley, Texas. Univ. Texas-Pan 

American Press, Edinburg,Texas. 
NuttallJ. 1835-1 836. Collections towards a flora of the Territory of Arkansas.Trans. Am. 

Philos.Soc. n.s. 5:1 39-203. [Greenia \ publish 1 'I o-ter 1944)] 
Pohl, R.W.and G. Davidse. 1971. Chromosome numbers of Costa Rican grasses. Brittonia 

Powfll, A.M. 1994. Grasses of th< Iran; n I adjacent ai i Univ. Texas Press, Austin. 

Shinners, L.H. 1 972. Shinners' spring flora it ilia Fort Worth region [,] Texas. 2nd ed. 

Prestige Press, Fort Worth. 
Silveus,W.A. 1933. Texas grasses. Published by the author, San Antonio, Texas. 
Smith, E.B. 1 994. Keys to the flora of Arkansas. Univ. Arkansas Press, Fayetteville. 
Smith, J. P., Jr. 1981 .A key to the genera of grasses of the conterminous United States. Mad 

River Press, Eureka, California. 

Sii m;i i ,E.T. 1 8/U.Nomenclator I lotanicu 

s.Ed it 


nda. Pars II. I it 



t i>it,i, Stuttgart. 

Rrrfil,E.E. 1971. Survey of occurrences 



soft endosperm ii 


ass genera. Bull. 

TorreyBot. Club 98:264-268. 

1 r. ,CB. 1841,Graminaagrostidea,ll. 

s rotur 

lu \c;n ' 



.Acad. Imp. Sci. 

St-Petersbourg. Ser. 6. Sci. Math., Phy< 



traits ,| 




1 1 G.( 1 ' "> 1 l - gen m i ol P >oi lea 


e} in t lie sou 

i United States. 

Harvard Papers Bot.9:1 1-90. 

A- 1 -,G. 1883. The grasses ofthe Unitec 

I Stafc 

;s. U.S.I 



/asey,G. 1885. A descriptive catalogue ofthe 


s ofthe Unit 

ed States 

.Gibson Broth- 

ers, Washington, D.C. 

'aison, Land M.J. [An/wo 1992. [he grass qenei.i of the world. ( AB. International, 
>Accessed 1999 October 5.) 

'uNDi-RUN,R.P1998.Guide to the vascular plants of Florida. Univ.Press of Florida,Gainesville. 

mskilvych, G. 1 999. Steyermark's flora of Missouri. Vol. 1 .Missouri Department of Conser- 
itKiu l< Id i on i i' in | 'h thi 1 i j r 1 1 i it il ' iid: n Press,St. Louis. 

E.GeneTowne Iralee Barnard 

(Pottawatomie County), Kansas was submitted t 
sity (KSC) for identification. The grass was identified as Themeda quodrivalvis (L) Kuntze,a 
plant native to southeastern Asia. , tn> > Hit • I tab , ih ■ in ,- in been collected in 
Louisiana (Brown 1945) and Florida (Wunderlin 1998), presumably as an escapee from 
cultivation. A search of herbaria records revealed that Themeda also has been collected 
in California (RSA). 

Themeda quodrivalvis can be recognized by the following description: Annual, 20- 
50 cm tall; blades papillose-ciliate on the lowen-i rgin li ul m mbianous, lacerate, < 
1 mm long; inflorescence a laxflabellateclusterof racemes,each subtended bya strongly 
keeled spathe;spikelet clusters with a fertile sessile spikelet surrounded by four infertile 
sessile spikelets and two infertile pedicellate spikelets,all partially enclosed by a spatheole; 
fertile spikelet terete, bearded, 4.7-7.2 mm long with a 3-4 cm long geniculate awn, 
disarticulating obliquely to form a pointed calk mm long;flower- 

ing Sep-Oct.The specimens may be assigned to variety helferi based on the presence of 
tuberculate hairs on the involucral spikelets and the plant stature (Bor 1960). The inflo- 
rescence of Themeda is a complex arrangement of spikelet clusters that is described in 
greater detail by Baird andThieret (1985). 

In a follow-up examination of the collection site, eight additional Themeda plants 
were found growing undei a bird feeder and in an unmowed area near the porch. The 
yard was semi-shaded, and the lawn was predominately crabgrass {Digitariasanguinalis) 
and bluegrass {Poa pratensis). Microscopic examination of birdseed from the feeder found 
numerous Themeda spikelets among the thistle (Guizotia abyssinica: Asteraceae) seeds, 
indicating that it was the source of the introduction. 

cial thistle seed with different lot numbers were purchased and inspected thoroughly 
(30.1 kg of seed). All of the sacks contained Themeda and other contaminant seeds,which 
were removed and compiled. Three hundred of the Themeda seeds and numerous other 
unidentified seeds were planted in vermiculite in a 22° C greenhouse and monitored for 
five months. In the spring, an additional 900 Themeda seeds were broadcast sown in a 

Themeda seeds removed from the birdseed were apparently inert. However, some 
Vigna radiata var. radiata (Fabaceae) and Arthraxon hispidus (Poaceae) seeds germinated 
in the greenhouse. Three unidentified dicotyledons also geiminated but died before 
reaching maturity Both U , u w^o , , ,nd ' >piaus are weedy annuals from 

southeastern Asia that have been introduced into the United States (Kartesz 1999). 

One year after the initial collection, examination of the void where Themeda was 
discovered did not tin I <u i in u tth , i m ' n numerous resi- 

dential yards wheie tin ,M« i i I It lid il lid not detect any exotic plants. 
Themeda may be unable to persist in Kansas, but in Louisiana it has flourished for more 
than 40 years in disturbed sites near cultivated fields (Reese & Landry 1 985). 

India and Ethiopia are majoi produces of thistle foi birdseed (Sharma 1 982; Vin- 
cent & Cusick 1 9m j in I ^ < t I i ui i nil 1 1. i i il h weedy species, are 
present in the h in e t I M i If if a mi eeds are treated to prevent ger- 
mination, the process is fallible. I ontaminalion oi inconsistent sierilization procedures 
apparently were u p u i loiint let fa t n , a, 1 he incidence of 
viable seeds in thistt ma 1 - n 1 i l ' t bit f i -, '< sents a potential source for ad- 


Thanks to Larry and Satah I ieml n hoinitiali\ i a I tin itass in their backyard, to 
Gail Wilson and Fmil I -i i f n i th it i i f stations, to Gordon 

consented to a florist n ut ey atoned their bud ie< lets upport for this project was 
provided by the Kin j n h'i il I ( -urn i t t it i d i 1 1 monal Science Foun- 
dation Long-Term Ecological Research Program. I his oapm is contribution no.00-l39-J 
from the Kansas Agricultural Experiment Station. 

Baird, J.R.and J.W. Tim hit. 1 985. Notes on Picture ,j ,r t u< .,, . ie) in Louisiana. 

Bor, N.L. 1960. Grasses ol Burma, Ceylon, India and Pakistan (excluding Bambuseae). 

iVuiam.on Press, New Yob.. 

a i I iP i > 1 1 i I ileal attributes fc 
United States, Canada, and Gieenland.l iist I dition.ln:Kai te 

ynthesis of the North American f Iocs Vetsion 1.0. Nortf 
i, Chapel HilfNC 


Reese, W.D. and G.P.Landry. 1985. Tl 

Sida 11:99-102. 
Sharma, S.M. 1 982. Niger cultivatio 

/.Press of Florida,Gainesville. 

Prk.iii and Nani yVoikman 1999 Landscapes in History: Design and Planning in 
the Eastern and Western Traditions,Second Edition. (ISBN 471 29328 8,hbk.) 

Mi, il n In m ||,i, I a nnuo New York,NY 101 58-001 2.589.95. 844 pp. 

Numerous b&w photos. 

.including current North 
iddle East, Asia, and North 

[.Wayni Smith and ,. Cotton: Origin, History, Technology, and 

Production. 05Third Ave., New 

York, NY 1 01 58-001 2. $250.00.850 pp. Numerous b&w photos. 
mm (he preface: Tl u u n / 'tun I 1 | i liinl I I tlmuqhout most tropical 

nd ubtropicalteqior Ml Mill i i m n .m nnmHii it 1 ust nn tin t 

rom herbaceous p> < m I t mi I in h~, I uu i i I i m limn iln i Idoiih hi 

■nt l ) hoit titt d< n t hi a n h jii ih ii ii i 1 li | .1 i ti hit tibm th it < hnji 

on/e In ]iih ii! | ' ! illi it I El I' i r 1 1 til trill .1 ii i n i i I | I n 1 It f >n i !• lib i I >i 

1) » 


Mohammad Athar James Harding 

^ ^ u, ,, ( , _ ,,,, ,tE^„onmentalHortu 

University of California-Davis 

Davis, CA 9561 6, U.S.A. Davis, CA95t t jaharding* 

\ survey was condL w, t d,aim. ihl< ,/, \ , |, ,i| t|l > from the Tahoe 

.asin. All were found to i lu it in i tun i t tr ti u ported for the first 

Ir ' l! ' : 'i'"" nesp.xiesinfw'.jeneno; .m :: " - >.'ac. diesn siicciesa'c distributed n tub.-, i „-■■;-]! ■.,:■, 
]enisteae,Thermopsideae, Irifoliean and Vi< km.;. Nodul dor. shape and frequency are also de- 
cubed. The results supp . t imh i t, it it ' I i in filar structures to classify legumes. The 

lealthy plants with lu h |i nt >li i i , u n I n u nn, nt unpi i tint t| 


T are 69 genera a 

species found in California (Hickman 1993). Most legumes me indigenous while 
are naturalized, often widely so (Witham 1994). Bacteria associated with legur 

atmospheric nitrogen and are helpful in improving and mult ii in i . I futility I 
cigiKullun tnd nmii il < i-\ inn Tht n I I [unm i-i > ,< laticui pr 

lation in 66 Papilionoid speues from Sauamcnto Valley, California and found that the 
ever, a majority of ihe lepumos found in ( alifotuia have noi been examined for their 

nodulating ability of some legume species from the Tahoe Basin of California's Sierra 
Nevada Range (39° N, 1 20° W). Plants were collected from accessible areas within the 
Basin, ranging from Desolation Wilderness in the wesl to the C aison Ridge in the east, 
Alpine Meadows in ti t ith ,i III |< .11 it I n i mllul i passes in the south. 
Geologic and clima.tic foices have mmbined lo form lahoe's unique and varied 
environment (Blackwell 1 997;Graf 1 999).Tahoe's floral diversity can be attributed in part 
to its central location hei < n plant ( mmuniti ft! i j Nevada western foot- 

variety of plant species tan f either bo explained by the a tea's rapidly changing and var- 
ied habitats, which have contributed to the large number of Tahoe plants with limited 
geographic ranp( u ti i 1 Ith it i t ti i in the basin is mixed 

due to variations in temperature, precipitation and soil, toniteious forests dominate 
(Blackwell 1997; Strong 1999). 

Legume species growing under natural nditmu t n I lot their nodulating 

ability. Periodic field trips were made from late sprint) to early fall in various parts of 
Tahoe Basin. Observations were made as described previously (Athar 1 996a). Legumes 
examined included indi i i u ndnm I iuml in it ml in s At least five plants 
of each species were examined to minimi/e eitot I eoumes were identified by speci- 
mens of mature plant ife, ml. ledistingu hod 1mm. olhei kinds of morphological 
modifications or pathoqenk root malformation, and Modulation (fata were recorded. In 
some cases, nodule sim it md uodul lm i , pamd and < imined ui lei th< 
microscope (Somasegaran & Hoben1994). 

Nodulation status was examined in Ml hapiiionoid soe, ies oiowinp under natural con- 
ditions in the Tahoi I m II h M i l ' ^d < r luCt d to various extents 
(Table 1). These result i mi i I ithth i ul ibl t |oit hi nodulation (Aguilar 
et al. 1994; Allen & Alloi I thai "oih MMI nt mi MMahmood 1 990; Corby 

1 988;de Faria et al. 1 994; Nasim et al. 1 998; Roggy & Prevost 1 999). Nodulation is reported 
for the first time in mi, i ;petie< ithin five geneia if Papilionoideae. These species are 
distributed in tribe i J i i t TI i i i I 1M lieae <md Vicieae. Joe 

presen< i 


Nodulating Species 1 

i ..ri'_-; i- ■■ 
I lonqjt-Ml 


Thermopsis monu 

IvJiiijcil ,i 

I l.nii.ldi.'r! 


: longafc 

Nodulating Specie 

llr * 


1 |i ing i 1 ' i 

as described by Polhill and Raven (1981) 
Brummitt and Powell (1992). Introduced s 

Modulating status 

B - Nodulation reported for the first time 
+ = Indicates sparse nodulation (1 to 5 no< 

du , espe 



d a witi 

IN lil 1 

compared the [i 1 lilt nl I -I I i n i n II in I l< new reports. The 

nodules observed in olhei species cotiobomte eatliei studies { Allen & Allen 1981 ; Athar 
1 996a, b; Athar & Shabbir 1 997; Corby 1 988). 

lateral roots, and were found in the 10 cm layer of the soil. Nodules of some legumes, 
particuhtK un > ft ' tit n i th it Ml d md were covered 

by a layer of damp litter. Similai obseivaiions vveio made bv Athar (1996a) for some 
logu nes from tl II I 1 ' 

for various species a Ml i mthtli n tb t i i >l th phut- Nodules varied from 
semi-globose to globose vvitli sttom.ob or smoom, iif i i elongated and branched 
forms (Table 1). They occurred sit ml m ,, b I tujut r ■ J' ' morphology strongly 
coincided with the descriptions of earlier workers (Allen <b Allen 1 <)H1; Athar 1996a;Corby 
1988;Pueppke& biou tut i ' M mm m t << i i latan & Hoben 1994). 
Nodules were mosM >M i bm n I'm t I li b n m tmts Nodule morphologies in le- 
gume species desenh. d b\ ^ ui im i to IthiMru me uniform at the tribal 

of establishment ol the -, t h een the two partners (Pueppke & Broughton 

1 999; Roggy & Prevost 1 999; Sprent 1 999). 

Attempts to isolate hnr jhi ; at t- -| m ,], ui <,, activity were not made for these 
nodulated legumes. Howevei, In .ih 1 1 >i ihlu h ■ imeti f mi i j.> gtowing in a nutri- 

ent-limited environment imply that they were nurtured by nitrogen-fixing nodules 
(Hartwig 1 998;Somasegaran & Hoben 1 994). This is supported by the generally accepted 
view that nodulated legumes have an internal regulatory system to allow them to ad- 
just nitrogen fixation to environmental conditions (Hartwig 1998; Sprent 1 999). 

l.upinus (! upines) were the most prevalent nodulated legumes inTahoe Basin with 
N ,pt'. ii s IoIId i (I b\' !"l /" Id ci J, i /i / < ivck h) and Of/ ■ , , u - , n 

10, 4 and 4 species each respectively (Table 1). The soils of theTahoe Basin generally am 
nutrient poor, especially near the surface where drainage greatly exceeds the rate of 

poor environment of the Tahoe Basin. Physiological processes that plants undergo to 
survive in the Basin help in understanding Tahoe's vegetation ecology. Nutrient limita- 
tion plays an important role n plant distribution. Plants cope with nutrient deficient y 
through mutualisms with mycorrhizal fungi or through associations with bacteria capable 
of nitrogen fixation. Nitrogen fixation is a crucial component of many plant communi- 
ties in the Sierra Nevada Range, where nitrogen is ways in si 

the legume- Rh i /ohiu' symbi i ii nutrient poor environment of theTahoe Basin may 
help improve natural ecosystem, and provide refuge and guality browse for wildlife. 

Special gratitude is expressed to Joseph H. Kirkbride, USDA Beltsville, Maryland for his 

I j 1 1 1 i n I I it i I luin n jmenclature and taxonomv aii I n| n lit 

it ninitn il i I i i i I Himenodulation.ThanksareduetoSteveShoer 

California Department of Food and Agriculture, Sacramento, California for helpful c 


J.I. Sprent and D. McKey, eds. Advances in legume systematics. 5. The nitrogen facte 
Kew, Royal Botanic Gardens. Pp. 25-31. 

\thar, M. 1 996a. Observations on nodulation status of some Papilionoid species of pc 
tential agricultural and forestry value from Sacramento Valley, California. Taiwan 

ohar, M. 1996b. New nodulating legume species from natural ecosystem of Pakistai 
Phytologia 80:385-388. 

A M | /<< 1 ^ | ihl i tu i Mli 


ity of legumes of Pakistan. List 5. 

Acta Bot.Gallica 144:67-72. 

Aiman.M. 199/b.Nodulation status of sorr 

e legume spec 

cs from Cache Valley and notlh 

ern Utah. Phytologia 81:145-1 50. 

AiHAK.M.and A.Mahmood.1 990. A qualit 

ative study oft 

ii nc : ilatin i ibCty of 1. nun - s 

of Pakistan. List 4,Trop. Agric. (Trinidac 


Athar, M.and 5.M. Shabbir. 1 997. Nodulation character 

sties of some of the forage and 

browse legumes. Phytologia 82:1 2-1 c 

Blackwlll, L. 1997. Wildflowcrs of tin 1 Tahoe Siena. L 

ne Pine Publishing, Redmond, 

VVashmgt on. 

Brummi i i , R.K. and C.E. Powell (eds.). 1 992. A 


names. Royal Botanical Undoes, 

Corby, H.D.L. 1988. Types of rhizobial 

nodules and 

their distribution among the 

Leguminosae.Kirkia 13:53-124. 

FARiA,, I <\ M < ,VPC i , and J I 5 u I "o4 cMLunonce of 

nodul iii. m ii I o it H peci In mi I i Mi i ci f lm e, <■ miai md I pirito Santo states in 
Brazil. In: J.I. Sprent ot u. I I ICe ul 91 nm ml. mm. I malic s 5 The nitrogen 
factor. Kew, Royal Botanic Gardens. Pp. 1 7-23. 

Grai,M. 1999. Plants of the Tahoe Basin: Flowering plants, trees, and ferns. California Na- 
| - Plan! Sen irt\ i, nnu nt, 

Hariwic,,U.A. 1998. 1 he angulation oi symbiotic N , fixalion:a corn eplual model of N feed- 
back from the eco tin tetM in t i i I M i | i I I silt EcoI.Evol. Sys- 
tem. 1:92-120. 

Hickman, J.C.(ed.). 1 993. The Jepson nunc il H iM i : .mi . ' e Ml una University of Cali- 
fornia Press, Berkeley. 

NAS!M,M,M.AiH-.h,aniM M i n I" 1 ' Tcmmii .n . mn mm nodulating legume 
I i . it in II i I i f h t M )ia 85:1 10-1 14. 

Pomin.R.M. and P.H. Ravln (eds.). 1981. Advances in legume systematics. Royal Botanic 
Gardens, Kew, UK. 

Puippki, S.G.and WJ. Bkoucihon. 1 999. Rhi/obium sp Strain NGR234 and R. fredii USDA257 

RoggyJ.C and M F P I " < u "mm n tern i « nun. r I in nesis in rain forest in 

ii n hi mat i i! m mil ind eoligiul mi i b M II I 1 1m > ' 1 

Sow i ran, P. and I I.I I 1994.1 ndbook fot rhi/ol h ml I n legume-/?/i/zob/um 

its hnoioav.Spiinom Veriag, New Yen k. 
Sprlni, J.I.I 999. Nitiei ih itionandgu IMMi.h i p i>WMn pi Lies in diverse en- 

u nnu nt Peisp< [.Plant Lo I ,'ol »yst 2 1 1" In 
SironoD.H, 1999.Tahoe:l rom timbei barons to ec ologiMs. Univwsily of Nebraska Press, 

Wiiham, C.W. 1994. Jepson prairie presence plain Cms k iist. Jepson Prairie Docent Pro- 


Loran C.Anderson 

Department Of Biological Science 

Florida State University 
Tallahassee, FL B^IBn : \ , 

The following appeart 

Cyperus retrofractus, Li 
of north Florida are a 

Los siguientes taxa pate i i ti in ' 1 i > B H m h A 

documentan tambien varias adi n i i f'ora del norte de Florida, y algunas especies que se 

nordestede Florida. 

Florida" was defined as the Florida panhandle west of the Suwannee River. Recent bo- 
tanical surveys, particularly in Clay, Nassau, and Putnam counties of northeastern Florida, 
have prompted me to expand the area of coverage to include that portion of the state 
from Putnam County northward. Particularly interesting sites were found at Black Creek 
Ravines Conservation Area just E of Middleburg in Clay County and Ralph E. Simmons 
Memorial State Forest just E of Boulogne in Nassau County. Exotics that appear to be 
adventive or naturalized are included.Nomenclature generally follows Wunderlin (1 998). 
Herbarium specimens are at FSU unless noted otherwise. These findings will update 
species distributions as mapped by Wunderlin et al. (1 999). 

Ajuga reptans L— Leon Co.: extensively naturalized in Tallahassee lawns, 7 May 1999, 
Anderson 18620; new to Florida. 

Aster laevis L. var. concinnus (Willd.) House —Okaloosa Co.: N side of Karick Lake, 1 1 
Oct 1996, Anderson ?779/;Santa Rosa Co.:shrubbythicketaround lake atCamp Paguette, 
6miNNEofMunson,27 0ct1995,/Wmon 76036,margin of Krull Lake,Wof Sweetwater 
Creek, 1 1 Oct 1 996, Anderson 17157; new to Florida. 

Duplicates identified by Almut G. Jones (pers. comm.), but I think they may repre- 
sent a new taxon. These Florida samples have disk corollas 3.7-4.8 mm long, whereas 
disk corollas from four samples of A. laevis (from neighboring states) ranged from 5.2-6.0 
mm. The Florida diskflowers have long lobes (1 .3-1 .6 mm long) and short throats,whereas 

with short, stout rhizomes and a branching pani< ulale miloiesc en< e, whereas the Florida 
plants have long, slender rhizomes and more racemiform inforescences. 
Chenopodium murale LI nue inTallahas 

see, 6 Mar 1999, m I -^ n i I lor ida panhandle. 

Conobea multifida (Michx.) Benth .— Leon Co.: railroad crossing at Capitol Circle NW in 
Tallahassee, 2 Oct 1992, Godfrey 84404, same locality, 6 Oct 1999, Anderson 7 9056; new to 
north Florida and econd >unt\ f r< ill il i la. Many authors (e.g., Gleason & 

' )|] 111 t \ ' <] II t I I ]■ 'i i ■ ' - ' ' .ii 1 iv I Mil. 

Cyperus retrofractus I in i . - li t h i ins State Park with 

Desmodium ochroleucum M. A. Curtis and D. rotundifolium DC, 10 Oct 1 998, Anderson 
78788; new to Florida l> GCm I 1 '■ I e " C. '. U i lln I lor ida panhandle, but 
the plants to which h< > tMiruii i i Ct :,--o-* Fernald;see Carter 

Linum macrocarpum Rogers.- Bay Co.: pine (latwoods near I ong Beach, 7 Jul 1963, 
Godfrey 62966; Franklin t o.: ApaLu hu . Ci National I orest:, F of Rte 65, 5 Jul 99, Anderson 
78870;new to Florida.The Bay County specimen was annotated as Lfloridanum (Planch.) 
Trel. by Rogers, but later Bruce Sorrie noted its capsules were 3.4-4.6 mm long and iden- 
tified it as Lmacrocurpum.The rranklin County spec mien was immature, with capsules 
3.3-3.5 mm long and shrunken seeds 2.2 2.6 mm lorn 1. 1 Ins specie-, was previously known 
only from the U Mabama (Rogers 

1 963).Anderson 16859 adds a second locality (eas; of Bayou LaBatre) in Mobile County, AL 
Lipocarpha aristulata (Cov.) G. Tucker. -Putnam Co.: wet depression beside railroad, 1 
miNofPalatka is «| I > \n o > ' -^ ' • mi. k ihc ^ < M G>% Anderson 18500; 
new to north Florid i I i il ipni urn [ i Ci < ' - A /7a (Cov.) Smyth. 

Mecardonia procumbens (M\\\ n ill uii m i In : nt in holder of lawn, S of 
rte 100, W edge of Palatka, 16 May 1 997, Anderson 17444, same locality, 19 Sep 1997, 
Anderson 18152; new to north Florida. 

RhynchosporathorneiKra Jack oih o.: marshy aiea Comet i no Like 1 mi Eof Marianna, 
27 Sep 1 957, 7<ra7 & Godfrey 5996, edge of small pond 2 mi NW of Grand Ridge, 1 Oct 
1997, Anderson on 1 mi N of Palatka, 18 Sep 

1 997, Anderson 18 u t Florida. Ih pecies is pi iousl known only from the 

type locality in BaC i unt , mini] nil lie It I ml lln. v specimen (origi 

nally labeled "Rhyn<lh< ; >m ; oo ■ ha| m ' a anie-taCd o 7V f/\wne/ by Krai in 

,vn weed at St. Johns River Com 

Thunbergia alata Bojer ex Sims. 

Leon Co.: escaped from c 

inTallahassee,22Jun 1998,Ander. 

■ ' -" t t o t n. ilh e i 

Thymophylla tenuiloba (DC.) Sr 

nail. Lee n ( esc eel f 

ized in Tallahassee, 7 Jul 1 L ) Q 9,' -• i to north Florida. 

Xanthosoma sagittifolium (L.) Schoii. -I con Co.: naturalized (vegetative) in swampy 

woodland by Ochlockonee Rd, NV\ fTa iha e< Nov 1996, Anderson 17247, same 

locality (flowering), 20 Sep ' to north Florida. 

Abutilon theophrasti Medik- l-control pond on Call Street in 

Tallahassee, 7 Jul 1 999, Anderson 18816; new county of record and second collection for 
north Florida. 

Aureolaria virginica (L.) Pennell— Clay Co.: steephead ravine at Black Creek Ravines 
Conservation Area, 1 Jun 1 999, Anderson 18691; new to northeast Florida and first col- 
lection in Florida east of Ochlockonee River. 

Boltonia asteroides (L) L'Her— Nassau Co.: bordering St. Marys River at Hwy 1 7 bridge, 
9 air mi NW of Yulee, 5 Oct 1 985, Anderson 891 1; new to northeast Florida. 
Callisia repens(Jacg.) L— Leon Co.:escaped from cultivation and naturalized along road- 
side ditch in Tallahassee, 20 Oct ]998, Anderson 1849 1; second county of record for north 

Carexcrebriflora Wiegand— Ch < ' . hid .t m| fl > -dplainof BlackCreek, 


Carexstyloflexa Buckley— CI lain of BlackCreek near junc- 

tion of the two forks, Black Creek Ravines Conservation Area, 1 Jun 1 999, Anderson 18687; 
new to northeast Florida. 

Chasmanthium latifolium (Michx.) Yates.- Nassau Co -shaded banks of St.Marys River 
at Simmons State Forest, 25 Jun 1999, Anderson 18788, 8 Jul 1 999, same locality,A/iderson 
18842; new to northeast Florida (fir t IF> r mii M 1 1< ' 1 < ,ot of ( ) ( hlockonee River). 
Croton willdenowii G.L.Webster.— Bay Co.:abundant in white sand of depression just E 
of Enfinger Rd, S of rte 20 (ca. 2 mi W of Econfina Creek), 27 Oct 1 999, Lisa & Ed Keppner; 
second county of record for the state of Finn i Tl ,f . . u , mm, previously known as 
CrotonopsisellipticaWWld. Penm II 1M i mt it i in the south part of its range, it oc- 

Hill in Washington County. 

CyperuspseudovegetusSteud -N i hi ' h i mm ions State Forest, 25 

Jun 1999, Anderson 18764; second oimi I i in Florida east of Leon County. 

Desmodium rotundifolium DC— Jackson Co.: Florida Caverns State Park with D. 
ochroleucum and Cyperus retrofractus, 10 Oct ]997, Anderson /8?85;third county of record 
for Florida. 

Fimbristylis decipiens Krai. — Leon Co.: lawn and open weedy slopes above pond E of 
Mary Ellen Drive in Tallahassee, 8 Oct 1 992, Anderson 14062, same locality, 25 Oct 1997, 

Hypericum gymnanthum Englem& A. Gray— Nassau Co.: set -page slope, Simmons State 
Forest,25Jun19Q ( MM ' ' > '^ n to 1 1 md,i ,ind fitst collection in Florida 
east of Ochlockonre Over. 

KyllingasquamulataThonn.exVahl.— Leon Co.:locally common at Tallahassee South- 
east Farm, S of Tram Rd, 25 Sep 1995, Home 773, same locality, 26 Sep 1995, Anderson 
15899; new to Florida panhandle. 

LeersialenticularisMichx— Nassau Co.:along stream in Plummer Swamp,4 air mi WSW 
of Yulee, 24 Jul ] 999, Anderson /OOp now to nmtheast I lotida and first collection in 
Florida east of the Ochlockonee River. 

Lithospermum incisum Lehm.— Putnam Co.tdry sand of open oak woodland in Palatka, 
27 Mar ]998, Anderson is > - >> . mid ui it\ it i . >n I lot northeast Florida. 
Murdannia keisak (Hassk.) Handel-Mazz— Leon Co.: under Meridian Rd bridge W of 
Lakelamonia,18 0ct1993,4nderson 74590, 27 Oct 1999, Anderson / 9082, shaded streamlet 
at Elinor Klapp-Phipp i itt J t d i< oihinhi t . >\ I >«, ■ let son 17068; second 
county of record foi hht I i pui'i it II t> In l> n 1 On I igqressive weed. At 

the Lake lamonia site, it appears to be displu mo '' >' > " > •< ei'-nenanum Cham. & 
Si hie. ht, which is an endanqeiod species in Fioiida p ouo POo). 
Oxycaryum cubense (Poeppig & Kunth) Lye.— Bay Co.: frequent along edge of drain- 
age ditch, W of Rte 389 at Girl Scout Camp, 5 Jun 1 999, Keppner 1 120. Reported (as Scirpus 
cubensis Poeppig S i uutln K i< II > I i -' I ut n >u h i liom the Florida pan- 
handle were found it m i it lni I hi ut I lit IP >).This collection docu- 
ments lis piosotH o; tiw sp«s ins has also been lounU lecentiy hi Alabama ami ueoiuiu 
(Bryson et al. 1996). 

Paederia foetida L— Duval Co ilb CI i n Rd near University Blvd in Jack- 
sonville, 1 3 Nov 1 998, Podris s.n.; Franklin Co.: St. George Island, 1 3 Apr 1 995, Jubinsky s.n.; 
LeonCoswoodland along Crestdale Ln,NE of Tallahassee, 30 Sep 1 997, Jubinsky s.n.; new 
counties of record for this Class I weed (FLEPPC 1 999.) 

Paspalum conjugatum Berg— Clay Co.: mesic woodland bordering St. Johns River, 
Bayard Point State Preserve, 17 Aug 1999,/PO 'm ,l t t , themtrimih 
Plantago major L. Nassau Co.: beside rte .' at M.tVliivs Pivot NiiclgcW of Callahan, 24 
Jul 1999, Anderson > >nd itu P tecord for northeast Florida. 

Plantago rugelii Doom h [ n Pin i Itei t i I Florida Caverns State 
Park, 1 Oct 1 997, Anderson 181 90; second county of record. Additional sites in Leon Co.: 

Platycladus orientalis (L.) Franco.- Marion Co.: naturalized, several trees of differing 

ages in pine-oak wood Linn Sot orry Rd,E oi rte I ( ) in Oca la National forest, 23 Oct 1998, 

Anderson 18507, sec in i i, It it O i G mt Glected since 1905, see Wunderlin 


Polygonum lapathifolium L.- Nassau Co.:edqo ot i ultivated lield, ca.4 air mi NE of 

Boulogne, 8 Jul 1999, Anderson 18852; new to northeast Florida. 

Pycnanthemum flexuosum (Walt.) B.S.I 1 . Nassau ( o.: semishado of seepage slope, 

1 999, Anderson 19009; new to n Hi . i I ti 1 and first collection in Florida east of 

Rhynchospora macrostachya Torr.— Nassau Co.: shallow pond on seepage slope, 
Simmons State Forest, 25 Jun 1 999, Anderson 18762. Reported for the state in 1933 by 
Small (but not by Godfrey and Wooten 1 979 or Wunderlin 1 998), but this is apparently 

beaksedge from two sites at Simmons State Forest {Anderson 19007, 19014) is puzzling in 
having achenes with short bristles tvpic al for R. corniculata (Lam.) A.Gray but with stout 
rhizomes as in R.careyona Fernald. 

Schisandra glabra (Brickell) Rehder.- Leon Co.:mesic woodland alongTrillium Court in 
Tallahassee, 1 May 1999, Anderson 18617; new county of record for this endangered spe- 

; (Steud.) Gleason.— Nassau Co.:campground at Cary State Forestjust 

Spermacoceverticillata L— Putnam Co.: roads 
son 18122, same general area, 5 Dec 1997, /And 

Vaccinium tenellum Aiton.— Clay Co.: overgro\ 

Area, SE of Green Cove Springs, 1 Dec 1 998, Sic 

Creek Ravines Conservation Area, 10 Jun 1 99 > I n 18665; S1 lohns Co.: powerline 

corridor, Moses Creek Conservation Area, 2 mi WNW of Crescent Beach, 9 Apr 1998, Ware 

s.n. These collections confirm its oc :urr. n n! f ri la VanderKloet (1988) noted a single 

1 943 collection from Clay Co.and in | I It i nn\ no longer be found in Florida; 

the same situation was noted in 1 996 (Luteyn et al.). Surprisingly, Wunderlin et al. (1 999) 

isted the species for over 20 counties in north Floric 
d" I, tnbution and all previous repod-, in, i, niv? Li >n - > intu , 

itJu-t I nent of the 1 genu 


1984. Noteworthy plants from north Florida. 5 

'86. Noteworthy plants from north Florida. II.:- 

1985 Ki i rl | I it mi ttiil i In HI Sida 13:93-100. 

1989. Noteworthy |>l nit It i i rth otida. iida 13:497 '4 

. 1991. Noteworthy plains horn north I louda.V. Sida 14:467-474. 

1995. Noteworthy plant ft no ihl i h v| Sida 16:581-587. 

Bryson, C.T., J.R. Mac-Donald, R. Cartfr, and S.D. Jones. 1996. Noteworthy Carex, Cyperus, 
Go h lhn,\ Kxlliiuhi and i '■ >. ova' 1 ' a \pwa< <m<- tion i Alabama, Arkansas, Georgia, 
l,iiMihi, M M i ippi Moitle noma [ennessee, and Texas. Sida 17:501-518. 

Carter, R., and C.EJarvis. 1986. Re-evaluation and leptotypification of ScirpusretrofractusL 
Rhodora 88:451-455. 

Ciiwtn, A.F. 1985. ( ude to ih 1 nl 1 fl nt < n i I i mhandle. Florida State 
'mi i t it\ Pt hi it I i til m i i Hi il is . 

Coin, N.C. 1 998. Notes on Florida's endangered and threatened plants. Florida Depart- 
ment of Agriculture & Consumer Set v., Division of Paint Industry Botany Section 
Cootrib.No. 38, Gainesville. 

FLEPPC. 1999. Florida's most invasive plants list. I louda Ixotu Post Plant Council. http:// 

States and adpt< > nl ( anada 2" l od. New Yoi k B< on i< al ( .arden Bron> 
Godfrly, R.K. and J.W \ 1'^° \puutu aid Pan plants ot outheastern United 

States. Monocop I. ■ I i Hi i n m a Pu s, At! or 

Kral, R. 1977. A new pecies ol ( poruca i n u miihwestern Georgia. 

Sida 7:42-50. 
Luteyn, J.L, W.S. Juian, S.R Vanoi r Kloei, Ld. Dorr, G.D. Wali ac i , K.A. Kron, RF. Stevens, and S.E. 

Cli-manis. 1996. Ericaceae of the southeastern United States. Castanea 61:101-144. 
PlnneipKW. 1918. Notes on plants >fth otlaai Unitt i tuti I Bull. Torrey Bot. Club 

Rogers, CM. 1963. Yellow flowered spo. ios ol litmni in eastern North America. Brittonia 

- i l\i a S.R I II i i i ii \oph i . ii i I - I Bwnch In 1 

Aoir i a; are Canada. 
/undfriin, R.P. 1998. Guide to the vas( ulur plants of Flonda. University Press of Flc 

.\o .. ust p i I | o o ' l' 1 at 


John L.Strother 

Stw ,'/«'■■>,;."••< 94-.V M65. U.S.A. 

Nesom (1999) and Gandhi (1 999) n i. i mil n ittribulions and datesforthe generic 
name Euthamia and for binomials in Euthai i rhe expressed slightly different inter- 
pretations; here is a third interpretation: 

The first known use of Euthamia as a botanical name was by Nuttall (1 81 8). Nuttall 

wrote of his Euti | to Solidago and 

Chrysocoma" (Fig. 1 ).Only a taxon of generic rank can be said to be "reciprocally allied" to 

Fig. 1. Pages 162 and 163 from volume 2 of Nuttall's The Genera of North American Plants. 

time, reciprocally allied to solidago ana Ltirysocoma. Nuitan \r\g. i j associareu u 

thets ijniminitolijd-MJ tenuilolia wi:h the generic name /' mhaniui and, in his proU 
I / ! i ] I I nil ill n i it) t i i )t n 

Elliott (1823),Cassini (1825), and de Candolle (1836) treated Nuttall's Futfw 
,|nneii( n im< | ubl I I in I I 11 1 1 1 h n i i M ' u i II n H i Nutlail 
and E. tenuifolia as published in 1 81 8. 

Regardless of typographic lapses orenors oi inconsistencies of numbering . 
I r nn i in i in h n if tr ill n iln lQ99;Nesom 19' 

papers cited b\ tin n n r i Mi i J i ill r ; names should be: 

i.N. 2:162. 1818. 

jtLGen.N.Amei pi it 2 1 H •> I ' \ t . , < 

a (Pursh) Nutt.,Gen. N. Amer. pi. 2:1 62. 1818. Basionym: Solidago te 

RN I PI Ni ! l 
Candoi i i , A.-P. ni . 1 836. Solidago. In: Prodr. 5:330-342.TreuUel and Wurtz, Paris. 
CASsiNi,H.1825.R/t/iom/u. In: I .Cuvier, Did. sci. nal.,ed. 2. 37:471. Paris. 
Ei i loi i, S. 1 823. Solidago. In: Sketch dot. S. Carolina 2:368 -392. J. R. Schenck, ( 

Reprint: 1971. Hainci Publishing Co., New York. 
Gandhi,K.N. 1999. Nom i I iiilu In twin 1 in Hemisphere plants 

Pap. Bot. 4:295-299. 
N 1 1 1 I ) ' h i i i i i I itui in 



OCTOBER 24, 1909 - DECEMBER 5, 1999 



.were back on pto^ it | i it i | ut i 

ch a good show, you need to know all the plant names. I c 
3 following week I mti a- , in ould become my r 
aat; adding names to tl w i h tde show used in a 

successful effort to 

have two smal 

" i ' in tilti nlJ 1 u.tln l I jli n ii t 

derness System. By 

the time thee 

'er, 1 had taken all the botany courses 

offered atTulane,w> 


a master's octree, and found a new career 

For over thirty 

years, Profess. 

ir Ewan t u Fed the lives and minds of Tulane stu 

dents. None of the h 


Jdents who enrolled in biology 431 (Plant Systematics 

in the catalogue bu 

t really a cour: 

se in "Plant Appreciation") could ever walk ao > th« 

campus in the same 


It ' it | it i t 1 aid ' on I i 'via t it t 

and his beloved hist 

ory were wove 

tint ant ! 1 i Foli n er mi ed a ! it 1 tut 

nor forgot one! 

For one whose focus would b 

e the FMory <<< vfam eJoseph Ewan left a renaAablv 

thin personal paper i 

.-rail/The facts c 

M islif. am in: .1, i not easy to find Born in f t tla iei 

phia, educated in Lo: 

s Angeles, he re 

'O - ' 1 1 1 1 in 1 n hi It u 1 tFt 11 11 t it t 

California, Berkeley in 1 934. In 1 935, he married Ada Nesta Dunn, a fellow student who 

diarce: many of his i 

nterests and h 

is life for the next 64 years. They had three daughters: 

Kathleen, Dorothy,ar 


Ifivegiand 1 1 1 1 not ttucto t the F 

/ of Colorado (1937-1944), botant r nh rh I it- I t mic Administration (1944- 
45), Assistant Curator, Smithsonian Institution (1945-1946) and Associate Botanist, 
reau of Plant Industn U' depatttiimii > 1 1 1 > It j- mi, Vn | n 1 947, he came to 
ane as an Assistant I ' f< in, • I, i| ti- n ademic ladder,and in 1972 was 

ned the Ida Richardson Professoi if I tan 1 Fat Ft In It i 1 mt ill 977 when he was 
aointed Emeritus Professor.For nearly forty years,Tulanestudents,faculty,and staff mem- 
-s were exposed to his mind, hr astlnn- I i- ami his enigmatic wit. 

Along the way E m 1 1 A 1 1 mm 1 r . II 1 I t ill ti il Science Founda- 
-1 Fellow (1 959-/5 1 1 i miih titn in I t t, ( ut-, r< IF. t]t ) mCiM tatinipmt. t 

He received tht i i i ' ' 1. tmi i tl it I ii i t IGtui ,H 

London in 1977, thi I losii Paym Luepiei I'vledul f mm lis Guidon Club of America in 
1978, and with his wile Nesta Dunn 1 wan, mo Henty Shaw Medal from the Missouri 
Botanical Garden in 1 994. Ewan was an elected Fellow of the Linnean Society of London. 
The Botanical Society of America awarded him a Certificate ol Merit in 1 989.The College 
,i , i Ih in, in I i i ,1 \ mi I lu in. i Mn ii it i ii d- I mi i n uiuuiiy doctorates. 

"A bite of imrnortalityms the Ewanian concept of publication. His own first "bite',' at 
age 19, was /\ Report > '' ' ' > "f>' published in Condor. 

His early botanical interests focused on fems,and he was President of the American Fern 
Society in 1951.Taxonomic studies on Delphinium and other Gentianaceae followed 
but his interest had il o I i pi ii D I i nil I iblu ><naphy,and history 

pletely replaced taxonomy in his research. Questions to Ewan were most frequently am 
swered by questions ^nd one left his ollu e buidenud with nooks. He believed books 
were to be used and 1 I , i i h hi i uM r s As a new graduate 

student, I rememho nii mil I u , I n Co I down from above his rickety 
desk, a beautiful vellum copy of Baulnns hodiomw. with the disclaimer that it was only 
the second edition irom 1 071. Back in my cubicle, I was afraid to open it! 

Othe- samples from his"bitcs"include sm h tempting titles us:"! rom Calcutta and 
New Orleans, or tales horn Barton's grecnnouse';"Rools ol the California Botanical Soci- 
ety""Who Conquered tl ill a mi r tu > t I o un in indehiscent cap- 

sule" and "The Botany of Cook's Voyages; or around the world on six shillings a day." 

Then there are Ins many contributions to The Dictionary of Scientific Biography 
where ones finds, among otheis, sketches of George Lngelmen, Albert Spear Hitchcock, 
Elmer Drew Merrill. Frederick Pursh, and the irascible Constantino Samuel Rafinesgue. 
His introductions to the clussica Bomnica Ameiicaiui series are classics themselves. As a 
book reviewer, Euan o ,n nil mCll mn i mi In enue were always 

tantali7ing.Asin'At| lb linningisth mn toiv i >i nine pigs driven 71 miles by a peon 
carrying a pine ton 1 1 vhi h certainly tempts >n. i I: into Aichie Carr's"High Jungles 

one mt to 1,1 ml icqnii ha it i. nhceg's The New Orleans Garden." 

t'em ijjs m m, n ,t I it u I I I ' ' 1 ' n > I' fol 

I n up i M !U i] ) IP and John Banister 

and His Natural Hhh» * mn ' tt Dtut ,utl < I Gais wife,Nesta Dunn 

Ewan. In 1969 Ewan edit D ' «' w„ ,►/ ,; m ,,<> m,p > nes and contributed 
the Calendar of Events and chapters on Early History and Plant Geography. Here he graphi 

cally illustrated the l\T to En m, i mm i Hi i jo ' il m believed William 
Bartram's contributor i it r It ih I i | m i i a I i o it the United States. His 

canPhilosophicalM.oi I >• Gil I ,u n nun ,t ,i Pi miaphy of Philadel 

I by the Missouri 

Botanical Garden. 

In 1986,the Ewans moved from lulane. I he Missouri Botanical Garden had bought 
his prodigious library ml i i them nt h m< I mi. the\ ere housed in the old 
museum building and loi the first lime the I wans had spacious working conditions 
with large double desks for each of them and his 4,500 books carefully catalogued and 
shelved within ea a u 1G i n noui moth n > F u i i-u lector of the Garden, 
said "The Joseph Ewan colli turn in i thiol aluabl icguisition for the Garden. 
The historical significance of the collection, its relevance to the work we do, and the 
respect Joseph Ewan commands in the scientific community make this announcemer 

We a Tulane are left with his name on the door, lots of happy memories, and hi 
herbarium now over I " "' |)c.n i j lit et f u < I a teguest from Swil 
zerland forTulane's holdings of Macrocarpaea,<) genus monographed by Ewan in 1945 

typewriter which oc.ncT I n th< in . loom I i ■ many years. 

The 1989 bwania: I he Writing >IIocandNi tatwm lists iSH"bites of immortality 
The following ten years produced many othch'bites." Surely a veritable feast for generc 
tions of botanists ioi it il I u i m I i i mf h i nth it neiations to come. 

JoEbGGioandANNOL il^klllPihnl hiT i or) 1999. Wild Orchids of Texas. 

(ISBN 0-292-7471 2-8, hbk). Univ. of Texas Press, P.O. Box 781 9, Austin/TX 787 13-78 19, 
U.S.A. $29.95, hbk. i-xii, 1 -228 pp., color photos, distribution maps. 

Eh i ' ■ f Icxas is an nit tandinq r lilt tv\ | ios of oi hid kn i 

chapte- giving an overme i In, i i i i tsity and conservation 

Subsequent chapters qi h ( .,, P t, I „ im mi iitmtmu ion .l^uch topics as pollina- 
tion, saprophytic spe< i- in I ll tiiih iitn i In II, -.i t II 1 bv a detailed discus- 

h'Mi adloMiiiih ntd m ' I i I ,i i tth nun name is said to be de- 
■tvecl bom two classical u t | i i I > t I i t t to the lip tufted with 
< omrU.I nairs An insiqlitlnl li u i i | li n I I ih. j ollm ilu 1 1 
am victims of these <it ^pti r li i i i i I l i aai) in return for the 
1 - • ' th such information 
as common name ramm i Im nT I mi i i i I uii in -, taxonomic and no- 
menclature! history, habitat mdotb > -i J I i I mi ■ mm time, and a county dis- 
tribution map showmq pm> n l'», .in mi Mi 1 - i ihih i mill „i minis are given for each of 

'.at' s qcnera ind | ^ i ' il n I 1 ^iiin ■ it only praise is due this work, I could 

-vmeamwofthe mo t tun >\ nti in r i ,n it i tl i pott< I itstnl utiun il i < ud 

I hr authms havi'C.eaily ciivcn om-s . :■■;:.. ;■ i t : ui,.l im; om coot as usoU; ,:S misohle to 

"amiliar. there is a list tn il i i 'til i l th i it m si and interested lay 

n-otocraphy Joe Lqq nff. I i r II ( , h 1 photo are a testa- 

' nl In m ti 1 t i i i n i i ii n i I wledge of orchids.' the species photographed tun I. im I rm ml -Is !1 sn and olten do so only in extremely 

i- •■•:: ■: -S'-siale places. The complete i i i i thus it rt rlv the result of hard 

it imgnidiiwi i Th t i ttl |i , nflim m 1 by the pleasing layout 

n 'li ivious thou |t t i t 1 i Ih -. ' i it t i Mil n | | mil ml 

> tingquot im ,h , , i u n i n i i I \ omtmtj out particularly fas- 

the Texas Panhandle af f ir^nt at I t, I f lit lir i it<- ttqime during the last ice 

Ih nntirma ui is it I i t t t I i I , ,| make it valuable to 

professional botanists 1 he m is s and thmt Htm m lidiihh I nave u r ed the most 

up to d it mloitti iTiotuin mmitifi i n it- Imm I < mt I' tl mt obtain information for 

' E i 'mm ui i i i II M in h it 

1h d isd welcome mli< t i i n n it ' i il i ll il i ' t i t k> often valued more 

tli, in I n ivvli 'I i oi igan ,ms in the wild. 

dust jacket to the su|i il N t n\ \ nl M n il n INI i it v is clearly a labor of 

love for the authors. This is a hook that anyone intended in texas plants definitely should not 
n)\ss— George M Di<k, f ^ vid Botanical Research 

hr.titiit<<<>! Icxa\! oil Woilh, 1X76102. 

John H.Wiersema and Bi anca Lon. 1 999. World Economic Plants: A Standard Reference. 

(ISBN 0-84^, ; I" ' o nl if n Hi hn a, i , C .'"'Mi orporate Blvd. NW, Boca Raton, FL 
33431 U S A (1 mi _^ 1 in 4 MM) 1 25.00, hbk. 749 pp. 
While discussing some nl my mseait h on ihe Kallavvava he'halists oi Andean South America with 
one oi ll 

validate some of mv > il in Im i I'm in i ul n | I ml a iiaiiin mportaiu e, tl m 
reference volume i sun I nil I I m I mi il | I lit on cm I ll 

economic plants Data i n| >il If tl it | mil ui I I mtion Network (GRIN), 

which is the USDA/AR in i n u i i n i t u I i hi i ueh of this database is 

oltth lielt i dei ulna i llnlinn Jim i .nil II h Mi i i get an answer. 

betically. Part one is the < atalog of economic plants oiienng the sc ieniilic name, synonymy, corn- 
index of common names lhat includes the list oi uiminon names I nam part one with correspond- 

itesubjea loth i i ill i Ml i in ti nt n t Inl ii i mgc red Species (CITES) 

thors < ite tin i|i\) imp! i I ti | i i i ih| it t l I i i I u In I World Geographical 

mioninedit a, Atu i I iiti , i i | i i h | I i il i I n | Noit mm An an i 

Pacific and South''!- n i iH ill im m il n I i iinte gel with the divi- 

,i ;n, hoi in t in 1 i I i in intlu i i I i i 1 t I t 1 f k i i I i 

South Amenta in. lud. ' mm ml, t i n I ml auth America. 

peci(i< i!l\ ( xotu I a. i, i .1 i i uti ilni I a us, i i m mllas'Tharmacopeiaof 

catalog of economic \ I -mi i I I i i , , 1t h r i h i 

citation for Ape/w/ana. When I looked for berro in tf mn niuni^iud tot its saeniifk nam. 
both World Econon m f r i tie, i H , , I i it when I looked in the 

catalog of economic plant 1 >i * u ,u i n n nl nomic importance as a 

authors leave out. I decided to suppler, i t n i nm i ith ntioduc ed Plants in the Indig- 

Economic Botany, 54(1):90 1 ,r m t r aril n ight limit confusion. I 

consulted the catalog ot ur.i pi it i i i | I mt ih ii I nnntl and f mm it 

as used to treat 12 bod\ I n [hi n [i | iti It 

investigating P.major, I notk - . I tht mm nun. listed is llanten Bastien lists 
llanten as Plantuqi u i i I I it t ^ t trouble locating at all. 

Using another plant from Bennc (t irui h m. ^ „ - , u<. .insulted the catalog of 

South American plant r in )l tai inmit md Prance. 

(%!'!, idian Medicinal Crops. (ISBN 1 

f)bl i N itinti il I ^ ,t i i i i 1 i I i n >ui inlii n | ( i | ipr i t 

M-55, National Research Cout il I n , i i t i I ' D h ( ANADA (613 

990-2254;61 3-952-7656 fax) htt t _ i ( j_hj uunl m gtapli lilml _ 
I'll 1 I i [ I i hi 1 h i i i hi ni 

ending to the demand >f th m ultuia uinmu lyfo it re information on n.itivi iti. 
si plants, the National ! em h u il l mi. hi f to luced a reference book that sets 
dtidard for other i ti to t I r i o no ti m i to the Ii mil piant in n 
gross $100 million annu ill ill n i r ii in II i f t ) lu ing other crops, 

ir 1 i Pn it . iii I ol I ti , d '. '■ ' , , it in i, t it t t tin hi i\ ilt n.m - 
ly f res. upturn drugs, modi* in if pi mt m it iu« to a urn ntk luggish agn. ullunil m 

a'-plinlogy, < lassi^ 
sting facts. I or ear 

the web address tc 

) a 1 995 Health Canada document on herb 

s used as 



drugs, i he web links listed after the selects 

specific. For examr. 

:j' . 1 'Ml J til". 

study/newcrops/echinace.i.html plus nin. othoi sites. The addition o 

f web links makes informa- 

1 ollowinu (he information on plants < nun", ( lupteis devoted 1 

le regulatory and legal framework in Cana< 


id marketing me- 

section is quite int 

eresting- the authors discuss principal detf 

soft hen 


crops, and a list of 

non-native medicinal crops that could be 

grown ir 

i Canada. \ 

Vhile this section 

lion a full blown < 

:hapteron the subject, with references and web linl 

ss. Ihoro is 

also o section on 

miiitiid ,tk i, t < t Lit ' ■■ ml , 

i laws. 1 he last 30 pages consist of a list of 

experts, organizations, and publications, an 


Off III, . 

research in Canada, a list of general refere 

t ol web s 

hen dt ifimli, i 

glossary of pharmacological and medical 

terms rel 

evant to n 

ledicinal plants. 

was slightly 

disappointed that there was no general inc 

anMeJii inuh top; 

ateresting and I'm sure, invaluable to anyoi 

rting a medicinal 

crop, i veu though 

this book has a target audience (the agncc 

Itural cor 


1 wouldn't expect 

readership to bee: 

part of a 

general reference- 

to anyone interested in Canadian medicine 

il crops.- 

-Kevin D. 1 


John 1 Ka tr and ( AM 1 099 Synthesis of the North American Flora, 

Version 1.0. . en and the Uni- 

versity of North Carolina at C ha\ Mil B ft fotten Center,Chapel Hill.NC 27599- 

3375, U.SA 

I minium In I n nh i n u i | i i t f m l i 

dow, S.I.N 

i 01)0 )| i itin isysto, 

m,i 1) ROMon" 

a keyl i 'an I, 

This CD-ROM includes an updated ve 

s.M'. I- t - t." 

t ttwsaums.itr i h. Ci-'f 


omitting au 

p' i n 



I , in ;uinan\ efft. u ,1 .' i, ■ 

more straightforward functions.The Synth 

esis allows vou 

it displays the biokxjk il -itltil ut ,hh 11 tit I in n ,m I I he attributes covered 
are general plant habit 3 t, dicot, fern, gym- 
aquatic, submerged tin jti i ih I r i tl list goes on. 

These functic e beginning of the 

functions of the program I h i i i ul i m i , he ]umi. ,nd tour operations you 

can perform on the quu< in nun i m I i n n Mtnni f mm hen I found it somewhat 

convoluted if nol in hi qi 1 i ih - | I i , < i , | h i ,i i | i hi n , (union, intersection, 

J tr t 1 ii 1 ii t m be | tin 1 i i i | i tn I quel igi q I i mil t 

or taxonomic). As an e- 1 nf h , i t nh i , i if i i - i 1 1 h Ww nlll ml -, 

kota and perform the union op. win, ., he . |, , • ,,, [|, e piogram would then highlight all of 

the species that occur in the n ml n II i u ju ul i , id an attribute query 

such asall annual duet found in I Hi t i hi il i I i I il il i I juery with something 
like Asteraceae and all 

the lower hqht hand cornew Ihe ir r f s> -;tion oiwuiw.- ( i "s ro ays me taxa with the oarhwihr at 
tributes or in a partu ular ta>'H it - th ' i i m nt * w wi geographic regions. 

The restricted operation displ u ii i m ,t ,w . in t. i t. - the selected region or regions. Finally, 
th nut )| ntioiii tin tut n i ,l || i Ml i in Im ii i ji I displays all of the spe- 

ciesofthecircumscubed ju i n t t h- i Ii i th i m tliere are about 28,000 

species in the Synthesis (North AmniK i in el .< 'i ■ . i id about 6,000 species in Texas. If the 
"not'operation were performed on 1 i h n th p mm would display 22,000 

colors of fonts used in the display I query ststh, team differen! kinds ol information. This added 

JuDD,CS.Campbell,EAKiii ind I n i its 1 999 Plant Systematics:A Phyloge- 

netic Approach. (ISBN 0-87893-404-9 HIT nam, Inc. 23 Plumtree 

Road, Sunderland, MA 01 37 

$67.95, hbk. xvi + 464 pp., with CD-ROM. 

derable. homohonal literature distii 

by a considerable number of institut 

,iv( , l.mVk iimima tnekmirv level km m m i | mvlmjenem 
asoi and mom tcccnt i nations com emima the tuxomam 

il tu m [i i in In 'ii ill | i ritil II ii | i i I 
em; topi I mil Ian] e! 1 h it i 1 ' m tudcnt nl.i in hould master. I h 

? probably should be consulted by all instim luis of plain tuxonomv.givon emerging changes 
ur knowledge of evolutionary relationships. 

Apart fiomth* k»i iid Preface ■ I - Ii I i t h k n I i t e ]hti I if 

two appendices, sepaiaiotaxonumii and suhjm t mdices.und a CD-ROM. 

I h i| t> i On i ntit I II u n. i f r in' ,1 i id i- a I H' I it ti ill e tk | lulu 

etic context that pei 1 the ntii 1 <i lib uqi th< erm n, taorphy is introduced 
ca simplified dadugi ,m I mitb. hi I I n - 1 In - unnin the major lineages of 
its appears on the second pagc.Plivloueov n km a nil vino oum ipal kiointhout I he text,, is any 

In the second chapt.sp Viet hods .aid Pi inn. pi e-> o ; ' ndnnij: kstematii '.',' the principles of 

|,.i|onelk Aslemath ,iii m kepedm ai illnl iluinik I h l n atinenl is muc h too 

i n t tu I i t i ll Mi l I II i t I i i t i I | t | i 

,t> a teO tk i il il a li n . | I 1 1 i i 1 i il ti h a i il i I h at mii ni r 

n in t bd. p ml a 1 i 1 | II i illl nl ii 


ai< and pa 

c groups 

are show 

in fig 2.23 in the cor it I If hi i uli a 


cladograms early in t 

are used as terminal 

of < ' id- ii. am u.r pb. nil ' ! 

2. mv 


aitha e ie ,i\,: 


i.:omi m tbiid wveku! en mil 

ice thorough i on 

1 reh n ion of d mi in a mi ill n pin n 

n lin i t i nli cc 



amples of cladograrr 


perspective of begin 

no udmi'e Mi ii'| i i" 1 



is the inadequate coverage of phenetic and evoli 

i hmakicata. 

1 nkk.ierm; main 


ring Plan 

s:The H 

torical Background: 

aams to approximate mourn 

m level ol 


2 toward 

which a 

mhlvskaildskve. Iheauth 

is kstoucak prominent taxi 

s. Sometimes, howe 


poses a knowledge t 

itl gin ,in , tmlmt I a 


ck, such as 

inferem e to Be 


The detailed discussion of inflorescence 
muriate and indeterminate inflorescences 
:he basic knowledge heumnin tu ' it r > 

I M i i it in ii I H i [ i i Mm I n imiti I i ink dated to separah 

"The Evolution of Plant n i iti I it i l rt m of phylogeny, history, 

pec i. i, hU i disc u m ir n I I n , i p I j I i I n , i species concepts. Re- 
garding the latter,th> mil n I i r ,t nij m rli. phylogenetic species 
< >no | i ilthough it « iudcs m< wh> th ■<) i „ ,1 >m ,,| , i • f , i i < c i< . i mrntionc d ,il ill 

tartmg vith Chapter ^ Ph\l e ti I nhi t i r , of Tracheophytes, Ex- 
cluding Angiosperms Iih ill i I ii th i till i 111 mi ti. atments,following to 
a considerable extent thp I, iti m t ' th » u m I I i i up (1998). Along with 

i hapti i eight hu h > ei ingi i p mi r smi i 15 

families are treated in detail. These fan 

: aced in table 8.1 The more inclusive grc 

tracheophytes"(Lycopodiophytes, Psilophytos, Eguiset 

ophytes, Leptosporangiate ferns), 'he'" 


"Norm monoi ot paleoherbs""Magnoliid c 

|'ll ■ 'hllK )1 , I 1 -1 1 | I1H J | t I II 1H 

and a few smallei lineages;"l urosids I ai 

(. omak's and Eric ales;and"l uastouds i and II"). As indie 

aled above, the Ixxakisan excellent ve 

am Keys are typically piovided Id the inajdi amilms ui a given order. In some ca 

sicales), fairly detailed hi mi- a il \\ I i Itn nh the average studen 

p tin mi unl t 1 t il I I | i i i t t i t t u ing cladograms . 

ii i |ih,l ) i' ti' ti> th lit in nil it inh I ii i pli\lv i considered equi' 

sated with quotation m nf In in m m< I v i mil Put Hue .rnpnn 

loldlni d although t r it ni im il i Inn 1 ui|t n ate excessively detaik 

I he majority of leptosporangiate fern families aie suhs 

eturns latei in iahle 1 m Appendix . 

overlooked, such as AasfmAm Sys'i 

,n,j/i),' Ro/.a.MV.and Sni/f/i Mriuin 

i ] • i' I nm • ill n I i i i ii i iat< Ih it m 

'ini- * unable to call up an i- i ■ f i Ao/iMpo/mA 

mo traditional lamina! < lassitn ations of i longuist 

Ih ii i 1 . it ' th h b 

7iirer?)o/7 : Munuiuy Digital!),, and Antinhinam. '. ho expanded i'Lintagmaceae, 
( lude^:allit7Kliaco,icandHipniitida;.(viL\()l ; 'ornrnorasuihas/-V^a;/ l Jr,'5.C'us 

~Pvso and other taxowarrw ; ha ' ■: j< ■■_-. vvil . ha ksm;w ; T;r"knowlecino"of families and lorce ■,,', 


ilies in their previous 

incarnations, and th it t 1 I d rl N t Hi i i h I also caution tl 

challenged by I . i i i i i ii i ih u I n , i in o i in Ii .1 1 ut 

iaries.One can easily 

un i m i 1 n ihi in mi m 1 t n , tl t ' n .i let \ lologist by insis 

.ting that Penstemon 

hi I ti i in Tl ,nt i mi u i. i din t ih n p dm 1 11 is also fairly easy t 

o envision the older 

biologist wonder in i IhmIhi tt i f t hi ' i m i h n i 

I , ,i in i i i i il mi i hi- ii| i lit n u i i F n a i 

mies also provide an 

)'iiV changes i" the next can ab> .o'foi an omwaojmtv to dn 
u h as hi and when not to t ih> I in i on tlie"ron 
in yalc.. it would tie he king 'n . omnuw sense to pomilicate a 

. si knowledge of loi a plants. I wo videauues and i onu 

■ Mitch." in southeaster 1 i. olosh , o , .s^-' e - 1 e-e: o 

i h i i v ill . , , , n u n it I lit 

- n. ml then .; o|hh jhi m rth \<- I n m >h 

ii i 1 1 ilk in. tw 1 nttoductc 
a: an rich lor many beginning students.V 

m.ituetrd fot inlio 
a different title. The 
artl and Clark 1989) 

i I i l> i l 


-:i ::-:KC [-. 

Ann. Missouri BotGard. 85:531-553. 

Caniino, P. 1 999. Review of: ludcl, W.S., C.S. Campbell, I .A. Kelloejm P.I . Stevens. 1 999. Plant 
systematics:Aph>li i< i ti i| pi i h imii i itii i ind Massachusetts. 

S V st.Biol.48:826-828. 

Fru.dman,W.E 1Q l H) D ill I tt ' n n i in Ml inn d plant Us beautuj 

on thronpinol aiuiiosperms.Scionce AM'AS 1 9.S-1. 

FRimMAN,W.E.1994 flu . olution I . n u i n in i I [ I it 1 it J the developmental 
n in it I * L til. I,i t . i^fi I- ,p< in "M it Pi I F 8 ' 4r. 

Harm, D.L. 1988. A prime' of population annntn \, 2ivA Edilion Annuer Associates, Inc., 
MindoilniKi, MA. 

Harm, D.L. and Ac, i , P'8'lni |P l| pil i n imli 2nd Edition. Sinauer 
Associates, Inc., Sunderland, M.A. 

SmiiiiJ.RJr. 1977. Vas. ula> pFmt lamilim,. Mad Rivet Ptess, Inc., Fureka, CA. 

WAiTFRS.D.K.and DJ.Ki ,. imM.Vasailni psanl taxonomy. I out til I dit ion. Kendall/Hunt Pub- 
lishing, Dubuque. 

— Neil Snow, Depnim tt f n il in t t IN t'ln m Colorado, Greeley, 

CO 80639, USA. 

Vaugh.2000. Botanical Results of the Sesse & Mocino Expedition (1 787-1 803): 
.A Guide to Relevant Scientific Names of Plants. (ISBN 0-91 31 96-68-1 , hbk.). 
ml Institute lor Botanical I ut i. i l n i , i ,mi<\|iH fvi. iDi 1 1 ■ r 1 1 or ity,5000 Forbes 
enue, Pittsburgh, PA 1 521 3-3890. $55.00. 626 pp. 
;/-\;,;e: h* pe.rposo oi the, :>,.iv ':-, <■ taxonomk, in the oosc it is -m: ;oono 

Mil | MM III 1 Id I III II | I || [1 milk 1I| Ml I | i I III! Ill II Hi 

ni't'r oooiIoms woo have s'uHho 'Ac p.A'ic Jtions of Sesso tC Mooino, or <Aor spo. roeoo 
■.I io.' ■■ ,/ ; 'l:o-,pl!0:: ioqive noon-- o; pin. olen f o lot 11,0:0 < 

s applied by de Candolle, 

d Bernardia dodecandrc 

I 1 B 1 D 1 Sesse & Mocino Biological Illustrations. 

(ISBN 0-913196-60-6, CD-ROM). Carnegie Mellon CD Press, Carnegie Mellon Uni 
versity,5000 Forbes Avenue, Pntsboi ih ' 'A 1 521 3-3890. $40.00. Shipping/Handling: 
$4 Domestic; $6 Outside U.S. 


"Compartiendo Oportunidades y Retos para el 

iin jo usi ntabl let Ecosistemas Forestates" 

Mibifii],]i)lr- >\c \Ob 1 msisli'iiMS horcstalcs'.' 

I o i I. in i I ih t i | i i i j It il li i it nit j v tv i ii ii un 

de los Recursos Foh t i i I ' t i M - I i I itilic mun Foi t ih 

Lompiomisos htemacionales;6) Fauna Silvestre. 

I j I i I i. , 1MNL 

Ids: C»,'82 1) 248-9!) v 2-42 M y I ax: (52821) 
Email: y juvalle@ccr.( 

ollrntintioductory materials cm the history mid cieoloqvot 
th CentralTe<as im a t r ' i r i i i i ii \- i u | |jr ' i 

m . 10 t< II f I litl.1l lot I" li It III , iliitk I lilt i II n 

— Sharon hosier, A 1 qude tear Im, 
l n » r\ u- it' I 1, [ im ' 

- I'M I i it II ih ' ) | ihn' h I 
I ever\ gciiclonrr, laiulM <ipe.ii 




I - it's a new 

departure in Texas floras, <i breath of fresh air" 

' ,' ' " ' ' l ' illtll'l 

Manual o( the Va\at!ar rhiuw of icxa\ 


- m , • ■ , ' ' > t , ■ . i --I i ii ii , n ii 

it i I I'll Hi ii il. J li I i hi t II . I 

model of how a local flora should be prepared.' 

I 'iik tot I dmI I id, inn ( .aniens, Kew 








; VViiqiil MM:.: 
Botanical Researc 

Table of Contents 


Bonnie L Clark— 235 



J. Braem AND Karlheinz Si n. .has— 249 

John J. Pipoly III and Jon M. Ricketson— 269 

Discovery of Ardisia subgenus Acrardisia (JVV 
John J. Pipoly III and Jon M. Ricki tso\ -275 

R.i.i o. v —291 

FL M Rl 1 l»Vl i I n, -I MONTERO— 301 

of Boerhavia (Nyctaginaceae) 


Rii ' iLiii" P L M vs— 345 

L ii' i fi u in ii i r i ihi ii i ( i iihiiahua, Mexico 

\.Eduardo Estrada C a 

David J. Rosen— 399 

Thymelaea passerina (Ti -[ • f) \k /. mil. 

Wo i in ii.vr,, I 111 ( I i i i i Ia ii li ii in 403 


George Jones Goodman ( 

, 248, 256, 274, 284, 290, 324, 344, 402, A 


Asteraceae subtribe Boltoniinae Nesoi 
Asteraceae subtribe Chaetopappinae N 
A ii ■ ■ Pentachaetinae N 

Paphiopedilum c 

:rossii (Morren) Braem & Sengi ias, comb, i r rial nov— 251 

Paphiopedilum c 

K II l <0>NT!AMilM [ i II U tT 1 I 

Paphiopedilum crossiivar.sublaeve^R h Hi i n o „ i. „i , vm h stat. nov.— 254 



atum (Greenm.) H.Rob.&Bretteil var.tzimolensis (T.M. Barki. fy) B.L.Clark,c 

Senecio quaylei 

T.M. Barki ey, sp. nov.— 286 

icarpa LeBlond, SP. NOV.— 292 

runana Pipoly & RlCKETSON, SP. NOV —269 

JL," M III ! , i ETSTA1 NOV.— 261 



cobanensis (J.M. Coui i.) 1 1. Rs m. ^ Bo hi var. molinae (H. Rob.& Brettell 

■ . . -236 


randifolia(I l H.I i h 1 1 fell vak. serraquitchensis (Greenm.) 





I K5 662 12-2542, USA. 


-Senecioneae) is described, and 

r. molinae, Telanthophora grandifolia var.serraquitchensis, and Telanthophora sublaciniatus. 
y Words: Senecio, Telanthophora, Pittocaulon, Mexico, Central America 

■ describe una variedad nueva, Pittocaulon hintc- n < > \ se propon cuatro 

imbinariones nuuvas, Pittocaulon vclatumvar. tzimok in t , nl >p i > t i var. 

lanthophoraamu 1' c '■< "t n, is y -Telanthophora sublaciniatus. 

scent studies in certain Mexican and Central American segregates of Senecio, s.\. (Clark 
?96) have led to the recognition a new variety and the need for four new nomencla- 
iral combinations. These are recorded here to make the names available for curatorial 

miiH.Rob.&Brettell var. cerrograndensis B.L.Clark, var. nov.TYP E :MEXICO: 
isco:Sierra de Manant!an,on very steep E-facing slopes of Cerro Grande on road from Lagunillas 
Campo Cuatro & Juluapan, on limestone cliffs above camp 4, dry, deciduous « 

isis differs notably from var. / 
/ florets instead of 7-9. It is apparently restricted t 

n for a number of endemics (Vazquez et al. 1 995).The n 

Pittocaulon velatum(Greenm.) H.Rob.&Brettell var. tzimolensi 

comb. nov.5enec/opraecox(Cav.) DC. var. tzimolensis I b.\ I ni i hvtologia 69:142. 1990. 
Ti "hi, ) I m i tiopical deciduous forest 15 km S of Comitan on road to Tzimol 

., I I, . i i j 1 1— r r - 1200 m, 20 Mar 1981, D.E.Breedlove 50266 (holotype:CASI). 

B.L Clark, comb, et Stat. nov. Telanthophora molinae H. Rob. & Brettell, Phytologia 27:428. 

I^M, i UATFMALA Pi nin i . ni\i i n< n Alt , Fmi- n id id between San Rafael 
Pie de la Cuesta and Palo Gordo, W facing slope of the Sierra Madre Mountains, wet moun 
tain forest,elev 1800 Mm I i I L> i <o i „ - M n 

Telanthophora grandifolia (Less.) H.Rob.& Brettell var.serraquitchensis (Greenm.) B.L. 

Clark,combetstat n Mil it 

1,1 - ''"'' '/>' >> r"-',ji h,n - 1 1 ,r. '■nni)H.Rob.&Brettell,Phytologia2/:428.1 t )/4. !■>:■• : 

i I'M I \ en i ,uit h- -I, - n A, l • , i i 


al dissertation prepared at Kansas State I 
ion ofT.M.Barkley. Rupert Barneby of th( 
:he Latin diagnosis. 


lion in Scna.u ix.l. feu t >mr 



i V.WMAlisupd.Ued - 

absikao (Japanese) 

•cutifolium var.pioriverensis {? ll^VA) \t%itZ\Zilj&ZnT\,\%. 


s one of Papuasia's largest 
lent Area (CMWMA) was re 



the following ace 
is updated with s 


Canarium acutifolium (DC.) Merr.var.pioriverensisTakeuchi,var.nov. Type: PAPUA NEW 

GUINEA Chimph Pi in 'iihM' il 'U- M hmmmiI r- i I i. I i "i,, Pluvial forest, 06°47'S, 


Differta C.acutifolio (Di f it i if l I I n I i i i i 1 > cm longis, stipulibus 

n 1 i I ],]!]- nl 2 • ■ n mm li i t' in I I i J ' 12-14 mm 

Canopy tree, 25-30 m tall, deciduous, plank-buttressed to ca. 1 m height, outer bark fur- 
rowed, pallescent, slash ferruginous to ochraceous, clear-resiniferous, wood white, very 
dense. Branchlets longitudinally corrugate, ca. 1 .5 cm diam., minutely furfuraceous, pithy, 
not hollowed with age; vascular strands peripheral, appressed to the pith. Leaves spiral, 
phyllotaxy 2/4, 6-8 jugate, 1.0-1.4 m long, the flush emerging simultaneously with flow- 
ering; leaflets membranaceous, (ovate), elliptic-oblong, or oblong, c 
chis, (1 1-)20-31 (-43) x (6.5-)9.0-14.0 cm, apex rounded or 
truncate to (cuneate), margins entitt iuii i mi| lodromous, bifacially prominulous, 
secondaries 10-1 5(-1 6), at the leaflet center diverging 55-70° from costulae, gradually 
arcuate or turning abruptly at the margin, crossing nerves scalariform, reticulum dense; 

pusticulate, all parts with an indument of appressed furfuraceous hairs; petiolule terete 

or compressed, pulviniform at both ends, 9-25 x 1.5-2.5 mm; rachis cylindrical, shal- 
lowly sulcate, purple-green; petiole 250-390 x 7-10 mm, dilate at the branchlet, 
articulative or not, broadly concave on the upper side, rounded beneath; stipules persis- 
tent, inserted at the base of the petiole,acuminate, 1 4-1 6(-25) x 3-4(-6) mm, flat, paired, 
evenly tapered to the apex. Staminote inflorescence axillary (lateral), thyrsoid, pyramidal, 
51 -60 cm long, rachide branches to 15 cm long, all axes glossy green or reddish-green 
andwithan indumrntoi g il< nDlikdmn en hr In i '■ n n k ai uminate.the larger 
ones 7-10 mm long, higher order bracts ovate, 1-2 mm long, undulate; peduncle 170- 

tubiform, ca. 5 x 3 mm, 1/2-2/3 connate, lobes ovate, densely sericeous on inner sur- 
faces, glabrate or put ml mi m ! i II . Ii t i t imbricate, petals lanceolate-ob- 
long, 6-7 mm lotto a I i i ill kin i i r i ill ilk < m < ous hairs; stamens 6, 
uniseriate, egual, united at the base into a glabrous column ca. 1 mm high, filaments 
plane, 0.5-0.75 mm long, anthers basifixed, sagittate, mucronulate, oblongish, ca. 2.5 x 
0.5 mm, erect, pilosulous; disk dome-shaped or globular, rugose, glabrous, ca.l mm high 
and not exceeding the column, 6-lobulate, recessed at the summit; pistillode absent. 
Pistillate inflorescence unknown Infra, lescencc axillar\ (lateral), occasionally from defoli- 
ate nodes, ebracteate, puberulous, ca. 20 cm long, burn king only ai the top, peduncle 
170 x 4 mm. Drupe ellipsoid or (subovoid), 24-27 x 12-14 mm, obtusely 6-angled in 
cross-section,apex acute or bluntly rostrate, locules 3,2 of the cells sterile and rudimen- 
tary,exocarp with scattered subappressed hairv .ik \ • r w i | mading, not accrescent, 
lobes debate or rounded, 3.0 x 3.5 mm.adaxially densely sericeous, externally glabrate. 

Distribution and habitat.— Known thus far only from lowland forest in the CMWMA. 

Etymology.— The varietal name commemorates the type locality. 

Paraiypf: PAPUA NEW GUINEA. CmimhjP^uvincf: Crater Mt. Wildlife Management Area, Pio River, foot- 
hill forest, 06°47 5 141 '. I " n ml " liui ,■■ . < u i 1 X86B (LAE) . 

Malesian Canarium has been treau d i\ ku'i nit ' 1 ns5a, 1955b, 1956, 1959; cf. also 
Storms I 972). There are 20 species reported from New Guinea (ibid). 

The new taxon belongs to section Pimela (Lour.) DC. and is related to the complex 
consisting of C. acutifolium (DC.) Merr. and its allies. Because of the 6-staminate flowers, 
the novelty is compaiablo to i ,o< utitolium vni.oomjkom bul is more robust, with leaflets 
to 43 x 14 cm and with larger stipules to 25 x 6 mm. Although var.acutifolium has similar 

other varieties of k h i lu » i phi nologv ol vai p/omc/em/ i also 

distinctive. Accordin i t Ik i i \ i h it tl i t u i < mat tenstically leafless 
immediately piioi lo llowerino. I hen leport is substantiated by ike fact that other indi- 

later exhibited synchronous flowering and leaf flush. The correspondence between 
leafand flower en km i i ,l 1 i I i il |( hha denigricant mem- 

The new variety will key to the species using t 
(1972) and can be incorporated into the varietal c 
(Leenhouts 1956:292) with the following continuatio 
2. Nerves 12-15 pairs 
, i es 2-4-jugate, leaflets 7-18 x 3.5-8.5 cm, abruptly acur 

to17x 1.5 mm;fruitssubglobose,to 17,5 x 15 mm 

Leaves 6-8-jugate, leaflets (1 1- )70-31( Ai) x (6.5-)9.0-14.0c 


Coniogramme macrophylla (Bl.) Hieron.; coll. 12,197. Initially reported as 'sp. nov. aff. 
macrophylld (Takeuchi 1999: 953), the CMWMA plants have once-pinnate fronds with 
pinnae to 36 x 5 cm. Comparisons against extra-Papuasian material indicate that the 
survey collections are merely a vigorous growth form.The lax venation noted previously 
(ibid) is actually within the range of variation for the species. 


Archidendronhispidum(Mohlenbr.)Verdc.;coll.//,2/0.(Fig.l- hide' - ' l. 

was formerly recorded only from Northern and Milne Bay Provinces (Nielsen et al. 1 984: 

95; Nielsen 1 992: 1 33).The Crater ML provenance places the species further west, and on 

the southern side of the Central Divide. The plant is apparently rare throughout its 


There are two species in the series Ptenopae Nielsen to which A hispidum is as- 
signed (ibid). Although the congenerApfenopumVerdc. has terminal inflorescences, it is 
now apparent that A. hispidum is strictly cauliflorous.The Crater Mt. specimen confirms 
that series Ptenopae is characterized by an inflorescence of pedunculate umbels. How^ 
ever, unlike A. ptenopum, the gynoecium in A. hispidum consists of a single ovary, so the 

An accessory description is provided for the previously unknown flowers of A 

Inflorescence a panicle of umbels or (corymbs), cauligerous, pendulous, diffuse, to 
15 3 12 cm, all axes filiform, nitid green, ± densely hirtellous; floral bractlets chartaceous, 
oblongish or widest above the middle, ca. 1 .0 3 0.3 mm, falling early, aglandular, hairy; 
pedicels 5-1 1 mm long;rachide branches usually alternate, 20-50 3 0.2-0.4 mm,bracte- 
ate at the base, the rachis bracts scarious or not,acuminate,0.7-2.0 mm long, persistent, 
adaxially glabrate, externally hirtellous; peduncle 5-52 3 0.4-0.5 mm. Flowers (measure- 
ments from spirit material) seen in bud only, up to 1 1 per cluster, bisexual, perianth pen- 
tamerous,subfleshy,green l hirtellous;calyx obconic,1.0-1.5 mm long, margin subtruncate, 
erulose, or denticulate; corolla gamopetalous but connate only near the base, the lobes 
lanceolate or ovate, ca. 2.5 3 1.0-1.5 mm;androecium polyandrous (ca. 40-50), stamens 

1 . Archidendron hispidum (Mohlenbr.) Verde. Flowering habit. Scanned from Takeuchi 1 


globular, 2-locellate; < 


Syzygium hylochare (Diels) Merr. & Perry; coll. 1 1,847. (Fig. 2). The survey voucher was 
previously cited as 'Syzygium aff. roseum Merr. & Perry, possible sp. nov.' but has been 
rekeyed to the given binomial. Although the Crater Mt. specimen has leaves with very 
prominent, evenly-spaced, and more numerous lateral veins, it is otherwise referable to 
the new result.The stipitate base (4-5 mm long) on dried flowers and the comment of 
the stipes being very noticeable on immature fruits (Hartley & Perry 1 973: 1 68) are de- 
ciding.These latter features are well-expressed on the survey numbe (Syzygium hylochare 
is also apparently characterized by the appearance of large pustules on the lamina and 
inflorescence after drying, in the manner of S. malaccense (L) Merr.& Perry sens. lat.This 
identification aid can be advantageously appended to the couplet seguence for the 

Syzygium hylochare is much closer to 5. roseum Merr.& Perry than is apparent from 
the artificial key in Hartley and Perry (ibid). Specimens belonging to either facies should 
be compared against the conuetiM - i^ I,' • II- > n )ns from the southern districts. 

Piper arfakianum C. DC; coll. 12,453.Jhe survey collection differs from the typical facies 
inter alia by the much sparser indument and the pendulous spikes with unusually fili- 
form peduncles.The delicate peduncles are only 0.1-0.2 mm diam. (0.5-1 .0 mm in other 
provenances) and the rachis is similarly narrowed to 0.5-0.6 mm diam. (normally 1 .0-2.2 
mm diam.). On the fully mature spike the individual fruits are elongate-ellipsoid rather 
than the more typical sguat-obovoid shape, and laxly disposed so the rachis is occasion- 
ally visible between the berries. At 12 x 6.5 cm, the laminae are at the maximum end of 
the size range for the species. Although the Crater Mt. voucher was listed as 'Piper sp. 
?nov.'in the CMWMA checklist 1 1 il uchi 1 >9 » the collections at hand suggestthat 
the survey number can be accommodated within the variable concept for P.arfakianum 
established in specimen annotations by Chew (cf. also Jebb 1987). While the character 
states for the Crater Mt. population appear distinct from the species sens, str., future 
collections should speculatively be able to unite the herbarium variation into a single 
continuum. If this does not eventuate, then the Crater Mt. plants will be deserving of 
future recognition as a separate subspecies or variety. 

Piper is well-represented in the Crater Mt.Wildiife Management Area, with 17 spe- 
cies now documented for the locality 


the earlier compilation in 
LAE = staff collections from the 
es from the University of Papua 


ASCLEPIADACEAE (al! dets. by P.I. Forstc 
Hoyasp.;WT 12,752 



, d h in ii i; 

7 1I886A (type 


Vaccinium acrobracteatum K. Schum.; W, 

r 12,451 

.originally cit 



ted (WT 12,702) is 
tually for Codiaeum vanegatum (L.) Bl. 




a lagenicarpa Warb.; WT 1 

surveys were originally intended as part of a conpi'" •.- m\o v vssment through a con- 
tinuous elevationa! seguence,on a schedule eventually tocu m nate in the preparation 
of a plant identification guide. Because of the truncation of our itinerary in duration and 
scope, the major objectives are now out of reach. However the existing documentary 
base provides considerable opportunities for future investigators. As noted earlier, the 
lowland environment at Crater Mt. is still miativev unsurveyed and almost certainly pre- 
sents rich prospective opportunities lot discoveiy ( lakeuchi l c >oq). hven though some 
work has already occurred in the low elevation zone, previous efforts have been brief 
and sp,;[\illy lestm '.ee. I ho existing heibarium do. eavnlat/oi "s a so boh v skewed 
with past collectors tending to f vi o t h mm taxa such as understory herbs 

and shrubs.Aconscious attempt k ut forth 1 to nngsofcanopy/subcanopy phan- 

diii'i lis |i I lojioiti ii di 1 

a the Pio alluvial plain, are suggestive 

The Crater Mt. tract is positioned on the leading edge of the Australian craton, form- 
ing part of a southern geoprovincv whit ii is flonstkahy depauperate in comparison to 

1 997). The contrasting species content on north-south orientations has been attributed 
to the orogenic etiology of northern New Guinea environments, and to increased spe- 
ciation induced by the orogeny (Balgooy et al, 1 996; Welzen 1 997). At least part of the 
difference however, is an ai 1 ifact of the uneven state ol plant explc nation in PNG,of which 
a salient element is the lesser number of collections from the southern districts. Since 
mountain-building piocessesa f 1 n I 1 n 1 i- 11 i \ Guinea, the high- 

est species density from its austral geoprovince should be expected on the latter's north- 
ernmost margin, whom the plate has been forced upwards by tectonic collision with the 
island arc terranesThe novelties from Crater Mt a an be iatiopali/ed on this basis. Further 
exploration of thf v.'h i > 1 m 10-- ^tG I 111 1 in be expected to 

produce similar results. 

TheCMWMA mpie , nt 1 \ n i , , I m ,e pi v I • | . n , - ' . Iderness forest. Re- 
part of an overall program for d< I | n i nli il|h i nphu pstimates of the 
New Guinea flora. The existing CMWMA checklist consists of a diverse mixture of 
allochthonous and autochthonous taxa, including many cestui vtional records. While ipaium ih it lutu< linoitii .jilt- - | atterns suggested by 
geological correlation, new refmem. ni m hi t K t u from exhaustive work on the 
Papuan side. Additional surveys toward the lowland alluvial zone could eventually con- 
nect to the elevation h iuvmI- ii , nvd at Lakekamu (cf.Takeuchi & Kulang 1998; 
Takeuchi in press). At the latter site for example, Palmeria gracilis Perkins was found in 
lowland communities 1 oiiespondinq lo Australian piovenances, though the species is 
ordinarily strictly montane in Papuasia. A complet 
strable at Crater M 

Several generalizations can be offered from our ongoing inventory work in wilder- 
ness areas, many of them previously little-explored or unexplored. 1) Substantial num- 
bers of undiscovered species are seguestered in lowland habitats, even in accessible 
places, having been unrecorded because of the poor documentation from low eleva- 
tions (Takeuchi 2000). 2) Although montane areas are better documented than lowland 
ones, additional discoveries are more likely to result from working the south-descend- 
ing ranges, on the margin of the Austro-craton, rather than the Mamose-descending 
side. To be sure, virtually any sort of serious effort will result in substantive discovery. 
Even at this late date in Papuasian botany, every major expedition comes back with new 
collections.3) It is an imperative on inventories, for survey botanists to collect uncritically 
by taking everything encountered,in multiple numbers ratherthan being selective (Pipoly 
pers.comm.). Many of the most conseguential findings from current investigations were 
of taxa whose significance was only revealed in the herbarium. This is especially true of 
novelties from speciose genera. If investigators are too choosy in what they gather, these 

phasis on vegetative markers for plant identification, which not unexpectedly results in 
highly erratic nomenclatural systems. The significance and discriminatory value of ver- 
nacular names are considerably overrated. At Crater Mt.,this is exemplified by the Pawaian 
'way-e-be,' which was described as monotypic by village guides, but actually encom- 
passes Gymnacranthera,Horsfieldia, and Myristica;'m fact virtually the entire Myristicaceae. 
Numerous examples of comparable imprecision were documented during the CMWMA 
inventory. Especially when compilations are based on limited sampling,the local names 
will appear to be specific merely because of discontinuities in the polling. While it is 
often standard practice for investigators to report local names in revisionary work, these 
reports have little to acquit themselves unless they are cast in the context of a compre- 
hensive census accompanied by comparative evaluation of the vouchers. Our experi- 

tion grossly incompatible with formal science.The guirkiness of local naming systems 
has also been shown in the ethnobotanical polling from the recent Josephstaal surveys 
(Takeuchi 2000). 

The Integrated Conservation and Development (ICAD) strategy at Crater Mt. con- 

opment. Nearly all conservation programs in Papua New Guinea are now founded on 
this principle. As one of the largest of the ICAD experiments, the CMWMA also ranks 
among the floristically richest sites presently subsumed under this operational para- 
digm. Whether or notthe ICAD philosophy can achieve programmatic success in places 
such as the CMWMA is still open to question (cf.Saulei & Ellis 1998). What the Crater 
surveys demonstrate is that the long-term viability of ICAD at Crater Mt. could have dra- 
matic implications for a remarkable and precinctive flora. The nature of future scientific 
contributions to be forthcoming from this area is in many respects an imponderable, 
but the results achieved thus far clearly point to a considerable promise for further floristic 


The botanical surveys of Crater Mt. were supported by principal funding from the Liz 
Claiborne and Art Ortenberg Foun ) in n Ml tl> Mhn D.and CatherineTMacArthur 
Foundation. Staff of the Research and Conservation Foundation of PNG (Robert Bino, 
John Ericho,Paul Hukahu,Paul lgag,and Arlyne Johnson) assisted with logisticsand com- 


■algooy, M.MJ.van, RHovenkamp, and P V\ i I 99(. : > Phvtoqeography of the Pacific - 
floristic and historical distribution patterns in plants. In: A. Keast and S. Miller, eds.The 
origin and evolution of Pacific island biotas: New Guinea to Eastern Polynesia: pat- 
terns and processes. SPB Academic Press, Amsterdam, the Netherlands. Pp. 1 91 -2 1 4. 

lARTLEY,TG.and LM. Perry. 1 973. A provisional key and enumeration of species of Syzygi urn 
(Myrtaceae) from Papuasia. J. Arnold Arbor. 54:1 60-227. 

luxLhY,C.R.and M.H.P.Jebb. 1993. The tuberous epiphytes of the Rubiaceae 5: A revision of 

rBB, M.H.P. 1 987. Key to Piper species in Papuasia. In: R.J. Johns. The flowering plants of 
Papuasia. Dicotyledons. Part 1: Magnoliidae. PNG University of Technology Forestry 
Department, Lae. Pp. 103-106. 

eenhouts, P.W. 1 955a. The genus Canarium in the Pacific. B.P. Bishop Mus. Bull. 21 6:1 -53. 

1955b. Florae Malesianae Precursores XI. New taxa in Canarium. Blumea 8: 

1 956, Burseraceae. Flora Malesiana ser. 1, 5:209-296. 

1 959. Revision of the Burseraceae of the Malaysian area in a wider sense. 

Canarium Stickm. Blumea 9:275-647. 

T. Baretta-Kuipers, and P. Guine 

Mimosoideae).Opera Bot. 76:5-1 20. 
gram, CJ. and H.L. Davifs. 1 987. Terrane 

. 1984. The genus Archidendron (Leguminos. 
s and the accretion history of the New Guir 

m> i til i Li ,- -I ' ii oph KM 1 ) Ml 
aulei, S.M. and J.-A. Ei us (eds). 1 998.The Motupore Conference: ICAD p 
from the field. A report of the presentations of the second ICAD c 
Island (UPNG),Papua New Guinea 1-5 September, 1997. Dept.of Environment & Con- 

i M >h 1 .1 ii I linn- j N M n he* I | n < nl I i mn < f f > t 1 

BuMivPfsity MjiiMMvahm & Resource ' 1 i i m i 

Steenis, C. G.G.J. van. 1972. Addenda, corrigenda et emendanda. Canarium. Flora Malesiana 
ser. 1, 6:921 -928. 

Takeuchi, W. 1 999. New plants from Crater ML, Papua New Guinea, and an annotated check- 
list of the species. Sida 1 8:961-1 006. 

2000. A floristic and ethnobotanical account of the Josephstaal Forest Man- 
agement Agreement Area, Papua New Guinea. Sida 19:1-64. 

In press. Rhododendron loranthiflorum (Ericaceae) from mainland New Guinea. 

A distributional record and new subspecies. Edinb. J. Bot. 57(3). 

In press. New and noteworthy plants from recent exploratory surveys in Papua 


and J.Kulang. 1998. Vegetation Part 2: Botanical survey. In: A. Mack, ed. A bio- 
logical assessment of the Lakekamu Basin, Papua New Guinea. Rapid Assessment Pro- 
gram Working Papers no. 9. Conservation International, Washington, D.C. Pp. 36-39, 

Welzen,P.C.van. 1997. Increased speciation in New Guinea:tectonic causes? InJ.Dransfield, 
M.J.E. Coode, and D.A. Simpson, eds. Plant diversity in Malesia III. Proceedings of the 
Third International Flora Malesiana Symposium 1995. Royal Botanic Garden, Kew. Pp. 


iim Woi\( mi Bulmamn and Jack Blcki k. 1998. American Garden Literature in the 
Dumbarton Oaks Collection (1 785-1 900): From The Newengland Farmer to 
Italian Gardens:An Annotated Bibliography. (ISBN 0-88402-253-6,pbk.).Dumbarton 
Oaks Research Library and Collection, Washington, D.C. U.S.A. $35.00 pbk. ix + 243 

published in the United States between 1 785 and 1 900 and are housed in the collections of the 
Garden Library of Dumbarton Oaks.Traditionally, the library's dominant holdings have been in the 
areas of English, Fren< h m h h,n ml it i tut M l |in it th i terests of garden histo- 

Dumbarton Oaks expanded its holdings in 1 9th-century American literature. 

Although, as noted . iln ,iith. n rli n jditional form of bibliography no longer has the 
same importance as it did in the past, because more and more libraries place their complete hold- 
ings online, and printed bibliographies are still valuable for scholars because they often provide 
important additional information in introductory essays.annotations, illustrations, indices.and other 
supplementary mati i ji u ml i n ,e ilm i tti | esent bibliography lies. 

The book is organized in five sections: I. Introduction; II. Annotated list of titles; III. Chrono 
and V. Bibliography. The introductory essay presents a history of the 
ning, horticultural, and landscape, cemetery and park design. The an- 
iequate comments on each title's contents. The index is extensive and 
t values of the volume. (Tie wishes ioi mote and better illustrations 

logical li 

■unl Miles; 

literature of Americ 

. one of th. 

blbh.HIt,|. oi (| 




Guido J. Braem KarlheinzSenghas 


Ki ,VV rd Orchidaceae,Cypripedioideae,Cypripediaceae,Cyprii 
barbatum, sublaeve, potentianum, Taxonomy, Systematica Oi 
Code of Botanical Nomenclature, Saint Louis Code. 

During the preparatory work for the third volume of Braem, Baker & Baker, The genus 
Paphiopedilum - Natural History and Cultivation, the literature regarding Paphiopedilum 
callosum (Rchb.f.) Stein was reviewed.The taxon referred to as Cypripedium crossii Morren 
has been interpreted as a synonym of Paphiopedilum barbatum by Braem (1988) and 
considered to be a nomen nudum within the synonymy of Paphiopedilum callosum by 
Cribb (1 987, 1 998). A review of the original literature has conclusively revealed that both 

:ole,21 years before the publication of Cypripedium callosum (Rchb.f. 
icle, Morren simply states that the Cypripedium originates from Peru 
he name of its discoverer, Mr. Cross. No description or any further 
? available. However, plate 1 7, which is part of the article, shows a 

plant labeled as Cypripedium > < [hi illustration leaves no room to doubt that the 
taxon published by Morren as Cypripedium crossii is identical with the plant described as 
Cypripedium callosum by the younger Reichenbach in 1886.The Morren "note" has been 
interpreted as insufficient to be regarded as a valid and effective publication of Cypr/pe- 
dium crossii as an autonomous taxon, an interpretation generally based on article 42.3 of 

"Code." Article 42.3 (taken from the Saint Louis Code [Greuter et al. 
effective version) reads, 

2000], which is the 

clciiiih uiJingiclenliUcatn '.n. (',,;; ; , 7 ; ?,;/>/< ' .'a' I he pi ; i i^ose - <l 1 h ; 'i , 1 1 ! ! ' c i 'e , >/ < p I a c e < 

i single figure shorting 

And .irtn. 1,; 42.4 clarifies what is to be understood as an aralvsc 

"For the purpose ut ^ f <■>, > "wures, commonly s 

'-•Initio i , , , ,!■ ' ,,/,,, , 

' ' m i option 


The Morren note was accompanied by a color plate of Cypripedium 

crossii (Fig. 1). There 

can be no doubt about the identity of the plant published as Cypripedium crossii by 
Morren in 1 865 is identical with the taxon we have hitherto referred to as Paphiopedilum 
callosum (Rchb.f.) Stein. The plate contains details that allow for the identification; two 
flowers are shown. The color plate clearly depicts the different aspects of the plant and 
flower such as shape and tesselation of leaves,shape and color of all parts of the corolla, 
from the front as well as from the back, detailed shape of staminodal shield, the ovary, 
and the floral bract. In other words all aspects relative to the identification of a slipper 
orchid. Thus the plate satisfies the requirements of Art. 42.4, especially as that article 
clearly states (see above) that the"figure"or"group of figures"do(es) not necessarily have 
to be "separate from the main illustration of the plant. "If this were not so,the inclusion of 
the word "commonly"in article 42.4 would make no sense. 

It is ludicrous to disqualify the publication of Cypripedium crossii by Morren in the 
scientific journal La Belgique Horticole knowing that the valid code accepts publication 
in trade catalogues or non-scientific newspapers before 1 January 1953, and in seed- 
exchange lists before 1 January 1 973 (see Code, Article 30.3). As every botanist knows, a 
publication in a seed list generally means simple mention of the botanical (Latin) name. 
Thus, Cypripedium crossii is for all purposes to be regarded as validly and effectively pub- 
extensive article entitled "Cypripedium barbatum, Lindl.and its major varieties, cross! [sic], 
warnerianum, etc/This article was illustrated with a colored plate (Fig 2). 

The plate reveals that the plant Morren described is identical with the plant he had 
depicted in 1865,thus the plant we have hitherto generally addressed as Paphiopedilum 
callosum (Rchb.f.) Stein. Morren wr\tes,"Cypripedium crossi [sic] has a very wide, some- 

Twiddle by a crimson stripe. The petals are half-green and half-white with 
d rose extremities. The lip is dark rose-brown." 

383, Morren published a description of Cypripedium crossii, although he 
s plant to be a variety of Paphiopedilum barbatum Lindley. 

In view of the facts delineated above, there is no reason to deny Cypripedium crossii 
Morren full taxonomic validity. Whereas the validity of Morren's publication of 1 865 could 
possibly be a matter of discussion (but see below), the taxon was distinctly described in 
Morren's 1883 article, and clearly identified as an autonomous taxon identical to the 
species hitherto generally referred to as Paphiopedilum callosum (Rchb.f.) Stein. Both 
Morren publications predate the publication of Cypripedium callosum by the younger 
Reichenbach and therefore, Cypripedium crossii Morren is to be given priority in accor- 
dance with the rules of nomenclature. The taxon is to be transferred to the genus 
Paphiopedilum. The authors are well aware of the fact that conservative growers and 
hybridizers will argue that the name "Paphiopedilum callosum" should be retained be- 
cause a multitude of hybrids has b^ n 'Mi ten I indicating "Paphiopedilum callosum" to 
be part of their ancestry. This argument must be rejected. The registration of orchid hy- 
brids (and any other hybrids for that matter) is no concern to botanical taxonomy. Fur- 
thermore, the registration authority for orchid hybrids (Royal Horticultural Society, Lon- 
don) is by no means a taxonomic ruling body.Furthermore,if the horticultural argument 
were to be followed, the name "Cypripedium" would have to be re-instated for the genus 
Paphiopedilum (and other genera), etc. Last, but not least, the identity of (at least) some 
of the hybrids must be guestioned. If we would reject a valid name because its use is 
: inconvenient to horticulture, it would, indeed be best to ignore all 
f botanical taxonomy. 


Cypripedium callosum Rchb.f, Card. Chron. ser. 2, 26:326. 1 886. Cordula callosa i 

Rolfe, Orchid Rev. 20:2. 191 2. 
Cypripedium schmidtianum Kraenzl, Bot.Tidsskr. 24:1 3. 1 901 . Paphiopedilum c 



lished.The differentia 

Fig. 2. Paphiopedilum crossii (Morren) Braem & Senghas, from Morren in Belgique Horticole, 1 

consists merely in var.sublaeve having somewhat smaller flowers with a smallei d 

(cf.Cribb 1987, 1998). Reichenbach fil. (1888), in his original publication of var. sub/ 
simply notes that the plant came out of a bate h of ( vpilpcdium callosum," and th 
might be supposed to be a natural hybrid." Cribb also states (loc. cit.) that the pet; 
var. sublaeve"usua\\y bear warts only on the upper margin/This, however, also appli 

the plant he depicts as Paph </ r - n , n page 333 of his i 

i i mi i in i ll it tl ii \\u in I tit tin. i th, I i I r it iaeh fil. publica 
rendered in a Bnli h in rii ihn-'n p if h r m I 1 1 [ n Hi F irlmut any descriptic 
illustration, has never been questioned, whereas the publication of Cypripedium c 
by Morren, admittedly in a Belgian scientific journal, and admittedly in French, shoul 
ugh Morren's plant can be positively identified by the ex| 

Db 1998). 

enotypes that are * 

I Mime ol a 

Paphiopedilum crossii var.sublaeve (Rchb.f.) Braem & Senghas, stat.nov. Basionym: 

Willi n h\ I I I i I hr II i 

1 ' ''I' I ■' - F'l'l 1 F.vJi. v'i In i 1 i i- t - 1 l I < ^ - 1 97 '4. Paphiopedilum 
sublaeve (Rchb.f.) Fowlie, Orchid I 
(Rchb.f.) Cribb, Genus Paphiopedilum 1 88. 1 987. 
Paphiopedilum thailandense Fowlie, Orchid Digest 43:220. 1979, nomen nudum. 
? Cypripedium callosum var. warnerianum T. Moore in Warner, Select Orchid. PI. 3, 1. 1 1 , 

ii var.potentianum (GruR & Roth) Braem & 5enghas, stat. 

■m: Paphiopedilum /_> - ' i ii n i L > IQQS Paphiopedilum 

ii/'i'feKM'iiHiiihtiil- I .(In it. I. i, mi > ,i h /,, , Jrnied 337. 1998. 

indebted to Peter H.Peeters for mi n. th if ( tK i .^tin. i t nlat< 

Ft I I til NC I ' 
em, G.J. 1988. Paphiopedilum - A monograph of all tropical and subtropical Asu. 
Hipper-orchids Bnu A ^ I FA lv im, Germany. 
em, G.J., CO. Bakfr, and MA. Baker. 1 998.The genus Paphiopedilum - natural history a 

hi i i I I I i mi I A I F f liami FF. 
fcM,GJ.,C.O.BAKiK,and M t F i i ] i e he o< tin lari.pia , i natural history a 

nil i- n .1 t mi i i m h I m| Miami, FL. 

BRAEM,GJ.,CO.BAKLR,and M.L.Baker (in preparation). The genus Pophiopedilum - natural 
history and cultivation, vol. 3. Botanical Publishers, Miami, FL 

Cribb, PJ. 1 987.The genus Pophiopedilum. Collingridge, London, UK. 

Cribb, PJ. 1998. The genus Pophiopedilum. Natural History Publications (Borneo), Kota 

Greuter, W, J. McNeil, F.R. Barrie, H.M. Burdet, V. Demouun,T.S. Filgueiras, D.H. Nicolson, O.C. Suva, 
J.E.Skog, P. Trehane, NJ.Turland, and D.L Hawksworth. 2000. International code of botani- 
cal nomenclature (Saint Louis Code). Koeltz Scientific Books, Kbnigstein, Germany. 

Gruss, O and J. Roth. 1995. Pophiopedilum potentianum, a new species from Thailand. 

1 5:226, 1. 17. 
Morren, C.J.E. 1 883. Note sur le Cypripedium barbatum, Lindl. et ses principals varietes, 

crossKwarnerianum, etc.Belgique Horticole 33:96-98, t. 7. 
Reichenbach, H.G. 1 886. Cypripedium callosum, n. sp. Gard. Chron. ser. 2, 26:326. 
Reichenbach, H.G. 1888. Cypripedium callosum (Rchb.f.) sublaeve, n. var. Gard. Chron. ser. 3, 

id W. Keller and Karl L. Braun. 1 999. Myxomycetes of Ohio.Their Systematics, Biol- 

ogy,and Use inTeaching. (ISBN 0-86727-1 33-7, pbk.).Ohio Biological Survey Bul- 
letin New Series volume 1 3 Number 2 (ISSN 0078-3994). Ohio Biological Survey, 
1315 KinnearRoad,Columbus,OH 4321 2-11 92, U.S.A. (61 5-292-9645,614-688-4322 

fax; $35.00 pbk.(Wire-0 Binding or Perfect 
Binding) xvi + 1 82 pp., 1 6 color plates. 

sm, sometimes several inches or more across, creeping in mass or in a vein-like network over 


itin l 1 ii i i it hi i, |h, paddles,these 

d cell-like amoebas that glide and IrrU bv cnuulling h,n icria 

30i. i. m; themselves into an amivol umorb,)'. that eventual v 

juctured and colorful spor 

e producing bodies that we can see easily with a 10 ■ hand 

the biology, morphology, and taxonomy of 

iii'm n N i 'I H l<\ hm • i i mi i in i tlvsu ,i 

of its personal touch about I he author 
? contributed to kno I. l.j i i|n M ,\ <>tes. But this book i: 
a than the state ot On fj. „| v ,' u,, world's 600 species a 

are included.and the conhituiiioir. oi ihe '.dentists mentioned h; 

odiversity catalogued? In what locality t 
Ticroorganisms,that one might find in a 




Section of Integrative Biology 

University of Texas 

AustinJX 7871 3, U.S.A. 

Guy LNesom 

North Carolina B~x r> , 
University of North Carolina 
19 1280 US 4 

?"se usa despues para agrupar vanedades 

Fritsch (1 997) has provided a much needed revision of Styrax for Mesoamerica, Mexico 
and western Texas, recognizing 19 species. One of these, S.platanifolius Engelm.exTorr, 
was treated as having five subspecies, bringing to 24 the number of formal taxa of Styrax 
recognized for the region concerned. Fritsch recognized infraspecific taxa as"subspe- 

concepts of Hutten (1 967) and Thome (1 978)," who "use the subspecies category for 
It 1 ,! > itic taxa that are geographically as well as morphologically distru. i." 

Of course, most current botanists who employ "variety" also use it in reference to 
infraspecific taxa that are geographically and morphologically distinct, and we do not 
accept what is essentially the equating of these two categories of infraspecific classifica- 
tion. The use of variety in plant taxonomy for the first infraspecific rank dates back to 
Linnaeus and has historical precedence over the term subspecies. More significantly, 

ptoin ah I i the hams (liaioiiiKi the print pal 01 as oiufdix I ml- A [J,,-,! , 
taxa of tt'ii- I >'l>v. r u Links |ii I, i i i.iii i,| ^iiu,- , . i, ,|i lU in MJ | ., ,iiiuir 
ill i i 1 | h i [i < iii i 1 . i s s i s . i I i a i ■ r mi 1 I t a ifi , 

Mihtienii'.a.tM [lo.oahas tiu.. sera "... aul 'Si.aaes, s| 'a a ■es,snbspri so variolas., mioss 

Philosophical and interpretive differences regarding use of infra-taxon categories are 
magnified by this tension in the ICBN: variety and/or forma are the ranks to be used 
first in describing infraspecific taxa (Article 4.1 ), but subspecies is the term first in 
hierarchical rank below species (Article 4.2). Use of species and variety, however, is 

analogous to that for general use of kingdom and division .class and order, family and tribe, 

and genus and section. In each of these pairs the first used sub-rank (the second term) is 
code-prescribed as such (4.1), with the orthographically subsidiary and immediately hi- 
erarchical ranks subkingdom sub< , . ' < .ouenus generally used for an ad- 
ditional rank (4.2), as is subspecies. 

Varieties may be clustered by use of the subspecies category 

Varieties are recognized within a species when it is desirable to refer by name to mor- 
pho i a a na 1 1 HI lea nhait I ei in compt inn tl I | i In our concept and 
experience, varieties usually are closely similar allopatric entities that intergrade over a 
relatively short distance in regions of contact (as opposed to gradual, broadly regional 
intergradation),if they intergrade at all. In addition to morphology and geography, other 
factors may be used in evaluation of taxonomic status of su< h entities -genetic diver- 
gence, likelihood of natural h i i.li_a n aili uln ft si ^tuessy 1990). 

In a species witci . eta I varieta no recognized, two or more varieties may be 
grouped within a subspecies. In this sense, use of the subspecies rank may point to larger 

egories finds support in the ICBN, which implies that the term subspecies is used for 
clustering varieties. 
Recommendation 26A 

uos|)ocaes rank in clustering var 
ious families and genera in the 

!'•-■ u t - , ' i," inenuear, 

Eriogonum, Heterotheca, Ipomops ,achaeranthera, Monardo, Prunu 

Ptelea, Ruellia, Salix, Salvia, Scutellaria, Sidalcea, Silene, Solidago, Streptanthu 
Symphyotrichum, and Tetramolopium). 

Other perspectives c 

The issue of "variety vs 
peatedly.Cronquist (1988) and Stuessy (1990) gave detailed overviews of the issue and 
Hamilton and Reichard (1992) provided a review of current practice in the use of in- 
fraspecific categories. Our commentary does not break new ground, but it emphasizes 
t is a reminder that usage of these categories remains in- 
thout explicit rationale. 

Views similar to ours have been well-expressed by Kapadia (1963) and Holmgren 
(1 994). In a contrasting view, Raven (1 974) proposed to simplify infraspecific terminol- 
ogy by using only the term subspecies, nomenclaturally equating the term variety, this 
proposal accompanied by detailed suggestions for its formalization in the ICBN. Another 
form of this latter solution is to use subspecies as the first category for infraspecific taxa— 
then to use varieties (at lower rank) for subsequent subdivisions of subspecies (see Stuessy 
1990, Fig. 12.1). For Thorne (1978, p. 190), "Genetic variants without well-defined geo- 
graphic ranges are treated as varieties ...."Such proposals, however, to formally displace 
"variety"as the first infraspecific category have not been accepted, presumably because 
many botanists find utility in maintaining two classificatory units at infraspecific rank 
and because the basis for use of"variety"is historical and currently codified. 

The rank of variety has been used to describe taxa over a range of evolutionary and 
morphological differentation.Some taxa are more strongly differentiated than others. 
Traditional views of boundaries between taxa at specific and infraspecific rank also differ 
among genera and families. Replacing variety with subspecies would not change this. 

The International Code of Zoological Nomenclature (International Commission on 
Zoological Nomenclature 1 999) does not provide for the term "variety"for classificatory 
purposes.The latter term, as used by most botanical taxonomists and as prescribed by 
the ICBN, is essentially equivalent to the subspecies rankof zoologists.This is acceptable. 
The botanical code is for plant workers,the zoological code for animal workers and the 
two codes need not become one. Actually, it is informative to see the term "variety" in a 
title or abstract: one knows that the organisms concerned are most likely plants. 

It might be argued that the term "variety" has been misapplied by various workers, 
especially horticulturists and plant breeders, to designate mere forms. But most profes- 
sional plant taxonomists use the term "forma" for such population variants, while the 
nternational Code of Nomenclature for Cultivated Plants (Trehane et al. 1 995) refers to 
such an individual plant or genetic strain as a "cultivar" (cultivated variety). The horticul- 
tural taxonomists coordinate their taxonomy with the ICBN, providing an adjunct system, 
not one that contradicts (Brickell & Trehane 1 997). In short, horticultural usage is not a 
tenable rationale for generally adopting the rankof subspecies to the exclusion of variety. 

^ pragmatic argu 

Articles 4 
puHk re- 
hearing, i 

on, according 


Article 3 


viboioomm oi 


io n ngw" 

Infraspecific taxa in Styrax platanifolius 

Evidence from morphology ami i em nil, h: m n n nilv a single species 
should be recognized among the populations of Styrax in west Texas and adjacent 
Mexico— S.platanito , nni|li oil nim 1 i mi i| l n o i tim ttaxon,apparently 
most closely related to the California endemic S. redivivus (Torr.) Wheeler (Fritsch 1997). 
While variation in pubescence quantity in other New World Styrax species is essentially 
random, Fritsch (199/, p./ 1 1) found thaf'trichome morphology or abundance within S. 
platanifolius is distinctly regional and facilitates the tf li mi tali n >i imaily or completely 
allopatric taxa. Therefore, I have recognized five subspecies within S. platanifolius based 
on minor but distinctive different es. richomet hammers aie those most reliable for the 
delimitation of thesi it | i- , i tm f eun i u n it im e include leaf form, 
calyx gland density, and sulfate leatures oi the stem,"chaiac lets considered by Fritsch 

Ihie< if the ' < ill- ' , U lie i 

relatively compact geographic cluster on the Edwaids Plateau of Texas; the other two 
{youngiae and mo/fa) ate king it ud i n.illv alig net J in sum ran lot alums from trans-Pecos Texas 
into northeastern Mexico. Fritsch did not mention any aspm t of ntergradation among 
these taxa— while it apparently is true that tfie distinctions aie fairly discrete, the popu- 
lations are rare, com| ^- f >1 ei . ni in ii mhi 'i| i ]i it ion can be seen as 
completely isolated within the rugged terrain. Fritsch's comment (p. 743) that "subspe- 
cies stellatu resemIT mlopi i< noil more closely than dm ubspecies youngiae" 
might suggestthat ( I | | o , n i - u I he considered together as a larger 

infraspecific unit, but tf would render this 

In sum, the morphological and evolutional, >mtio utt : ,, ific taxa of Styrax 

plalanilolius (sense I else h I mm) c niiespomils to we « uncoivo of here as varieties. 
Varietal rank is gen ml i 1 if ' t n 1 m 1 m itegory throughout 

sEngelm. ex Torrey var.platanifolius. Automatically 
the publication of Cory's var. stellatus, as listed below. 
/raxplatanifolius subsp. platanifolius. Automatically established by F 

c. Styrax platanifolius var. stellatus Cory, Madrono 7:1 1 
Styrax platanifolius subsp. stellatus (Cory ) P.W. Fritsch, , 

d. Styrax platanifolius var texanus (Cory) B.L.Turner, c 

Styrax platanifolius subsp. texanus (Cory) P.W. Fritsch, / 

e. Styrax platanifolius var youngiae (Cory) B.L.Turner, c 

youngiae Cory, Madrono 7: 11 3. 1 943. 
Styrax platanifolius subsp. youngiae (Cory) P.W. Fritsch, , 


are grateful to Peter Fritsch for comments on an e 

arly draft of the manuscript, to 

i Strother for perspectives on terminology and pars 

imonyand to Rogers McVaugh 

Brickell, C. and P.Trehane. 1 997.The RHS Advisory Panel on nomenclature and taxonorr 

New Plantsman 4:1 1 5-1 1 9. Also < .asp 

accessed October 2000. 
Cronquist, A. 1 988. The evolution and classification of flowering plants. Ed. 2. New Yo 

Botanical Garden, New York. 
Fritsch, P.W. 1997. A revision of Styrax (Styracaceae) for western Texas, Mexico, ar 

Mesoamerica. Ann. Missouri Bot. Gard. 84:705-761 . 
Greuter,W., F.R. Barrie, H.M. Burdet,W.G.Chaloner,V. Demoulin, D.L. Hawksworth, P.M. Jorgense 

D.H. Nicolson, PC Suva, RTrehane, and J. McNeill (eds.). 2000. International code of botar 

cal nomenclature (Saint Louis Code). Koeltz Scientific Books, Konigstein, Germany. 

HAMiiTON,C.W.and S.H. Ri , ,m>. 1992. Current practice in the l 

and forma in the classification of wild plants. Taxon 41:485— 
Holmgren, N.H. 1994. Redefinition of Dodecatheon dentatum (Prir 

for use of varietal rank. Brittonia 46:87-94. 
Hulten, E. 1 967. Comments on the flora of Alaska and Yukon. Ark. Bot„ n.s. 7:1 -1 47. 
International CoMMr k u u 7 iN - i i ,ilt >n International code of zoological 

nomenclature (ed.4). International Trust for Zoological Nomenclature, London, U.K. 
Kapadia,Z.J. 1 963. Varieties and subspecies: A suggestion towards greater uniformity. Taxon 

Raven, P.H. 1974. Proposal for the simplification of infraspecific terminology. Taxon 23: 

StuessyJ.F. 1 990. Plant tax. >m >my: I he s\ .tematic evaluation of comparative data. Colum- 
bia Univ. Press, NY. 

KMi.lvl.l New subspecific < >mbi iti n fo uihem< ihfornia plants. Aliso 

Trehane, P., CD. Brickell, B.R. Baum, W.L.A. Hftterscheid, A.C. Leslie, J. McNeill, S.A. Spongberg, and 

6).Quarterjack Publishing,V 


Recent molecular evidence indicates that most genera of North American Astereae comprise a 
monophyletic assemblage. Within this f ' i h American « I i I are ihi li tin t i iihi i i| 
previously hypothesized to be n the Southern Hemisphere— the 

Monoptilon group.the Pentachaet i v | ith ■ a qroup. Additionally, morphological 

and molecular evidence indicates tlnttb „ M » oracantha.and Batopilasia are closely 

related among themselves and constitute a coherent group. These four North American groups 
are recognized here with formal names at subtribal rank:Chaetopappinae subtr.nov. {Chaetopappa, 
Monopt/Von), Pentachaetinae suhtr rr ' . nt ■ ' - Astranthiinaesubtr. 

Dov.iAstranthium.Dichoetophortider, e/ - n id Boltoniinae (Batopilasia, Boltonia, 

China n an\ 

■: MTv'FN 

Adicionalmente, la evid 

encia morfologic.i y rnoiec 

;ular indican que los generos Boltonia, 

Chloracantha, y Batopilask 

i estan muy relacionados ent 

■.][,■ ron nombres formales en el rango: 

Chaetopappinaesubtr n 

ov. {Chaetopjpt 

), Pentachaetinae subtr. nov. {Pentachaeta, 

Rigiopappus, Tracyina), Astr 

ium, Dichaetophora, Geissolepis, Townsendia), 

y Boltoniinae [Batopilask 

i, Boltonia .China i a •> d 

ectotipifica Homochrominae, colocandola 

A primarily morphological overview of the tribe Astereae (Nesom 1 994a) recognized 1 4 
subtribes, four of which were hypothesized to be primarily North American 
(Machaerantherinae, Chrysopsidinae, Solidagininae, and Symphyotrichinae). Three dis- 
tinctive North American generic groups were hypothesized to be most closely related 
to subtribes of the Southern Hemisphere— the Monoptilon group (subtribe Feliciinae = 
Bellieae, primarily Africa and western North America), the Pentachaeta group (subtribe 
Bellieae), and the Townsendia gnu i hnU-'n h ,cominae, primarily Australia and 
North America).The genus Boltonia was hypothesized to be related to genera of subtribe 
Asterinae, primarily an Eurasian group (Nesom 1 994a, 1 994b). 

Noyes and Rieseberg (1999) used nucleotide seguence data from nuclear riboso- 
mal DNA representing a broad range of Astereaean genera to test various hypotheses of 

relationship and classification in the tribe.Their results showed that all genera examined 
of North American Astereae comprise a single, strongly supported clade^thus mor- 
phological parallelism rather than homology underlies hypotheses suggesting that dis- 
persal from the Southern Hemisphere and Asia accounts for the origin of the Monoptilon, 
Pentachaeta, and Townsendio groups and Boltonia. 

Notwithstanding these unexpected patterns of relationship, the infra-tribal generic 
groups under consideration have consistently been recognized as coherent (see cave- 
ats below regarding Aphonostephus, Geissolepis, and Bo/ton/a). The morphological dis- 
tinctiveness of these groups is egual to others already n 
with recognition of their origin from within the broad 
provided here with formal names at the same rank. 
ChaetopappinaeNesom,subtr.nov. bn <,iu\r,:Chaetapai 

Small, annual or short-lived perennial herbs, taprooted, decumbent (Monoptilon, some 
Chaetopappa) to erect. Leaves entire, oblong to oblanceolate-spatulate, alternate. Heads 
mostly solitary; phyllaries flat to convex, with broad, sharply delimited, hyaline margins. 
i Hi I I" t( hiti n m il\ i dling Disc flowers sometimes with sterile ovaries; 
style branches with obtuse or truncate to triangular collecting appendages. Cypselae 
eglandular or glandular, terete and multinerved (most Chaetopappa) or obovate, flattened, 
and 2-nerved (Monoptilon, some Chaetopappa); pappus of persistent bristles, or scales, 
or pales, or of bristles and scales, commonly in multiples of 5 (in Chaetopappa), or absent. 
Base chromosome number, x = 8. Genera included: Chaetopappa DC, Monoptilon Torr. & 
A. Gray ex Gray. Distribuiio piimaiil i ha tat in the southwestern and south- 
central USA and northern Mexico. 

This is essentially the "Monoptilon group," earlier placed in subtribe Feliciinae = 
Bellieae (Nesom 1994a) and as "Incertae sedis" (Nesom 2000). The two genera of 
Chaetopappinae form a monophyletic group sister to Euthamia Nutt.ex the Noyes 
and Rieseberg analysis. In the analysis of Lane et al. (1996), the phyletic origin of 
Chaetopappa lies imm I t I 1 t < i > ind Townsendia, although it does 

not do so in the original analysis from which the DNA data were drawn (Morgan 1 990). 

The southern European genus Bellium L. was earlier included in the Monoptilon 
group (Nesom 1 994a), but with the strong indication that Monoptiloi 
are North American in origin, it is probable that the closest relation: 
with the other Old World genera. Similarities between Bellium and An 

Pentachaetinae N 

florum disci. Cypsela* 
pappus setarum vel 

Annual herbs, taprooted. Leaves alternate,entire,filiform or linearto narrowly oblanceolate. 
Heads solitary and long-pedunculate; phyllaries with hyaline margins. Ray corollas yel- 
low to reddish, less commonly white, strongly coiling (lamina absent in some Pentachaeta). 
Disc flowers: style branches with linear-lanceolate collecting appendages. Cypselae ter- 
ete to slightly compressed, narrowly oblong to oblanceolate in outline,beaked in Tracyina, 
eglandular; pappus 1 -seriate (1-2 seriate in Tracyina), of persistent bristles frequently in 
multiples of 5s, sometimes flared at the base and partially connate, or sometimes com- 
pletely lacking, or of long scales {Rigiopappus). Base chromosome number, x = 9. Genera 
\nc\uded:PentachaetaNun.,Rigiopappus A.Gray, Tracy/na S.F.Blake. Distribution primarily 
in grassland habitats of California, USA, with several taxa of Pentachaeta reaching Baja 
California, Mexico; Rigiopappus also occurs into Oregon, Washington, ldaho,and Nevada. 
This is the "Pentachaeta group/earlier placed in subtribe Feliciinae = Bellieae (Nesom 
1994a) and as'lncertae sedis" (Nesom 2000). The close resemblance and relationship 
among these three genera have been noted by Blake (1937),0mduff and Bohm (1975), 
and Robinson and Brettell (1973); they were placed as a coherent unit within the 
"Chaetopappa group" by Bremer (1 994).The Pentachaetinae form a monophyletic group 
sister to Ericameria Nutt. in the Noyes and Rieseberg analysis. 
Astranthiinae Nesom, subtr. nov.TvPF genus: Astranthium Nutt. 

Herbae annuae biennes vel perennes plerumque radice palari. Capitula solitaria, plerumque 
longipedunculata;phyllaria marginibus late hyalinis; receptacula convexa vel conica.Corollae radii 
plerumque albae vel caeruleae, midfascia abaxiali lavandula, non reflexae aut circinnatae.Corollae 
discii tubo brevi. Cypselae oblanceolatae vel obovatae, complanatae, 2(-3)-nervatae, laeves vel 
papillatae, plerumque pubescentes trie i i matibi , hidiatis;pappus setarum vel corona demissa 

Annual, biennial, or perennial herbs, taprooted, often with a branching caudex, rarely 
fibrous-rooted, mostly strigose with short, white hairs. Leaves alternate, spatulate to linear, 
entire or few-toothed. Heads solitary, mostly long-pedunculate, rarely sessile; phyllaries 
with broad, hyaline margins; receptacles convex to conical. Ray corollas white to bluish or 
pinkish above (rarely yellow in To wnsendia), usually with a lavender to blue or pink abaxial 
midstripe, not reflexing or coiling. Disc corollas short-tubed; style branches with triangular- 
lanceolate collecting appendages. Cypselae oblanceolate to obovate, flattened, 2(-3)- 
ribbed,the surfaces smooth or papillate,glabrate or usually pubescent with duplex hairs 
with glochidiate, bifurcate, or entire apices (achenes winged and fringed-ciliate in 
Dichaetophora); pappus 1 -seriate, of barbellate bristles or a low crown of setae/bristles 
and scales (2-awned in Dichaetophora). Base chromosome number,x = 9 (or x = 3,4,and 5 
\r\Astranthium;x = 3 in Dichaetophora). Genera iuclucie&.Astranthium Nutt., Dichaetophora 
A. Gray, Geissolepis B.L Rob, Townsendia Hook. Distribution mostly in western North 
America north of Mexico {Townsendia), Mexico and south-central USA {Astranthium),Jexas 
and adjacent Mexico {Dichaetophora), and e 

This is the "Townsendia group," earlier placed in subtribe Brachyscominae (Nesom 
1 994a) and as'lncertae sedis"(Nesom 20Q0).Aphanostephus DC.also was earlier included 
in the Townsendia group (by Nesom and various others), but molecular data from 
eral sources (Morgan 1 990; Lane et al. 1 996; Noyes and Rieseberg 1 999; Noyes 2000) < 

Dichaetophora, and Townsendia comprise a monophyletic group essentially sister to the 
Conyzinae and Chrysopsidinae in the Noyes and Rieseberg analysis. Geissolepis is p 
etically interposed between the Townsendia group and these two related subtribes, 
a set of morphological features places it closer to the Townsendia group than the 
Conyzinae or Chrysopsidinae, and the genus can reasonably be included in the 
Astranthiinae. Alternately, it presumably would be treated as a monophyletic subtribe. 
Geissolepis is similar to the Astranthiinae "core" genera in its solitary heads, white, 
straight ray corollas, conical receptacles, and glochidiate cypselar vestiture. It is distinct 
within the subtribe in its combination of a prostrate habit with creeping,fibrous-rooted, 
lignescent stolons, succulent leaves, resin canals on the phyllaries, cypselae, and disc 
corollas, paleate receptacles, ray corollas without an abaxial midstripe.gradually ampliate 
disc corollas, subterete cypselae with 8 resinous ribs, and pappus of short scales with 
argins.The chromosome number has been reported as 2n = 1 6 (Ralston 

Boltoniinae Neson ubtr.nov.lvi ,i nus Boltonia L'Herit. 

us ac foliis persistente viridi-glabratis. Folia 

iria vel laxe aggregata; phyllari; 
resinosis.Corollae radii albaeve 

Perennial, herbs or subshrubs {Chloracantha), rhizomatous, with persistently green- 
glabrate stems and leaves, thorny in Chloracantha. Leaves essentially all cauline, entire or 
few-toothed. Heads solitary or very loosely corymboid to paniculate; phyllaries primarily 
herbaceous, apically rounded to obtuse, with three orange-resinous nerves. Ray corollas 
white to slightly bluish, coiling. Disc corollas orange-veined; style branches with deltate 
collecting appendages. Cypselae terete and multinerved or flattened, 2-nerved, and 
winged (Boltonia). Base chromosome number.x = 9. Genera included:8oto/3/7os/'o Nesom 
& Noyes, Boltonia L'Herit., Chloracantha Nesom, Suh, Morgan, Sundberg, & Simpson. Dis- 
tribution in northwestern Mexico {Batopilasia), eastern USA {Boltonia), and Mexico and 
the southwestern USA to Louisiana (Chloracantha). 

Batopilasia, Boltonia, and Chloracantha apparently are closely related among them- 
selves (summary of ideas and evidence in Nesom & Noyes 2000). Batopilasia and Boltonia 
are sister genera in the Noyes and Rieseberg analysis, the pair in a sister relationship to 
the Symphyotrichinae and Machaerantherinae. Chloracantha was not included in this 
molecular analysis but is morphologically similar to Batopilasia, as observed in the origi- 
nal description of the species (Sundberg & Nesom 1990). 

Boltonia is set apart from the other two genera by its conical or convex receptacles, 
short-tubed disc corollas,and flattened,2-nerved,orange-veined, often winged cypselae 
with an abbreviated pappus. These specializations prompted the observation that 
"Boltonia is morphologically isolated in the New World" (Nesom 1 994b, p. 1 63), but mo- 
lecular evidence has found its close relatives. It is related neither to Old World Astennae 
(Nesom 1 994b) nor to genera of the Townsendia group (Bremer 1 994). 

With the formal recognition of the four subtribes above, six North American genera 
of Astereae remain without a clear hypothesis of subtribal affinity.These are among the 
"'primitive' Asters" (Nesom 2000), placed by Nesom (1994a, 1994b) mostly in subtribes 
Asterinae or Symphyotrichinae: Doellingeria Nees, Eucephalus Nutt., lonactis Greene, 
Oclemena Greene, Oreostemma Greene, and Tonestus A. Nelson. 


Homochrominae of Bentham & Hooker as invalid 

published— it was provided with a description a 

based on a legitimate genus name (Homochroma 1 

among those genera they placed in the subtribe. F 

the nomenclaturally"sister"taxon to the illegitimate Heterochrominae Bentham 8 

(this name not based on an included genus), but a reasonable lectotypificc 

Homochrominae already has been effected by Solbrig (1 963), who specified th 

Homochrominae Bentham & I 

Hooker,Gen.Pl.2:1 74.18731 

ECTo™ E :(Solbr 

DC. (= Zyrphelis Cass.). 

With this typification, Homochrominae becomes a syno 

nym of Belli. 

Gren. & Godr, Fl. France 2:83, 

1 04. 1 850; type, Bellium L 

.) along wit! 

(Phytologia 76:205. 1994; type 

.Felicia Cass.). 


1 apprecate the helpful comn 

nents of Ted Barkleyand K. 


1 937. Tracy ina, a new c 

1:73 ■ 

■tics and classification. Timber Press, Portland, Oregon. 

ANF,M.A.and J. Li. 1993. Chromosome number reports in Compositae with emphasis on 

tribe Astereae of the southwestern United States and Mexico. Sida 1 5:539-546. 
ANb, M.A., D.R. Morgan, Y. Suh, B.B. Simpson, and R.K. Jansen. 1996. Relationships of North 

American genera of Astereae, based on chloroplast DNA restriction site data. In D.J.N. 

Hind and H.J. Beentje (eds.). Compositae: Systematics. Vol. 1, pp. 49-77. Proc. Interntl. 

Compositae Conf, Kew, 1 994. 

using restriction site analysis of chloroplast DNA and a taxor 

Machaeranthera section Psilactis. Ph.D. dissertation, Univ. of Texas,/ 5 
Nesom, G.L 1994a. Subtribal classification of the Ast 

Nesom, G.L. 1994b. Review of the taxonomy of Aster 

emphasizing the New World species. Phytologia 7 
Ni si >m,( ..1 . 2000. Generic conspectus of the tribe Astei 

Central America, the Antilles, and Hawaii. Sida, Bot. 
Nesom, G.L. and R.D. Noyes. 2000. Batopilasia 

Chihuahua, Mexico. Sida 19:79-84. 
Noyes, R.D. 2000. Biogeographical and evolutionary i 

(Asteraceae) from ITS sequence data. PI. Syst. Evol. 22( 
Noyfs, R.D. and LH. Riesfbfrg. 1999. ITS sequence data s 

d reflect deep geographic 


8 41 .T 

Ornduff, R. and B.A. Bohm. 1 975. Relationships of Tracyina ar 

Ralston, B., G.L. Nesom, and B.L.Turner. 1 989. Ch romosome nu r 

with special reference to the tribe Tageteae. Sida 1 3:359 
Robinson, H. and R.D. Brettell. 1973. Tribal revisions in the Ast< 

Rigiopappus. Phytologia 26:69-70. 
Solbrig,O.T. 1963.Subfamilial nomenclature of Compositae.Taxon 12:229-; 
Sundberg, S.D. and G.L. Nesom. 1 990. A new species of Erigeron (Asteraceae: A 

Chihuahua, Mexico. Phytologia 69:278-281 . 

The relationship c 


Fairchild Tropical Garden Research Center 
1 1 935 Old Cutler Road, Miami, FL, 33156-4299, U.S.A.; 

Jon M.Ricketson 

Missouri Botanical Garden 

RO. Box 299, St. Louis MO 63166-0299, U.S.A. 


nily Myrsmaceae for the florula of the "Rio Cenepa" region 
;s in the genus Stylogyne.Stylogyne 
id and its phylogenetic relations are 


The neotropical genus Stylogyne A. DC. comprises approximately 60-70 species, a num- 
ber of which remain undescribed, owing to lack of adequate material. Understanding 
the systematic biology of this genus has long been problematic because of its sexual 
liability. Androdioecious, bisexual, polygamo-dioecious and dioecious species of Stylogyne 
have been documented (Pipoly 1 989, 1991), as well as the consequent morphological 
variation due to sex expression. Since no comprehensive study of the genus Stylogyne 
has occurred since C.Mez's treatment in Engler's Das Pflanzenreich (1902), the genus is 
in need of additional study. Our current studies continue in the genus Stylogyne (Pipoly 
& Ricketson 1 999; Ricketson & Pipoly 1 997), as well as the entire family for our treatment 
of the Myrsinaceae for Flora Neotroplca. 

During preparation of a manuscript for the Florula of the Rio Cenepa Drainage Ba- 
sin Project of the Missouri Botanical Garden, a new species was found and is described 


n t ii i - ' - , lot, ii v h, , n laiedistat. 

free or small tree 4-8 m tall, to 5.7 cm in diam. Bronchlets ca. 1 mm in diam., terete, the 
bark yellowish-brown, longitudinal H koH , ,l,i ,. >u\hollow,densely lenticellateleaves 
pseudoverticillate; blades chartaceous, oblong to oblanceolate, 32-39 cm long, 5.5-1 3.4 
cm wide,apically acute to obtuse, basally tapering gradually to petiole base, the midrib 
channel prominently raised above, decurrent on the petiole/stem junction, the midrib 
prominently raised below, the secondary veins numerous, brochidodromous, 

below, the hydropotes scattered, bright orange below, the margin flat, entire; leaf base 
and petiole,when distinguishable,deeply canaliculate,obsolete to 1 mm long,the petiole 
margin often abruptly tapered at petiole apex to almost appear auriculate, deeply 
canaliculate and marginate, glabrous. Staminate inflorescence and flowers unknown. Pis- 
tillate inflorescence lateral, a condensed raceme, 3-10 mm long; floral bract girdling pe- 
duncle, chartaceous, elliptic, 2.7-3 mm long, 1 .3-1 .5 mm wide, apically acute, densely 
and prominently orange punctate and punctate-lineate,the margin irregular, somewhat 
erose apically otherwise entire; pedicel cylindrical, 2.5-3 mm long. Pistillate flower 5- 
merous, white; calyx carnose, membranaceous, 2.5-2.7 mm long, the tube ca. 0.2 mm 
long, the lobes nearly free, ovate, 2.3-2.5 mm long, 1 .5-1.7 mm wide, apically obtuse, 
with one or two orange punc tations medially, glabrous, the margin hyaline, entire; co- 
rolla membranaceous, 3.8-4.2 mm long, the tube 2-2.2 mm long, the lobes connate 
basally, ovate to lanceolate, 1 .8-2.2 mm long, 1 -1 .2 mm wide near the base,apically acute, 
conspicuously orange punctate and punctate-lineate,glabrous,the margins entire, hya- 
line;stamens 3.7-4.3 mm long,the filaments 2.8-3.2 mm long,filamentous,free,epunctate, 
glabrous, the anthers free, narrowly ovate to lanceolate, 1 .2-1 .3 mm long, 0.5-0.6 mm 
wide at the base,apically emarginate, basally umHte, Oncntuc km, illy dehiscent by slits, 
the connective conspicuously punctate; pistil obturbinate, 3-3.3 mm long, 1 .3-1 .5 mm 
in diam, glabrous, the ovary 1 .5-1 .7 mm long, the style 1 .3-1 .5 mm long, the stigma 
punctiform, the placenta cotyliform, with 4 open chambers above, the ovules 4,exposed. 
Fru/'tglobose,5-6.2mmlong,5-6.2mmdiam apicallytrun te,r I at maturity, densely 
and prominently pollu i Him ate punctate, the exocarp thin. 

Distribution.— Stylogyneaguarunana is known only from Imaza District, Bagua Prov- 
ince, Amazonas, Peru, in the Rio Marahon Drainage Basin around the Comunidad 
Yamayakat (Fig. 2), from 300-480 m. 

Ecology and conservation status. —Stylogyne aguaninana H know from only four 

11 i i' Ii i ih [ ' i t ih Mm i i | I h inhabitthearea. 

Fig. I.Sfy/ogyne^oownonoPipoly&Ricketson. A. Flowering 

late inflorescence. D. Detail of pistillate flower. E. Fruit and fruiting inflorescence. 

etal. 296 (MO). E drawn from C Diazetal. 7895 (MO). 

I t PERU..'. 


/. Bagua, Distrito Imaza, Comunidad Aguaruna Yamayakat, can 

)96 (young bud), £ Rodriguez et al. 1 167 (CPUN n.v., I l"G, MO); I 
at, 05° 03' 20"S, 078° 20' 23"W, 380 m, 6 Nov 1 996 (ster.), R. Vasqu, 

Fig. 2. Distribution of Stylogyne aguarunana Pipoly & Ricketson (•), in the area around the Comunidad Yamayakat, n 

The long leaf blades, that are nearly sessile and pseudoverticillate, clearly distinguish 
Stylogyne aguarunana from all other members of the genus.The multibracteate inflores- 
cence rachis is similar to those of Stylogyne bracteolata, but can be easily distinguished 
from that species by its smaller or obsolete petioles, and 5-merous flowers. Stylogyne 
aguarunana belongs to a group of taxa with 5-merous flowers and anthers and 
onq spindly liLimenls, generally 3 ormoie times longer than the anthers. 

Missouri Botanical Garden, the Flora Mesoamericana Project, and the 
ilGarden,forfunding that allowed J. Ricketson (MO) to visit J. Pipoly (FTG) 


us in our work, including Gerrit and Jeany Davidse, Linda Oestry, Mary Bard, and Catherine 
Mayo (MO), Barney Lipscomb, Debra Track and Jim Rivers (BRIT). Illustrations were pre- 
pared by the junior author. Reviews of the manuscript by Gerrit Davidse and Roy Gereau, 
and meticulous copy editing by Barney Lipscomb, improved the presentation of the 

Mez, C 1 902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich IV. 236(Heft 9):1-437. 
Pipoly, J.J. 1 989. Notas sobre el genera Stylogyne A. DC. (Myrsinaceae). Ernstia 53:1 -9. 
PipOLY,JJ.1991.Sfy/ogyneradr/gues/ono(Myrsinaceae):a new androdioecious species from 

Amazonia. Novon 1:202-203. 
Pipoly, J.J. and J.Ricketson. 1 999. Novelties in the Myrsinaceae from the Venezuelan Guayana. 

Sida 18:1167-1174. 

Karstfn H.K.Wodr!ch. 1 997.Growing South African Indigenous Orchids. (ISBN 90-541 0- 
650-6, hbk.). Rotterdam (in USA: A.A. Balkema Publishers, Old Post Road, 
Brookfield, VT 05036-9704, U.S.A (Fax 802-276-3837, e-mail, $85.00 hbk., 253 pp., 94 color and 109 b&w photos and 
lh In mtifulh In ti m I , lum i 1 I ut in ' i ii ,ndph on how to grow South African 
orchids, from summ urn il In n t n In I, r i In ir t n h | iiuijlir t 1 

ting up a tissue culture homo. Although, the foe us imhi spec 10, native South Africa, much ol 
the general discussion can be applit ti 'i il ultun l i , i In I I il m the book, the discus- 
sion moves from genu If ii hi In t ii I ind climates in South 
Africa. Next is a general discussion of design ol i uliur i . i inl n home, orchid physiol- 
ogy and nutrient reguii< muil id m ml < i ud Ii i ih n, t I uipter looks at the three 
major growth forms of or. Inch and the hoilu . ultura! reguirements common to each. Within the 
treatment of the respective growth habits, each genus and species is broken out for specific in- 

photograph of its inflorescence or individual flov 

^er. There are sections on orch 

propagation using culture media both with and \ 

pll meter. An appendix even includes desions fo 

laminar-flow hood. The author does not forget cor 

vation legislation.The book is so complete and wel 

l-designed that I loumi only on. 

the inks contain a volatile compound That has the 

odor of overly ripe ouava Ihr 

■re and would n 

library.— Roqet IV k 

nager, Botanical -A 




Jon M.Ricketson 

t free apical portions of the filaments, anthers whose wide longitu- 
t c rom each theca meet apicalty to form a subcontinuous opening at anthesis, short and 
conical flower buds.and relatively numerous ovules on the rather thick placenta, precluded 
:rment in any known neotropical subgenus of Ardisia. Curiously, the aforementioned fea- 
;mprise the diagnostic character states unique to the Indo-Malesian Ai 

indSf) i i ia Because subgenus Stylardisia is defined by its protogynous flowers (the 

rang the bud apex long before the bud opem i < I n i< . ' .^iiiv ■ mf lit m 

up and therefore placed in subgenus Acrardisia Mez.We suggest that Ardisia rarescens is a 

' and that this represents another species whose distribu- 

c relationships, ecology a 

a por hendiduras longitudin 

n embargo' 

itneel ill I vj ,,| 

The pantropical genus Ardisia Sw. is by far the largest in the family Myrsinaceae, contain- 
ing perhaps as many as 500 species (Chen & Pipoly 1 996). Its circumscription has been 
problematic owing to a lack of comprehensive treatment since that of Mez (1902) in 
Engler's Pflanzenreich, almost a century ago. While reviewing the status of Ardisia rarescens 
Standi., we observed the combination of:dextrorsely imbricate petals, extremely short 
free apical portions of the filaments; anthers whose longitudinal slits from each theca 
meet apically to form a subcontinuous opening at anthesis;short, broadly conical buds, 
and relatively numerous ovules on the rather thick (thicker than long) placenta, all char- 
acteristics of Ardisia subgenus Acrardisia Mez. With subgenus Acrardisia's distribution from 
Sri Lanka eastward through Malaysia and Indonesia to the Philippines and New Guinea, 
it is strikingly similar to that of Hymenandra (A. DC.) A. DC. ex Spach, a group that we 
suspect is of boreotropical origin (Pipoly and Ricketson 1 999). A distribution such as this, 
most closely fits those groups cited as partial evidence to support what Wendt (1 993) 
discovered in his study of lowland Mexican wet forests, and what Lavin and Luckow 
(1 993) attributed to the Boreotropics Hypothesis, proposed by Wolfe (1 975) and Tiffney 
(1985a, b) to explain the distribution of the "boreotropical flora."The hypothesis pro- 
poses that the biotas of North America and Europe, including tropical North America, 
were once more widespread in the northern hemisphere and transgressed the North 
Atlantic by direct land connections or over limited water gaps until the late Eocene or 
early Oligocene (Lavin & Luckow 1 993). A logical extension to Wolfe's and Tiffney's con- 
cept, presented by Wendt (1 993) and Lavin and Luckow (1 993) is that the boreotropical 
flora not only existed in the Eocene, but also left a significant number of direct descen- 
dant lines in present lowland tropical floras of northern Latin America. Therefore, we 
would expect that many of the early Tertiary fossil taxa from both North America and 
Europe were most closely related to extant species from tropical southeast Asia, and to 
some extent, Central America and the Greater Antilles (Lavin & Luckow 1 993). It is the 
latter notion that is congruent with the Amphipacific distributional pattern like that found 
in our new concept of Ardisia subg. Acrardisia. Jh\s extension to the boreotropics hy- 
pothesis was cited by Wendt (1988, 1989, 1993), in discussing the relationships of 
Chiangiodendron (Flacourtiaceae),and by Zona (1990) in discussing the biogeography 
of Sabai (Arecaceae). A similar distribution for the genus Alstonia (Apocynaceae) was 
cited by Gentry (1983), but he did not invoke the hypothesisperse.Conran (1995), in his 
study of the Liliiflorae, found that three taxa defining the Southeast Asian/northern 
Australasian clade (Stemonaceae, Hanguanaceae and Uvulariaceae) were widespread 
northern taxa that have spread southwards. 

The geographic distribution of subgenus Acmidi\ia is eniimly consistent with the 
area cladogram presented by Lavin and Luckow (1 993, Fig. 1 ), where Central and South 
American elements are ultimately derived from among diverse North American lineages, 
these lineages having a sister group relationship to paleotropical groups. While Wolfe's 

(1 975) hypothesis could be correct even if no modern descendants of the boreotropical 
flora were found in the Neotropics owing to extinction, the three criteria proposed by 
Lavin and Luckow (1993) to test the hypothesis were: 1) a center of diversity in North 
America (including "tropical North America" as they define it), 2) an early Tertiary fossil 
record in North America, and 3) a pantropical distribution. 

For the first test criterion, "North America" includes both tropical and temperate 
elements,the tropical ones south of theTropic of Cancer. In the case of subgenus Acrardisia 
it is clear that only one species is presently known, but until we have tested hypotheses 
of phylogenetic relationship among other related Ardisia species, in particular the group 
described by Lundell as lbarrea,and containing the rather common species Ardisia 
paschalis Donn.Sm.,we cannot be sure. Subgenus Acrardisia itself may be paraphyletic, 
because the only characteristics separating it from the extremely closely related subge- 
nus Styiardisia Mez are that it does not have a stigma that perforates the flower bud and 
is receptive before the flower opens (thus protogynous), and that its ovules are pluri- 
rather than uniseriate. A group even partially defined by the lack of a structure can be 
problematic because it is not clear if protogynous flowers have occurred more than 
once within the genus Ardisia as a whole. 

Unfortunately, no data is available for use in the second test criterion owing to lack 
of fossils known for the group. As for the last criterion, a pantropical distribution, the 
closest related group, Ardisia subgenus Styiardisia, is concentrated in Borneo, but occurs 
from northern India and Bangladesh through Indochina, eastward through Malesia as 
far as Sulawesi, with no neotropical members known at this time. However, the entire 
genus Ardisia is truly pantropical, as is the tribe Ardisieae. Until a phylogenetic analysis is 
complete for the tribe, a rigorous test cannot be performed. Despite the absence of a 
cladogram, the pattern of distribution among the genera of the tribe Ardisieae fit the 
general pattern one would expect if the distribution was boreotropical. 

In summary, we hypothesize thatyArd/s/'a subgenus /Acra/-d/5/'o appears to fit the overall 
pattern consistent with a boreotropical distribution. We must emphasize that until a 
phylogenetic analysis among the subgenera of the genus/W/s/a,and more importantly, 
among the genera of Myrsinaceae is complete,there is no reliable way to unequivocally 
determine if the group is boreotropical or Gondwanan. However, at this point in our 
work, we find it useful to point out the strong correlation and call attention to the value 
of examining generic limits on a worldwide basis when preparing treatments for a large 
flora such as Flora Mesoamericana. 

Ardisia rarescens is known only from 27 herbarium specimens, and apparently is 
restricted to the eastern slope of the Sierra Madre de Chiapas, from central-southern- 
most Chiapas,Mexico,along the slope and into northern San Marcos and Quetzaltenango 
Departments, Guatemala.This region, including the famous VolcanTacana, houses many 
endemics and a myriad of taxa with boreotropical affinities, particularly in the pine-oak 
and Liquidarnbar forests in the region. 


i iiit.iti in iiilniii I iN i i itit description for floral partsar 

from organs rehydrai I nn h nut | i i u I I ilum •. iter Measurements 
from these range from 10% to 15% greater than those measurements taken directly 
from dried material. Data regarding stem diameters, inflorescence rachises, pedicels, leaf 
and fruit shape and size were taken from dried herbarium specimens. 

Morphological terms in this treatment follow Lindley (1 848) and Pipoly (1 987, 1 992) 
for the inflorescence, rachis pedicels and floral parts. Description of leaf morphology 
follows Hickey (1 984), trichome description follows Theobald et al. (1 984) and basic cell 
and tissue terminology follow Metcalfe (1 984). 

Ardisia Swartz subgenus Acrardisia Mez, Pflanzenr. IV. 236(Heft 9):1 16. 1902. 

Subshiub i ) m ill u i i n 111 ,1 in I j| n ipill id I in ul hirlHI >u i iru nm 
ous tomentose or rarely, glabrous. Leaf blades mostly entire or obscurely crenulate, but 
never regularly serrate. Inflorescence terminal, subtended by a foliaceous bract, the 
branches corymbose to umbellate or rarely racemose in fruit. Flowers 5-merous; buds 
shortandbroadlyconical;sepalsandpetalsdextioi ,.'K iinhri v. ,i miens with extremely 
si mil apical portions ol (lie filaments, anthers dehiscent by wide, apically confluent lon- 
gitudinal slits; pistils not protogynous, the ovules numerous, plunsenate. 

Distribution.— Appioximatel > p< > , thioughout ndo-Malesia,with onedisjunct 
at the border of Mexico and Guatemala. 

Ecology.—- Premontane to montane humid to wet forest,often on calcareous or sand- 
stone-derived soils, 500-2400 m elevation. 

Ardisia rarescens Standi. (Fig. 1 ), Publ. Field Columbian Mus„ Bot.Ser.4:248. 1 929. Amatlania 
rarescens (Standi.) Lundell,Wrightia 7:40. 1982 Typi MEXI< O.Chi i Tenodel Boqueion, 
withoutelevation,Sep1913(fl)C.APurpus7032(HOLOTYPc:F,Fnegnot,H24- i, i 

Shrubtosrnall trees 3-1 2 mtaW.Branchlets slender, terete, 2-5 mm in diam, densely glan- 
dular-papillate, glabrescent. Leaves loosely pseudoverticil late; blades chartaceous, elliptic 
to slightly oblanceolate, 3.5-1 6.2 cm long, 1.1-5.3 cm wide, apically acuminate, the acu- 
men 0.5-1.4 mm long, basally acute to cuneate, decurrent on the petiole, midrib im- 
pressed above,prominently raised below,the secondary veins 1 5-27 pairs, slightly raised 
above and below, prominently black punctate and conspicuously punctate-lineate,gla- 
brescent above and below, the margins entire, revolute; petioles slender, canaliculate, 
3.5-10.2 mm long, glabrescent above and belo • //v. «.iui nnal,erect,bipinnately 

or tripinnately paniculate, 6-22.5 cm long, 5.5-20 cm wide, pyramidal, usually longer 
than the leaves, densely, erect glandular-papillate, the branches terminally congested 
into / I il i - I ml |x iui i k i) - in I (.], inlloi > IihiuhIii n 

inflorescence branch bracts foliaceous, chartaceous, ovate to oblong, 2.3-3.7 cm long, 

Fig. l./lrrf/s/'flrflresfens Standi. A. Flowering branch. B. Detail of abaxial leaf surface. C. Detail of inflorescence. D. Detail of 
flower. E. Detail of stamen, showing adaxial surface. F. Detail of stamen, showing lateral margin. G. Detail of stamen, 
showing abaxial surface. H. Fruit. A-B drawn from isotype, C. Purpus 7032 (BM). C-G drawn from isotype, C. Purpus 7032 

1 .3-1 .9 cm wide, apically acute, early caducous; secondary t 
duncles, but 0.9-2.2 cm !ong;floral bracts persistent, membr, 
0.7-1.8 mm long, 0.3-0.7 mm wide, apically acute, the midr 
ondary veins obscure.densely and prominently punctate and punctate-lineate.glabrous 
above,sparsely erect glandular papillate below,the margins entire, hyaline,sparselyg!an- 
dular-ciliate; pedicels slender, terete, 6.7-1 0.3 mm long, inconspicuously punctate and 
punctate-lineate, densely, erect glandular papillate. F/otvers 5-merous, membranaceous, 
pink to reddish-violet; calyx 1.6-1.8 mm long, the tube 0.3-0.5 mm long,the lobes ovate 
to lanceolate, 1.1-1.5 mm long, 0.6-0.8 mm wide near the base, asymmetric, apically 
acute to rounded, prominently punctate and punctate-lineate, glabrous within, tomen- 
tum of scattered, erect glandular papillae, the margins entire, minutely erase, hyaline, 
sparsely glandular-ciliolate; corolla 5-5.3 mm long, the tube 0.9-1.1 mm long, the lobes 
connate basally, ovate to lanceolate, 3.9-4.4 mm long, 2.5-2.8 mm wide near the base, 
apically acute to rounded, prominently punctate and punctate-lineate,glabrous through- 
out,the margins entire;starmens 3.1-3.3 mm long;the filaments 1 .9-2.1 mm long,apically 
free, 0.3-1. 7 mm long, connate basally into an elobate tube, 0.4-0.6 mm long, free from 
the corolla tube, epunctate, glabrous, the anthers free, ovate, 1 .4-1 .7 mm long, 0.8-0.9 
mm wide near the base, apically apiculate, basally sagittate, dehiscent by wide, apically 
continuous longitudinal slits, the connective epunctate; pistil obturbinate, 4-4.2 mm long, 
glabrous; ovary 0.9-1 .1 mm long, the style 2.9-3.3 mm long, slender, erect, inconspicu- 
ously punctate;stigma punctiform;ovules 1 1 -1 5, pluriseriate. fru/'f globose, 5-6.4 mm in 
diam., conspicuously and prominently punctate, the style base persistent. 

Distribution— Ardisia rarescens is restricted to the extreme SE corner of Chiapas in 
the Sierra Madre Mountains,and in the adjacent area in the Departments of San Marcos 
and Quetzaltenango in Guatemala, growing at 1,300-2,400 m elevation. 

Ecology and conservation status— Ardisia rarescens occurs in montane rain forest and 
loud ton i l - i i' ,_,] i ' ii i I iii nhuiiuh n buiili \ < . \ I <> ,1 ihn it n< d 

Etymology— The specific epithet comes from the combination "rare" meaning far 
apart, sparsely and "-escens"to indicate a process of becoming without full attainment 
reached, referring to the sparse tomentum. 

Common Name.— "Cereza morada" (J.Steyermark 33656). 
Specimens examined. MEXICO. Chiapas: Mpio. Motozintla de Mendoza, 45-50 km NE of Huixtla 
along road to Motozintla, 1 ,900 m, 1 7 Nov 1 971 (fr), D. Breedlove & A. Smith 22592 (LL, MO); 28 Dec 
1972 (fr),D. Breedlove & R. Thorn 110 til \U mr if i ( ' i I, Jh Mendoza, 5W side of 

Cerro Mozotal, 1 1 km NW of the junction of the road to Motozintla along the road to El Porvenir 
andSiltepec,2,100m,21 Nov 1976 (fr), D. Breedlove 41608 (MO); 23 Nov 1981 (fr), D. Breedlove & B. 
Bartholomew 55740 (LL, NY); Municipio of Motozintla de Mendoza, between El Rosario and Ojo de 
Aqua along road to Nqn ill i I h l» ih it , ) Above El Rosario,8mi 

S of Motozintla, 1,800 rn, iO Jul 19// (II), / ( loat 40740 (LL, MO); Mt. Ovando, without elev., 24 Die 
P'-,o(P> / Uj/<;i/ ( /<)n-()(US) Mt Pi il in 1P1 ^uMi I'm il ' Matuda 1641 (LL);4Aug 

1937 (f\),E.MatudaS ( I II 1 t 1 n Nov 1939 (fl), E.Matuda 

3942 (A, NY); Mt. Ovando, Escuintla, without elev., 1-16 Jul I940(fl),i Matuda 4 180 (A, GH,LL, MO, 
NY);Saxchanal, Sierra Madre,2, 700 m, 1 Jul 1941 Ph ' [ I L MO NY) Carlas, near 

;Q45 ,; 

TEX) mi 

1 ,834 m,23 Jul 1 948 {f\),E.Matuda 18148 (F); Along the dirt road to Siltepec, past Ejido Benito Juarez, 
ca. 1 2 km from the turnoff from Mexican highway 1 90, S of Motozintla, 1 5° 20' N,92° 1 5' W, 2,1 00 m, 
9 May 1 987 (f\),J.Miller& J.Myers 2778 (BRIT, F, MEXU, MO);Tapachula, Finca Chinince, 1 ,500 m, 1 7 Aug 

Bogueron to Cerro Boqueron, 1 5°?1 5' N, 92° 1 7' W, 2,400 m, 9 Feb 1 990 (fr), P. Stafford etal. 347 (BM, 
MO); Municipio of Motozintla, Buenos Aires, 1 ,900 m, 3 1 Jul 1 986 (fl), £ Ventura y E. Lopez 3985 (MO). 
GUATEMALA. Quezaltenango: Volcan Zunil, 6,1 00 ft [1 ,859 m], 5 Aug 1 934 (fl), A Skutch 948 (F, NY); 
Lower S-facing slopes of Volcan Santa Maria, between Santa Marfa de Jesus and Calahuache, along 
great barranco between Finca Pirineos and San Juan Patzulin, 1,300-1,500 m, 6 Jan 1940 (fr), J. 
Steyermark 33656 (F, LL).San Marcos: 6 mi SW of town of Tajumulco, NW slope of Volcan Tajumulco, 
along Rio Malacate.2,300-2,800 m,26 Feb 1 940 (ster.) J. Steyermark 36666 (F);Above Finca El Porvenir, 
between "Todos Santos Chiquitos"and"Loma de la Paloma:, S-facing slopes of Volcan Tajumulco, 
1,400-1,700 m, 8 Mar 1940 (ster.), J. Steyermark 37283 (F, LL); 1,300-1,500 m, 16 Mar 1940 (ster.), J. 
Steyermark 3798 1 (F, LL); Near Aldea Fraternidad, between San Rafael Pie de la Cuesta and Palo Gordo, 
W-facing slope of the Sierra Madre Mountains, 1,800-2,400 m, 10-18 Dec 1963 (fr), L.Williams etal. 
26002 (F); (fl, fr), 26004 (F 2-sheets); (fr), 26 7 J (NY); Outer slopes of Tajumulco Volcano, Sierra Madre 
Mountains about8-10 km Wof San Marcos,ca.2,300m,31 Decl964-1 Jan1965(fi U - 

We thank the Missouri Botanical Garden and the Flora Mesoamericana Project, for fund- 
ing that allowed J. Ricketson (MO) to travel with J. Pipoly (FTG) on visits to the C.L. Lundell 
Herbarium (LL), housed at the University of Texas at Austin. Without access to that critical 
collection, assembled by C.L. Lundell over a period of nearly 60 years, the present study 
would not have been possible. We also thank the curators of the herbaria cited for loans 
of specimens. We are grateful to the staff of LL and TEX, especially Billie L.Turner, Tom 
Wendt,CarolTodzia, Beryl Simpson, and Jose Panero,for their cooperation and hospital- 
ity. We are also grateful to those who have been so instrumental in assisting us in our 
work, including Gerrit and Jeany Davidse, Linda Oestry, Mary Bard, and Catherine Mayo 
(MO), Barney Lipscomb, Debra Trock and Jim Rivers (BRIT). Illustrations were prepared by 
the junior author. Reviews of the manuscript by Gerrit Davidse and Roy Gereau and 
meticulous copy editing by Barney Lipscomb, improved the presentation of the paper. 


Chen, J. and J.J. Pipoly. 1 996. Myrsinaceae. In: 
Science Press, Beijing, People's Republic 
Louis, MO, U.S.A. 15:1 -38, lllus. 1-20. 

Conran, J. 1 995. Family distributions in the 
tions. J. Biogeog. 22:1 023-1 034. 

Gentry,A. 1 983. Alstonia (Apocynaceae): Am 

Hickey, L. 1984. A revised classification of tr 
25-39. In: C. R. Metcalfe and LChalk (edil 

Mogeographical implic 

r paleotropical genus in Central A 

Oxford, U.K. 

Lavin, M.and M. Luckow. 1993. Origins and relationships of tropical North America in the 
context of the Boreotropics Hypothesis. Amer. J. Bot. 80:1 -1 4. 

Lindlcy, J. 1 848. Illustrated dictionary of botanical terms. Excerpt from illustrated dictio- 
nary of botanical terms by John Lindley. Pp. 346-383. [Reprint, with an introduction by 
Alice Eastwood, SUmiord University, School of Earth Sciences 1964], 

Metcalfe, C.R.I 984. Some basic types of cells and tissues. Pp. 54-62. ln:C.R. Metcalfe and L 
Chalk, eds. Anatomy of the Dicotyledons .Vol. 1 .Systematic anatomy of leaf and stem, 
with a brief history of the subject. Clarendon Press. Oxford, U.K. 

Me/, C. 1 902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich IV. 236(Heft 9):1 -437. 

Pipoly, J.J. 1987. A systematic revision of the genus Cybianthus subgenus Grammadenia 

Pipoly, J J, 1 992. The genus Cybianthus subgenus Conomorpha (Myrsinaceae) in Guayana. 
Ann. Missouri Bot. Gard. 79:908-957. 

Pipoly, JJ. and J. Ricketson. 1999. Discovery of the Indo-Malesian genus Hymenandra 
(Myrsinaceae) in the Neotropics, and its Boreotropical implications. Sida 1 8:701-746. 

Theobald, W.L, J. KRAHuuK,and R.C.Rollins. 1 984,Trichome description and classification. Pp. 
40-53. In: C.R.Metcalfe and L Chalk (editors). Anatomy of the Dicotyledons. Vol. 1. Sys- 
tematic anatomy of leaf and stem, with a bnel Iiim m J lb' hioct. Clarendon Press. 
Oxford, U.K. 

Tffney, B. 1 985a. Perspectives on the origin of the floristic similarity between eastern Asia 
and eastern North America. J. Arnold Arbor. 66:73-94. 

Ti f nly, B. 1 985b. The Eocene North Atlantic land bridge: its importance in Tertiary and 
modern phytogeography of the northern hemisphere. J. Arnold Arbor. 66:243-273. 

WendtJ. 1 988. Chiangiodendron (Flacourtiaceae:Pangieae),a new genus from southeast- 
ern Mexico representing a new tribe for the New World flora. Syst. Bot. 1 3:435-441 . 

Wendj.T. 1989. Las selvas de Uxpanapa.Veracruz-Oaxaca, Mexico: evidencia de refugios 

I 1 ' ' i t i> Jit lftinit es and origin i tl no| n flom of ll 

Mexican Atlantic slope rain forests. Pp. 595-680. ln:TRamawoorthy,R. Bye, A. Lot and J. 
Fa, eds. Biological diversity of Mexico: Origins and Diversity. Oxford University Press., 

WolieJ. 1975. Some aspects of plant geography of the northern hemisphere during the 

late Cretaceous and Tertiary. Ann. Missouri Bot. Gard. 74:264-279. 
Zona, S. 1 990. A monograph of Sabal (Arecaceae: Coryphoideae). Aliso 1 2:583-666. 


Matuda, E. S-206; 0680; 1 641 ; 3942; 41 80; 4306; 55 1 1 ; 1 81 48. Miller, J. & M 

Purpus,C. 7032 (type) 

Skutch, A. 948. Stafford, P. et al. 347. Steyermark, J. 33656; 36666; 37283; 3 

Ventura, E.y Lopez, E. 3985. 

Williams, L. et al. 26002; 26004; 261 01 ; 26799A. 


Virginia Scott Jenkins. 2000. Bananas. An American History. (ISBN 1-56098-966-1, pbk.). 

Smithsonian Institute Press,470 L'Enfant Plaza Suite 71 00, Washington, DC 20560- 

0950, U.S.A. $ 1 6.95 pbk., 232 pp., 36 b&w photos. 
From the author of I 1 ' - 1 t n t a f of it it \inet 

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Ik titer, ted Kcvii •< Solan < < at ' limit >/ Texas, 



Theodore M.Barkley 

Botanical Research Inst, n,u , 
509 Pecan Street 
Fort Worth, TX 76 1 02-4060, USA. 

b irkle @bril 

iecio quaylei, a new species from Texas, U.S.A., is described. 

describe Senecio quaylei, una especie nueva de Texas, U.S.A. 

orth central Texas, recently submitted an 
3S presumed that the plant was simply a 
recent or otherwise unrecorded introduction, but a survey of the literature and several 
herbaria, plus the opinions of several colleagues,suggested no possible matches, so it is 
offered here as a new species, Senecio quaylei. 

The only known occurrence of the plant is a conspicuous colony of about 1 5 indi- 
viduals, growing waist high in a weedy roadside ditch, along with Cirsium, Geranium, 
Sonchus, Verbena, and much dead vegetation from the previous year. The new species 
vaguely resembles Senecio ampullaceus Hook., a Texas endemic of disturbed open sites 
in the central part of the state, however, S. quaylei is glabrous and notably coarse, to 12 
dm tall, while S. ampullaceus is conspicuously hairy but unevenly glabrate in age and 
typically only 3-7 dm tall. In addition, the new species has large, broad, and clasping 
cauline leaves.These characteristics combine to give the new species a distinctive gross 
aspect. Four of the plants were collected to make the specimens needed for documen- 
tation, and they were divided into eleven herbarium sheets. 

Structurally, the stems are hefty, with the proximal third some 10-15 mm in diam- 
eter, but they are hollow and thin-walled.There is a conspicuous purplish-red layer in the 
sub-epidermal region of freshly cut proximal stems;the red color fades as the specimen 
dries. The leaves have unevenly scattered light-brownish spots that are caused by a spe- 
cies of Coleosporium, a heteroecious rust,f/c/eDr.JoeHennen,a mycologist at BRIT and an 
authority on rust fungi. 

The biology of S. quaylei is unknown and only suggested from inference. It appears 
to be an annual, with a short, narrow taproot and a tuft of abundant, thin, fibrous roots 
that are weakly branching. The most mature ovaries in the specimens collected are 
wrinkled and unfilled, indicating that they are sterile. The pollen is of uneven size and 

shape, suggesting it is of reduced or doubtful viability. I he ( hiomosome number is 

The resembla nee of Senecio quaylei to Senecioampullaceus and other typical sene- 
cios (Barkley 1999) in both structure and aspect clearly places the new species in Sene- 
cio s. str.; rigorous quantitative analyses are yet to be done. It is possible that additiona 
data from cytology, pollen morphology, and comparative biochemisty,may suggest that 
the new species is allied to Packera, but that notion seems remote. Senecio quaylei is of 
poor fit in the keys to Senecio in the treatment in the North American Flora (Barkley 

n w the V,ws u> inlcurmmus assembiauoa uioup with unburn, eeu tleshv-fibrous toots 
and of very different gross aspect. If the initial couplet of the key is ignored, it falls into 
the Annui group, along with Senecioampullaceus. 

Recognition of Seneobguay/e/' generates speculation on its significance in the flora. 
The possibility that it is merely an exotic waif cannot be discounted, but neither can it be 
said that it is not a rare member of the regional flora. This notion is compatible with the 
review by Ertter (2000) on the occurrences and recognition of distinctive species of lim- 

Senecio quaylei T.M. Barkley, sp. nov. (Figs. 1 , 2). Type: U.S.A. TEXAS, Parker Co.: waste ground 

adjacenttol ike f 'in r II i n in i i i il m t It i n ilit t 

Chapel Road intersection, ca.2 mi N of Hwy 1 80, E of Mineral Wells,32° 45' N,98 o 02'W,29 Apr 

ii m ' i i ii I I II i I II Mi ih 

Annual, 8-12 dm tall, okibtous thiouohout or with a few in uespicuous hairs on the 
peduncles and phyllaries.Sfem5single,striate,the proximal third 10-15 mm in diameter, 
narrower distally, hollow, with a conspicuous reddish subepidermal layer when freshly 
cut; arising from a short, thin taproot surrounded by abundant, thin and sparingly 
branched fibrous roots. Basal and proximal cauline leaves with blades ovate, mostly 12- 
24 cm long and 8-12 cm wide, margins wavy, with a few scattered minute denticles, 
midvein prominent and lateral veins less conspicuous in dried specimens, tapering or 
gently contracted to a distinct petiole, ca. 2/3 the length of the blade. Middle cauline 
leaves with blades nearly as large as the basal leaves, ovate to broadly lanceolate, sessile 
and clasping. Distal cauline leaves lanceolate to linear-lanceolate, 6-14 cm long, sessile, 
the distal most further reduced and bractlike. Capitulescences terminal or arising from 
the axils of the upper !eaves;fundamentally corymbiform cymes of 20-40 capitula,or a 
close cluster of corymbiform cymules. Involucres cylindrical ot u.sbinate to weakly cam- 
panulate, each subtended by a weakly defined calyculus of 2-7 linear bracteoles, 1-3 
mm long, margins hairy. Phyllaries mostly 13, ± 8 mm long, green with hyaline margins, 

Corollas yellow. Ray florets mostly 8, pistillate; corollas ca. 1 1 mm long, tubes 4 mm and 

;. l.Senecio quaylei. Habitat from type locality. Photograph by Bob O'Kenr 

laminaeca.7mmlong,2.5mm wide.D/s/c/7orefs20-40,bisexual;corollas 7(-10) mm long, 
tube and limb (including lobes) of about equal length, corolla lobes triangular,! 0.5 mm 
long. Cypselae not seen, oldest ovaries wrinkled, empty, 1 -3 mm long, pubescent through- 
out. Pappus of abundant white, minutely barbellate bristles in a single series, 6-7 mm 
long. Chromosome number unknown. 

Etymology.— The specific epithet quaylei commemorates Jeffrey Quayle, the dis- 
coverer of the new species. 

Distribution. — Endemic to Texas. There are no known collections other than the type 

This key covers the range of the recently published Illustrated Flora of North Central 
Texas (Diggsetal. 1999) 
1. Capitula discoid, rj\ rl m < ,H ; . t . -,< Til and then with laminae of corollas less 

It is a pleasure to acknowledge the assistance of Joe Hennen, Barney Lipscomb, ( 
Nesom, Robert O'Kennon, John Pruski, Peter Raven, John Strother, Debra Trock,and c 
ers who offered advice on the nature of this plant. Guy Nesom kindly translated the ii 
diagnosis and Linny Heagy prepared the illustration. 

1 999.The segregates of Senecio,s 

Mexico. Sida 18:661-672. 

. 1978. Senecio. In: C.T. Rogersor 

Diggs, G.M., B.L Lipscomb, and R.J. 
north central Texas. Sida, BotT 
of Texas, Fort Worth. 

Eh ■•:■■, B. 2000. Floristic surprises 


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i i ii i in i ih in u i i i It in i 

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pages of further reading and an index follow these chapters. Throughout the book Sumner covers 
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mj I t it i i e ] ii ii, ili in I mi rh t i I i i\ rnthropology and 

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i, Botanical Researc 


Solidago villosicarpa, sp. nov., is described from the outer coastal p 
lina, where collections have been made from four counties. Although it apparently is ex 
two of these four c nintie it h ih if ni 'i J i n 

species may be more frequent cii i iT- lm 1 n uirentlv known. Solidago 
is placed as a member of sect, and subsect. Solidago and is dr tir i ti 'inn m n i 
cent stems, glabrous to glabrate leaves, thyrsoid inflorescence, large heads with brighl 
low rays,densely villous achenes,and late flowering. 

Se decr'ibeSolidago villosicarpa, sp. nov., de la llanura costera externa del sudeste de C habitat y evidenteadaptacion a la perturl i rn 
que la nueva especie puede ser mas frecuente a lo largo de la costa de Caro ina T 
conoce actualmente. Solidago villosicarpa se coloca como un miembro de la sect, y si 

Early collections of a distinctive goldenrod species were made from"live-oakscrub"on a 
barrier island in Brunswick County, North Carolina, in 1 949 and 1 950. Specimens from 
the 1 950 collection were annotated as S. sciaphila Steele (Fox et al. 1 952), a plant other- 
wise known only from sandstone and calcareous habitats in the Upper Mississippi River 
region (Minnesota, Wisconsin, Iowa, and Illinois). A subsequent collection of the North 
Carolina entity was made in 1963 from a sandy roadside in neighboring New Hanover 
County, N.C.,and annotated asS.erecta Pursh (NCU).Three more populations were found 
1991-1998 in Onslow County, N.C., during a natural area inventory of Camp Lejeune 
MarineCorps Base by the North Carolina Natural Heritage Program (NCNHP).and a popu- 
lation was found in Pender County in 1 998 during another NCNHP inventory. Only the 
Onslow and Pender county populations are known to be extant. 

The original collections of the North Carolina entity from 1 949 and 1 950 have a 
curious history. According to Fox etal. (1952) the 1950 collection {Godfrey 50963 &Boyce) 
was identified as Solidago sciaphila"by Dr.ArthurCronquistand it was rechecked by him 

SIDA 19(2): 291 -300.2000 

after several duplicate specimens were sent lo him. Godfrey als< ) c decked specimens of 
the collection against material of the species at the Gray Herbarium, and he concurs 
with Dr. Cronquist in his determination. We are at a complete loss to account for the 

The 1 949 collection (Godfrey 501 32) is not mentioned in the 1 952 Rhodora article. A speci- 
men from this collection was found in the folder for undetermined Solidago specimens 
at the N. C.State University herbarium (NCSC) during the current investigation. The label 
of this specimen (herbarium #1 9998) has a typed and penciled portion. The typed por- 
tion reads "Solidago ...In live oak scrub on the sand dunes, Long Beach.'The penciled 
portion reads "sp. aff. 5. sciaphila Steele, less closely aff. S. glutinosa sens. lat. May need a 
name.A.C.3-22-50.""A.C"undoubtedly is Arthur Cronquist.These circumstances suggest 
that the 1 949 collection had been forgotten by the time the 1 950 collection was deter- 
mined. Adding to the mystery, the three specimens from the 1 950 collection at NCSC 
(herbarium #'s 26549, 26550, 34039) were annotated as 5. sciaphila by H.E. Ahles, not 
Cronquist. Yet them i im mention ot . ,/ ... miing in the Carolinas in Radford et 
al. (1 968) (Ahles was responsible for the treatment of Asteraceae), nor apparently in any 
other flora, treatment, oi checklist since thai time. No specimens from the 1949 and 
1950 collections are known from other herbaria, including the New York Botanical Gar- 

This distinc tive,iaie,and narrowly endemic North Carolina plant does indeed "need 
a name,"and is here described as a new species. 

I ; , Tiqs.1 6 

nate; stem color medium brown to datk bio i i I . ui| I lov and light brown, 
stramineous, or purple above. Basal leaf rosette present; basal and lower cauline leaves 
the largest, petiolate,toothed;larger blades 7-14 cm long (not including petiole) by 4-7 
cm wide, elliptic, bn » . I | • tlhf ii. obovate, apices obtuse, subacute, or broadly 

acute; bases cuneate-attenuate, often somewhat abruptly narrowed; petioles 2-7 cm 
long; petiole base non-auriculate but sheathing the stem for 1/4- 1/2 of its circumfer- 
ence; blade margins toothed, the sinuses 1 -2.5 mm long from base of cavity to tip of 
tooth mucro,the mucros mostly 0.2-0.4 mm long,blunt;teeth margins ciliate-scabrous 

' ..iJiiki ;u viilosicarpa LeBlond. Habitat showing example of paniculately branched thyrsi 




ML - * '1 

Figs. l-6.Solidago villosicarpa LeBlond. Fig. 3 (top left) simple 
elongate thyrse inflorescence form. Fig. 4 (top right) example 
of the paniculately branched thyrse. Fig. 5 (bottom left) close- 
up of the inflorescence. Fig. 6 (bottom right) a well-developed 

i surfaces, the mid-nerve flattish and squarred at the edges, pale 
ate nerves distinctly reticulate on both surfaces; adaxial surface gla- 
brous to sparsely pubescent with short stiff trichomes mostly along the mid-nerve and 
larger veins; glabrous to glabrate abaxially; texture thin, papery, brittle when dried; color 
drab green to olive-green, the lower surface barely if at all lighter than the upper. Middle 
and upper cauline leaves gradually reduced upwards, sessile, becoming entire; cauline 
leaves 1 5-50 below inflorescence. Inflorescence a simple or paniculately branched thyrse; 
when simple (elongate terminal thyrse), the terminal axis bracteate,straight, narrow,and 
cylindric, 7-22 cm long by 3-6 cm wide; when paniculiform, producing straight, elon- 
gate, thyrsoid axillary branches up to 20 cm long; bracteal leaves at the base of the 
branches similar to the cauline leaves,gradually reduced in size upwards. Short second- 
ary branches and peduncles moderately densely to densely invested with stiff, spread- 
ing and ascending, straight and recurved trichomes 0.1-0.4 mm long, which appear res- 
inous and segmented; peduncles 0.5-9 mm long. Heads at anthesis 1.4-1.7 cm wide 
NRMMnvil from ray tip to ray tip. Involucre 5-8 mm long by 3-5 mm wide at anthesis, the 
summit 6-8 mm wide at maturity. Phyllaries strongly imbncate,the outer shorter,c 
somewhat cucullate,the inner broadly linear;outer phyllaries 1 .0 -2.0 mm wide,appressed; 
inner phyllaries 0.8-1.5 mm wide, with rounded to subacute apices becoming s 
I H gu him ( in ,i ]l ill -ill, il nit- anh ,) n now bin d in i \ .> i bdi I, bio n 
colored centrally and laterally, the margins hyaline, often lacerate, long-cilia 
ciliate-fimbriate (at least near the apex) with cilia 0.1-0.3 mm long. Ray florets 4- 
head, limb in vivo 5-7.5 mm long, 1 -2 mm wide, bright lemon-yellow. Disk florets 1 0-18 
per head, the corolla lobes 1 .5-2.2 mm long, the entire disk corolla 4.9-6.8 mm long; 
stigmatic lobes 0.9-1.1 mm long, anthers 2.1-2.2 mm long. Pappus (4.2-)4.7-6.1 mm 
long, occasionally some bristles clavate. Achenes villous with ascending hairs 0.3-0.5(- 
0.7) mm long, the achene body 2.6-2.9 mm long when mature. 

Additional specimens examined:NORTH CAROLINA. Brunswick Co.: in live oak scrub on the sand 
dunes.Long Beach, 22 Oct 1 949, R.K. Godfrey 501 32 (NCSC); in live-oak scrub thickets on sand dunes, 
IV -. Hanover Co.: sandy road- 
side, Pembroke Jones Park, Wrightsville Sound, 29 Nov 1 963, A. McCrary 1813 (NCU). Onslow Co.: in 
pine-oak forest with open understory, Camp Lejeune Marine Corps Base near mouth of Frenchs 
Creek, 22 Oct 1 991 , R.J. LeBlond 2622 (pers. herb.);same locality, 1 8 Oct 1 992, R.J.LeBlond 3127 (NCU); 
in pine forest with open understory, Camp Lejeune Marine Corps Base near Salliers Bay, 08 Oct 
1995, R.J.LeBlond 41 10 (\ r holo inn k>< ilit\ I I n, i m / 1 AS Weakley, and K. 

Patterson (RJL per ful i in h ihtv If- Dd 1998, RJ. LeBlond 5082 (DUKE, NCSC, NCU); Camp 
Lejeune Marine Corps Base, SW of Mock up Road, 0.9 mi SE of NC 1 72, 05 Nov 1 998, RJ. LeBlond 5 124 
and E.Davis (NCU). Pender Co.: in pine hardwood foiesi mi WNWof i larks Landing on Long 
Creek, 30 Sep 1G98 ' , > m uw/e (NCU). 

mm long. The size combined with the bright lemon-yellow color of the ray limbs make 
this one of the showier goldenrods.The density of the achene pilosity completely ob- 
scures the body surface with hairs 0.3-0.5(-0.7) mm long.The combination of thyrsoid 
inflorescence, persistent and glabrous to glabrate basal leaves, upwardly reduced cauline 
leaves, and pubescent achenes place the new species in Solidogo sect.Solidago subsect. 
Solidago in Nesom's 1 993 overview of infrageneric goldenrod taxonomy. North Ameri- 
can members of subsect. Solidago are (Fernald) Fernald,S.g/o/nerafaMichx.,5. 
multiradiata Ait. (including =5.cur/e/-/Fernald) ,S.nana A.GrayS.p/umoso Small, S.sciaphila, 
S. simplex Kunth, S.spathulata DC.,and S.spithamaea M. A. Curtis. Solidago villosicarpa dif- 
fers from other members of the subsection by a combination of stem pubescence, floral 
head width at anthesis, involucre length (5-8 mm), pappus length (4.2-6.1 mm), length 
of disk corolla and lobes (4.9-6.8 mm), and nature of achene pubescence (villous, the 
hairs 0.3-0.7 mm long). Also, it flowers late September to early November, one to two 
months later than the others. It superficially resembles S.bicolor L.,S.hispida Muhl.,and 5. 
squarrosa MuhL, members of sect. Solidago subsect. Albigula in Nesom's is 
readily distinguished from all three by its pubescent achenes, from S.bicolor and S.hispida 
by its glabrous (-glabrate) leaves, and from S.squarrosa by its appressed outer phyllanes. 
The following key distinguishes southeastern U.S. Solidago taxa with thyrsoid 
inflorescences, basally disposed leaves conspicuously larger than middle and upper 
cauline leaves, and pubescent stems. 

1 . Inner phyllaries very narrow, <0.5-0.75 mm wide at mid-length, tapering to slender 

2. Stems irregularly or decurrently short-hairy (-glabrous) below t 

3. Leaves glabrous (-sparsely pubescent adaxially);involucre 5-8 mm long; ray florets 

4-8; pappus (4.2-)4. 7-6.1 mm long; achenes villous S. villosicarpa 

3. Leaves pubescent on both surfaces; involucre 3-6 mm long; ray florets 7-14; pa- 
ppus 2.5-4 mm long; achenes strigose-puberulent or glabrous 
4. Achenes strigose-puberulent; disk flowers 14-27; larger leaves 3.5-1 2 cm long; 

plants flowering in spring S.verna 

4. Achenes glabrous (sometimes sparsely hairy when immature);diskflowers 7- 
1 6; larger leaves 8-20 cm long; plants flowering late summer-fall 
5. Ray limbs white (rarely yellow), often turning yellowish in drying; phyllaries 

ly with a well-defined green tip S.bicolor 

5. Ray limbs deep yellowtoorange-yellow;phyllaries w IL '■ ih- ; 

or not at all greenish S. hispida var. hispida 

.Ithough similar to Solidago sciaphila in overall habit, 5. villosicarpa is distinguished by 
?veral characters, particularly within the inflorescence (Table l).The stem of S.villosicarpa 
, pubescent throughout, while that of S. sciaphila is normally glabrous below the 

Disk corolla length (limb ; 
Disk corolla lobe length 
Pappus length 

inflorescence (Fein il i I i ,l , i nun, n.t |ui t I 

occasionally can have sparsely to moderately pubescent stems (frequently so just below 
the inflorescence).The basal and lower leaves of S. villosicarpa tend to be sparsely pubes- 
cent to glabrous adaxially and glabrous abaxially, while those of 5. sciaphila tend to be 
either glabrous on both surfaces or, less frequently, sparsely pubescent on both surfaces. 

With so few collections and kirn ip tu'tmi tl i I ^ dagovillosicarpa 

is only partially undm i id I 049 md I QSO Brunswick County collections are from 
"live-oak scrub"or"live-oak scrub thickets"on the Long Beach coastal barrier island. This 
likely is either Maritime Evergreen Forest or Maritime shrub m< h.iiale and Weakley 1990). 
Maritime Evergreen Forest typically has a canopy dominate, i bv ( kicrcus virginiana, usu- 
ally with Plnus taeda and Q. hemisphaerica. Charactmistic umlmsiory species include 
Juniperus virginiana i 1 1 1 ins andOsmanthus 

americana. Wind-borne sand and salt spray often produce dense thickets along the 
ocean-facing side of sm ii loiesh.ln areas < losei to mm ocean or more exposed, the Mari- 
time Shrub community forms. It is characterized by a dense growth of such shrubs as 
Ci'tothamnusceiifern {-My in m omem var.c critcia)d'e\ vomitoiinj-lnci has is halimitolia.and 
nil]'- m ■ 1 /and m 

The three Solidago villosicarpa sites in Camp Lejeune Marine Corps Base in Onslow 
County have been altered by past logging. Canopies at all thiee sites are dominated by 
pine {Plnus taeda), with hickory {Carya glabra var.megacarpa or Calba) and/or oak (Quercus 
falcata, Q. nigra, and 0. stellata the most common). Frequent understory species are Q. 
margarettiae,Liguidanithi! ivra< ithia, and >•■ m 1, o I Ins composition suggests the Dry 
or Dry-Mesic Oak-Hickory Forest natural community of Schafale and Weakley (1990), 
perhaps transitional to Coastal riinoe i mk|uvn I on ,1 i,e,,, ,t the ,ite near Salliers Bay, 
where Q. virginiana is a subcanopy component. Two of the sites occur on excessively 
drained Wando fine sand entisol, and the thuo 1 n 1 II inined Marvyn and 

N>)i tul', fximy fine sand ultisol. 

The site in Pender County near 1. larks I anding occurs on slopes above a drain; it, Got d ' n 1 || 1 , > ilustris. Understory 

and shrub species include Ilex opaco, Lyonia lucida, and Vaccinium arboreum. This mix 
suggests a former longleaf pine community grading downslope to a pocosinstreamhead, 
and likely influenced by proximity to tidal freshwater swamp 300 feet downstream (5. 
villosicarpa itself appears to be an indication of that influence).The upland soil at this site 
is classified as well-drained Baymeade fine sand ultisol. 

Two of the four currently known sites— in Camp Lejeune near Salliers Bay and near 
Mock-up Road— are within one mile of the ocean, and each was impacted by the hurri- 
canes of 1 996 and 1 998, with considerable canopy blow-down. Increased seedling es- 

responsetothe hurricane impacts. At the Salliers Bay site, 300-400 flowering individuals 
and 1 000+ vegetative rosettes were estimated in 1 998,compared with 50 flowering and 
100 vegetative rosettes under a closed canopy in 1995. The Mock-up Road site, first dis- 
covered in 1 998, had 500+ flowering/fruiting individuals.The other two sites are consid- 
erably inland, with the Frenchs Creek site in Camp Lejeune seven air miles from the coast 
and the Clarks Landing site in Pender County 1 6 air miles from the coast. Each site, how- 
ever, is located on low uplands or upland slopes adjacent to fresh or slightly brackish 
tidal creeks or floodplains. Due to their more inland locations,these sites were much less 
impacted by the 1 996 and 1 998 hurricanes. Comparative population data are available 
only for the Frenchs Creek site: 25 flowering with 75 vegetative rosettes in 1998, and 40 
flowering with 1 50-200 vegetative rosettes in 1 992. Another observed habit of Solidago 
villostcarpa that may be associated with hurricane impacts is the tendency of plants in 
canopy openings to produce more robust inflorescences with paniculate thyrsoid 
branches, while plants in shaded areas tend to produce a simple elongate terminal thyrse. 
These variable conditions and evident adaptation to disturbance suggest that Sol- 
idago villosicarpa, wft\\e likely restricted in range, may be more frequent along the Caro- 
lina coast than is currently known. 

I am indebted to Alan Weakley for his recognition of the resemblance of the Camp Lejeune 

this paper in draft. Material assistance was provided in the field by John Hammond a 
Davis of Camp Lejeune Marine Corps Base Division of Fish and Wildlife.and by Bruce 
Karen Patterson, and Alan Weakley. Fieldworkthat resulted in the discovery of the Onslow 
and Pender county occurrences was conducted under the auspices of the N.C r 
Heritage Program. Jeff Nekola and Bruce Sorrie provided critical help in the exami 
of habitat and character states of Solidago sciaphila populations in northeastern 
am grateful for the assistance provided by the herbarium curatorial staffs at the University 
of North Carolina-Chapel Hill (NCU) and N.C. State University (NCSC).The illust 
were prepared by Margret Mueller and photographs were taken by Bruce Sorrh 
using living specimens from the type locality on the date of the type collection. C 
U.S. Geological Survey topographic quadrangle place names are used when ava 

<> < ' 'f J' 'i i, A l l »80 Vascular floia of thi- southeastern U.S. Vol. 1 : Asteraceae. Ur 

North Carolina Press,Chapel Hill, NC. 
Fi rnai 1 1, M.I . 1 950. Gray's manual of botany. 8th ed. D. Van Nostrand, New York, 
] W.h, PI d in jnd ML Bi i u 1°5J Note- dim! ml ul m -III ml 

plants— III. Rhodora 54:165-182. 
C.i i ason,H.A. 1 952.The new Britton and Brown illustrated flora of the northeast* 

States and adjacent Canada. 3 vols. New York Botanical Garden. 
t.>n j.H.A.andA.C , n^i. I 991 .Manual of the vase ular plants of northeaste 

States and adjacent Canada. New York Botanical Garden, New York, NY. 
Kallunki, J. 1 998. Personal communication (e-mail) to R.J. LeBlond, August 1 3. 

Botanical Garden, New York, NY. 
Nesom, G.L. 1993.Taxonomic infrastructure of Solidogo and Oligoneuro 

1 ' im i ml I hi i ii h nli I mi t | mi 'h tologia 75:1-44. 

University of North Carolina Press,Chapel Hill. 
Schafale, M.P. and A.S. Wfaki ey. 1 990. Classification of the natural communities of North 
Carolina (Third Approximation). N.C Natural Heritage Program, DPR, DENR, Raleigh. 




Jose Martin Vazquez Lopez 1 , Bruce F. Benz 1 ' 2 , 
Miguel Olvera Vargas 1 , Sergio Graf Montero 3 

'Universidadde Guadalajara, Centro Universitario de la Costa Sur 
DepanamentodeEcologiayRi'i,,' > ,., 
Apdo. Postal No. W8,Autlan de Navarro 

Secretaria del Medio Ambiente Recursos Naturales y Pesca 
Autlan, Jalisco, 48900 MEXICO 


i nil of ihi n mini mil ofomte (Olaten aanwiiabi 

lanagement.The poin 

n category, harvesting, art 

to cuadrante para estimar la densidad de tallos. 

de varianza de la densidad y categorfas de tallos para comparar los rodales 

cativasen unaovarias 

Numerous genera and species of the Bambusoideae play an important role in forests 
because they occupy a wide diversity of habitats and exhibit extremes of morphologi- 
cal diversity (Soderstrom & Calderon 1 974). The genera of bamboos in Mexico include: 
Ototea, Olmeca, Guadua, Chusquea and Olyra (Judziewicz et al. 1999). The antiquity of 
Bambusoideae species'use in Mexico is not known with certainty though we suspect its 
most ancient inhabitants used them. The bamboo otate {Ototea spp.) has been used in 
Mexico since prehispanic times for a wide variety of purposes including house con- 
struction, walking sticks,stakes,and for basket making.amonq others ("lorres 1985;Anaya 
1 989; Benz et al. 1 994; Bye 1 995). Despite its long history of use and considerable study of 
its distribution and taxonomy, it is clear that information about the species' ecology and 
management is sorely lacking. 

Ototea is a genus indigenous to Mexico and Central America, occurring in Pacific 
watersheds from Sonora to Chiapas and Central America, besides the Mexican states of 
Veracruz, Puebla, Queretaro and Mexico (Guzman et al. 1 984).There is a disjunct popula- 
tion in northeastern Colombia where it is called caha brava (Judziewicz et al. 1 999).There 
are two species in the genus, 0. fimbriata and 0. acuminata, the last one with two sub- 
species, 0. acuminata subsp au i • ml ' -- ita subsp. aztecorum. These spe- 
cies are used foi bad « t n I in . ill i i hinom uO on i nstiuction, canes, fur- 
niture, crop supports, and house rafters (Guzman et al. 1 984; Judziewicz et al. 1 999). 

This research was carried out in the ejido of Platanarillo, in the Municipality of 

Calderon &Soderstmn ul j ,v nmo iuzman,Anaya&Santana) isan important natu- 
i'al ' u - Ihi u pre h i hi in ih ' I <ic in i , i i i i ■iihu Tin in ilo i 

Durango, Nayarit, Jalisco, Queretaro, Mexico, Guerrero y Puebla (Guzman et al. 1 984). 

Otate forms dense thickets of erect, two to eight meter tall individual shoots whose 
apex often overarches surrounding vegetation.This species occurs mainly in gorges and 
on pronounced slopes, on thin mil t n I m >i i i I In illv derived from cal- 
careous rocks (Guzman et al. 1 984). Otate spreads asexuallv I w i hizomatous growth.Young 
individuals are totally covered with culm leaves. Stems (culms) emerge annually in the 
humidseasonandreachtheirmaximumh ght iihin hreetol ui nonths.ln Platanarillo, 
otate is used in house and corral construction but its principal utility and value is tied to 
its suitability for making handicrafts, principally baskets. There has been resurgence in 
loi I mi. i i ui ih. mol Di h i ilt- i ti « iii -ii In i hi oi 'i ni ill . I fini, mi ,1 , i , j, , 
of stakes for cultivation of tomatoes, chayotes and other vegetable crops. 

Craft production in the ejido Platanarillo is of great socioeconomic importance 
because 32 families, or about 40 percent of the residents, engage in basket manufacture 
as a principal means of generating household income.Otate harvesting for this purpose 
has been carried out over the years under a traditional management scheme, that con- 
sists of the selective cutting of young stems.This traditional form of management is of 

The obj< 

active of this \ 

/vork is to de 

■scribe how tr 

been affected by tradH 

:ional extract 



The Ejidi 


d is located b 


latitude and 103°56'ai 

id 1 04°00' W 

' longitude in 1 

passes 3 

,,028 hectares 

at an altitud* 

2 ranging frorr 

Nearly o 


ea is located or 

a calcar* 

?ous formatior 


tary origin wit 

■aphic coordinates 1 9°21 ' and 1 9°29' N 
the state of Colima.This ejido encom- 
n 900 to 1800 meters above sea level, 
n the southwest flanks of Cerro Grande, 
h 1 to 45 degree slopes. The remain- 
der of the ejido is located on the north slope of the Sierra Perote,a volcanic formation 
with 25 to 45 degree slopes. Both form part of the northern-most extent of the Sierra 
Madre del Sur. Soils on the slopes of Cerro Grande are litosols and andosols,and on the 
Sierra Perote, regosols and cambisols (INEGI-SPP 1 981 ). 

Lithosols are generally shallow, rocky and infertile soils located in areas with pro- 
nounced slope.These soils are not particularly apt for agricultural purposes. Cambisols 
are superficial soils rich in organic material with weakly developed horizons found on 
moderate slopes. Nevertheless, cambisols are more appropriate for agriculture than litho- 
sols. Andosols form from volcanic ash. They are very light textured and have a high ca- 
pacity for retaining water and nutrients.This type of soil scarcely occurs in the study area, 
generally in areas with high slopes. Regosols are similar to cambisols in being rich in 
organic matter and having weakly developed horizons found on moderate slopes.They 
differ by regosols having almost no horizon development. Regosols support agricultural 

The climate in the area surrounding the ejido is mild,mid-latitude humid subtropi- 
cal with dry winters and hot summers according to the modified Koeppen classification. 
Average annual precipitation is 1,350 mm. It presents a marked seasonality with a dry 
season from October to May and a wet season from June to September (Martinez et ai. 
1 991 ).The predominant vegetation types in the ejido are the tropical deciduous forest, 
tropical sub-deciduous forest,and deciduous oakforest (Vazquez et al. 1 995).Otate popu- 
lations are an important component of the tropical deciduous forest that occupies the 
greatest extension in the ejido. These populations of otate occur mainly in communal 
land in the ejido, but also in areas with assigned rights, where basket makers must get 

Sites dominated by otate stands were delimited with aerial photos and a satellite image 
SPOT1 HRV2; band 321 , scale 1 :50,000 taken on March 30th of 1 987 and ground truthing 
in the ejido. Otate populations occur over 340 ha in the ejido. An inventory of the otate 
populations was conducted during the months of May June 
document the presence and evaluate the quality of stands. 


t least a 

s.These populations cover about 1 00 
d remain today in spite of harvesting. 

tensity was calculated as the numbet ol stems per hectare 
r method of Cottam and Curtis (1 956). Eight stands subject to 
were sampled systematically. Fifteen to twenty points were 
nts were placed on Inn ai imm e. id tubuted systematically 

cording to observed spatial pattern of the stand. Estimates of den: 

Bity were calculated 

using the equation proposed by Cottam i 

ind Curtis (1 956): D=dnr 2 


density and dm is equal to the mean distance of stems from the center point in each 

quarter. Analysis of variance with multiple t 

)ost-hoc comparisons usir 

ig Duncan's method 

was used to compare population densitie 

Culm diameter at breast height (dbl 

i) was measured using a 

caliper. Height was 

measured using m n fl dstab , A, ,triioi< 

J ached their maximum height, they were clas- 

.ified ntot )ui grcn th stag* itegi u n< 

■w, young, adultand dead. 

New refers to stems 

that emerged during the last rainy season 

i and were easily recogni; 

:ed by having com- 

plete spiculaU' l u ' m '■ leaves; young tele-v V 

3 those stems that had en- 

lerged two to three 

years ago, losing some culm leaves in the 

intervening time; adult s 

terns refer to those 

with few or no culm leaves. Dead stems v\ 

'ere easily recognized sine 

:e they had neither 

foliage nor culm leaves and had brittle stems. Average age of shoot d 

eath was unknown. 

The average density of stumps (cut stems 

) was considered an indie 


an indicator of regen- 

Vigor was characterized 

nth' *' .or 

notably healthy, complete 

I foliage; regul 

h some d in > i' b h i -< t>'i 

s to gray and c 

/, i hi! ,ihlv damaged leaves. 


Population vigor varied from good to regular for new and young s' 

generally had regular vigor. Site characteristics were similar for stain 
slopes of Cerro Grande while tl ^ i i f I u i >n Sierra Perote > 
favorable (Table 1, stand 8). 

Stand eights hi! ted ih ti it. t r< ei rat n and the highest 

Accessibility 1 Area Aspect Slope in Rockir 

or assigned rights). 


2 Rockiness refers to abundance and 

tion and of regeneration (Table 2). Stands five and eight had the largest diameter of 
young, mature, and dead stems. Stands one, four and five had relatively large-diameter 
stems. Stands six and seven had small-diameter stems (Table 3). The tallest stems oc- 
curred in stands one and five; the shortest stems in stands two, three and seven (Table 

(Table 3 and 4). 

In summary, stands number six and eight presented significant differences from 
the rest with respect to density, degree of extraction, and stem diameter. Stand eight 
exhibited the highest level of extraction, the greatest density of regeneration, and the 
largest— diameter and height— stems compared to the rest of the stands. In contrast, 
stand six exhibited no evidence of extraction, had the largest density of dead stems, 
showed the lowest density of regeneration, and had some of the smallest diameter stems 
(Tables 2-4). 

According to our interviews with local basket makers, stands one and four were 
intensively extracted in the past. Stems are extracted from stands one and four by the 
greatest number of basket makt i I n i lulanty, each collecting between 

40-60 stems per fortnight during September through November.Stands two,three and 
eight are subject to an intermediate level of extraction, at most ca. 40-60 stems per 
month. Stands five,six,and seven currently have the lowest levels of harvesting,the eguiva- 

Young 1412±1302 2^11^2^63 3556±2599 291 1 ±2922 4810±2603 4871 ±321 

abc abed bede abede cde cde 

Adult 4025±6057 2089±2141 4550±6686 3081 ±3306 6480±3623 8227±6393 

Dead 1304±1065 4402±6397 5270±4653 3106±3463 7614±9690 9556±9355 

) ,Mu„ l ap,ocfM,na,i,lan, | 

2 3 

0.62 2.4±0. 



sare not signifl 

antly differe 




i the ejid 















S.b5±l -'" 


2.46±0.71 2.4 

2.82±0.65 3.47±0.95 2.67±0.90 2.01 ±0 


Harvesting in the ejido is based on selection of young stems because these make better, 
more flexible, baskets. Over the years, this kind of management has produced a stand 
structure with an abundance of adult stems and appears to promote regeneration. 

Field observations indicated that stands one and four have the greatest prolifera- 
tion of woody species (Acacia spp.and Lysiloma spp.) and these stands have some of the 
lowest densities of young, adult, and dead stems. According to local informants these 
two sites were once occupied by dense stands of otate that were intensely exploited in 
the past, suggesting that prolonged and intense extraction impeded regeneration of 
adult stems in these otate populations and permitted the invasion and establishment of 
trees and shrubs Thi inn ,> un . tiiHokm . >tm in i tout d mug the past was due 
mainly to their proximity to the communities inhabited by a large number of basket 

producing a high don it t n .-. la? i> di i?imtm dmu no mid be due eitherto high 
levels of extraction reducing competition or its location on rock-free acidic soils with 
high fertility. Stand six had the lowest level of regeneration— it had the highest density 
of mature and dead stems— perhaps because the population has not been subject to 
thinning by basket mnko < oi.u ting siems foi I >asktt mum ium< lure. These results lead 
us to hypothesize that the basket makers of Platanarillo have exceeded sustainable lev- 
others (two, three, five, seven and eight) appropriate to eoolouioi conditions and the 
populations ability to regenerate. 

Numerous hypotheses have been offered to explain massive flowering of bam- 
boos. This phenoniPN n h i i i n mi o up m iht be determined by physiologi- 
cal changes caused k > n . in i n In nns, attempts to satiate seed predators, the in- 
ta< tionot Ml Hit 1 1 ( lanzen 1976;Keeley 

& Bond 1999). The fl i . iiuipnm i >t these otate populations is known from local 
folklore (Santana & Lemus 1 992). According to local people, the last massive flowering 
event occurred 30 to 35 years ago.Seven informants between 40 and 55 years old report 
having seen the flowering process once before, and two info- -ma tits, 75 and 80 years old, 
have seen it flower twice before. 

In 1993 weobset it tin tl n i it t to t pkt n otth ji k 

but not all individuals in a stand apt ired t i et n ill 1 1 1 l\ Widely separated 
individual culms flowered while nearby culms did not. Within one year's time from the 
appearance of one flowering in I id il h indi ]n,| /.ill have flowered. Now, in the 
year 2000, most of the populations have flowered completely. Nevertheless,the flowering 
process has been seguentia! and prolonged, not simultaneous: some populations still 
have not flowered. By the time the last population in the ejido flowers, the first popula- 
tion to flower will have new shoots can bo oxtrac ted for basket manufacture. The 


our observatic 

Continuous harvesting of young stems from stands three, five, six, seven, and eight ap- 
pears to be possible from the information we have obtained thus far. These stands had 
either a high density of regeneration (stand three) or had high densities of young and 
adult stems (stands five,six,and eight). Stand seven had the lowest level of regeneration, 

It appears contradictory that population six located at the greatest distance from 
the largest number of basket makers possesses the poorest guality for basket manufac- 
ture, while population eight located closest to a significant number of basket makers 
exhibits the greatest intensity of extraction, hast the greatest amount of regeneration 
and possesses stems with greatest useful dimensions. We offer the working hypothesis 
that human extraction of otate stems in Platanarillo has actually fostered growth of more 
stems with better qualities. 

Otate is a very important resource for the ejido,especially for the poor people (most 

extraction poses management challenges because the intensity of extraction could be 
considerably greater than potential regeneration, while labor investment and the im- 
mediate economic benefits are significantly higher than actual costs. 
Suggestions For Management 

A controlled management program must be implemented in order to regulate harvest- 
ing practices. Basket makers agree that regulation of extraction is necessary and pro- 
pose some actions of control, like protection against fire, excluding livestock, and com- 
mercial harvesting of adult stems. Establishment of permanent research plots is also 
necessary in order to continuing monitoring and evaluating the populations' responses 
to different harvesting treatments. These measures could lead to improved manage- 
ment by establishing cutting level as well as a systematic registration of the phenology 
of the species. This will be very important since the process of flowering is nearly com- 
plete in all populations of the ejido.The monitoring process should involve local people 

We acknowledge the support and participation of the basket makers in the Ejido of 
Platanarillo that allowed us to complete our work. This research was supported by a 
grant from the Mexican Secretary of Environment, Natural Resources and Fisheries to 
the Directorship of the Sierra de Manantlan Biosphere Reserve. 

R: Fl K! KlLs 

Anaya C, C. 1989. Estudio de la subfamilia Bambusoideae (Poaceae), con revision 
taxonomica para el estado de Jalisco, Mexico. Tesis de licenciatura. Facultad de 
Agronomi'a.Universidadde Guadalajara. Guadalajara lalis< o,f h i< o 

BhNZ, B.F.,F.J. San [ana M.,M.R.Pink)aL, J.ChVAu os E.,LRoi3iFsH., and D.DeNiz LI 994. Charac- 
terization of mestizo pla in tl t i i i u una i I 
Ethnobiol. 14:23-41. 

Bye,R. 1995.Ethnobot n tth , m M t, ,| j, t fests ln:Bullock,S.H.,H.A.Mooney 

and E.Medina (ed i >imII\ Liv It f i i I oi est s. Cambridge University Press. Cam- 

bridge. Pp.423-438. 

Cottam,G. and J.T.Curtis. 1956. The use of distance measures in phytosociological sam- 
pling. Ecology 37:451-460. 

Guzman M.,R., M.C. Anaya, and FJ. Saniana M. 1984,] | t^neio Otaii-n (Bambusoideae), en 
Mexico y Centroamerica. Bol. Inst. Bot., Univ. Guadalajara. 5(1 0):2-20. 

Aguilera-Herrera, N. 1989.Tratado de Edafologia de Mexico. Tomo I. Universidad Nacional 
Autdnoma de Mexico. Facultad de Ciencias. Departamento de Biologia. Laboratorio 
de Investigaciones de Edafologia. Mexico, D.F. 222 Pp. 

INEGI-SPP. 1 981 . Sintesis Geografica del Estado de Colima. Coordinacidn General de los 
Servicios Nacionales de Estadistkom hhhimIh <■ Hi 'imatica.SPP. Mexico. 

jANZtN, D.H. 1 976. Why bamboos wait so long to flower? Ann. Rev Ecol. Syst. 7:347-391 . 

Judziewicz,E.J.,LG Cl i>,\ [ ,t,i ;, andiVU a r. Few. American Bamboos.Smithsonian 
Institution Press. New York. 

KbbLEYj.E.andWJ. Bond. 1999. Mast flowering and semelpauiv in oamboos:The bamboo 
fire cycle hypothesis. Amer. Naturalist 154:383-391. 

Martinez R.,LM.,JJ. Sandoval, and R.D.Guevara G.1 991. Climas de la Reserva de la Bidsfera 
Sierra de Manantlan y su region de influencia. Laboratorio Natural Las Joyas. Univer- 
sidad de Guad il -i . , i i i t , h 

OiviraV.,M., S.Mori n. > ( . and B.I i-koa R 1 990 Sitios permanentes para la investigacidn 

silvi'cola. Manual para su establecimiento. Libros Tecnicos del Instituto Manantlan. 
Universidad de Guadalajara. Guadalajara, lalis< o. M< <ico 

Soderstrom, T.R. and C.E. CALDbRON. 1974. Primitive forest grasses and evolution of the 
Bambusoideae. Biotropica 6:141-153. 

Torres, B. 1985. Las pi v i ut n lf l i mt in iuL in ites del siglo XVI.En: 

Rojas R.,T.y W.T.Sanders (comps.) Histona de la agricultura.Epoca prehispanica, siglo 
XVI. ( (doccion Bibliotora del INAI I. Mexico, D.F. 

Saniana M„ F.and S. Lemus J. 1 992. La floracidn de los otates, un hallazgo sorprendente y un 
estado cn'tico de la planta.En:Cartapacios.Suplemento cultural, Ecos de la Costa. Colima. 


Richard Spellenberg 

; ' i* Biology, M5C3AF 


ons are reported on anthesis, stamen and style m 

ovement, and insect visitation in five 

erhavia, one a pantropical perennial (B.coccinea W 

'.intermedia M.E.Jones, B. spicata ChoisyB. torreyano 

-S.Wats., and B.wrightii A. Gray). Obser- 

I flowers are open for 

^nlv\i oo 

rtion of one day. Insect visitors were Hymenopter* 

a, Diptera, and Coleoptera. All species 

miii in i ill ii i flowered species had rr 

lore visitors. Autogamy is believed to 

lal method of reproduction, eitherthrough insect po 

lination or self-pollmati motil 'i . < • 

no evidence of wind p^Tm,! i 

>d populations. Chromosome numbers are faitl. In Ji i t Ml i i annuals. New chro- 

number reports are made for B. coccinea [n = 26), 6. 

diffusa l.(n = 27), B mt, > > - • 


n i" i i(n ca. 26), B. mathisiana F.B.Jones (n = c 

a. 26), fi. spicata {n = '<•) mil ' i ' ,' 

Cn = 27). 11 

: is suggested that high chromosome number, prevalent aut i mi, 1 11 t 1 


>r hybridization, produce a population structure of 

locally uniform ions that diflet 

slightly tc 


i lead to difficulty in classification. 

Se inform 

a de las observaciones del movimiento de los esta 

mbres y el estilo durante la antesis, asf 

la visita de insectos en cinco taxa de Boerhavia, 

lil - 

nco anuales norteamericanos (B. intermedia M.E.. 

ion( s, / p it a < hoisy, 6. torreyano S. 


I i rl ii lu.T, •> abren solo una parte de un dia Lo ir i i it r t in i 

Hymenoptera, Diptera,y Coleoptera.Todas las especies recil 

de fiores 

grandes tuvieron mas visitantes. Se cree que la 

autogamia es el metodo normal de 

No hay evident la cf im mo j i mm m i |i,i- i - 
Dn bastante altos,especialmente en las anuales. 
:cinea(n = 26),B.diffusaL(n = 27),B.intermedia 
,,-'./, ca 2o), B. mathisiana F.B. Jones (n = ca. 26), B. spicata (n = 26), y B. 

wrightii in = 27). Se sugiere que el alto numero cromosomatico, la autogamia predominante, con 
reproduccion cruzada ocasional o hibridacion.producen una estruchura poblacional de poblaciones 
ocalmente uniformes que difieren de ligeramente a mucho de otras poblaciones, un patron que 
puede llevar a dificultades en la clasificacion. 

Nyctaginaceae comprise a small family of approximately 30 genera and 390 species 
(Mabberly 1997) consisting mostly of American genera, several of which are noted for 

: problems. Among those geneici i ->,,,, , ( „ h, | m 
addition to being highly developed in North America, contains some rather difficult 
groups, for example, a pan-tropical complex of perennial forms (ft diffusa L.B.coccinea 
Mill., etc.) and at least two North American groups of annuals (B.spicata Choisy complex; 
ft erecta L. complex). As discussed by Ornduff (1 969), insights into reproductive aspects 
may help to understand variation seen within and bel een pi ipulations and this, then, 
may be useful in taxonomic interpretations. 

In the case of Boerhavia, populations in the held often appear homogenous within, 
but differ to a greater or lesser extent with neighboring populations. This population 
structure is conspicuous in the perennial B.coccinea where there are numerous races of 
maroon-flowered populations that differ in general robustness of plants, in nature and 
distribution of pubescence, or in number of fruits in the terminal clusters of the 
inflorescence. In addition, a few populations in this species vary markedly with respect 
to flower color and other characteristics. For example, an isolated consistently white- 
flowered population with bright green, lightly pubescent foliage occurs on a rock out- 
crop on the plains of southern New Mexico. Elsewhere in the region a yellow-flowered 
race with dull green more heavily pubescent foliage has been discovered adjacent to 
maroon-flowered less densely pubescent plants (white: New Mexico, Doha Ana Co, ca. 3 
mi S of Cambray on Providence Cone, 25 Aug 1 985, p< i ib< / and Zucker 8244 [NMC, 
NY]; yellow: New Mexico, Doha Ana Co., Doha Ana Mts., 5 slopes Summerford Mountain, 
14Sep ]969,Spellenberg2141 [NMQ. Within theannual species thereare alsoa number 
of examples. Boerhavia aiata S.Wats, (in the B.erecta L complex) grows on the rocky coast 
in and near Guaymas in southern Sonora, Mexico. Without conspicuous habitat differ- 
ences it contacts and intergrades very locally with ft intermedia M. E.Jones, common on 
the hillsides in the immediate vicinity (Mexico, Sonora,Guaymas,26 Aug 1 97 3>, Spellenberg 
and Willson 3627, 3629 [ft aiata], 3630, 363 i [intermediates], 3628, 3632 [ft intermedia] [all 
at NMC, variously distributed to CIIDIR,MEXU,IBUG,RSA, NY, UC, etc.]) (herbaria acronyms 
from Holmgren et al. 1 990). 

Perhaps because of the curiosity of nocturnal flowering in a number of species of 
Nyctaginaceae and/or the presence of chasmogamic and cleistogamic flowers on dif- 
ferent plants within populations of a species or,commonly,on the same plant,a number 
of authors have reported on floral tepr du ti ft Mures and insect visitation in several 
genera. Several papers report that Nyctaginaceae have flowers that attract insects but 
often self-pollinate by anthers contacting the stigma as the flower closes {Boerhavia, 
Chaturvedi 1 989; Mirabilis, section Mirabilis, Hernandez 1990) and/or through cleisto- 
gamy in plants that are also chasmogamous (Acleisanthes, Ammocodon, Selinocarpus, 
Spellenberg and Delson 1 977; Cyphomeris, Ma hrt and Spellenberg 1995; Mirabilis, sec- 
tion Oxybaphus, Cruden 1 973). Self-incompatibility is known also in Abronia (Tillet 1 967; 
Williamson & Bazeer 1997) and M 'ubii tti't)''i e L I ifei !964;Pilz 1978). 

Identification ot speumen ,oi Boerriaviu i often equivocal >pe< les are variable and 
often are different Kit i i ute -,- il|^ h t- | , i M t the fruit Differences 

ments in floras during the past 50 years attests to the difficulty of 
satisfactorily circumscribing species in some groups of Boerhavia. Often in such groups 
of plants (in general and not just in Boerhavia) a combination of biological characteris- 
tics contribute to the source of the difficulties faced by the taxonomist. Here I make a 
comparison of pollination and floral action of five New World taxa, one perennial and 
four annuals, and relate these observations in a general sense to the variation seen in 

Taxa observed were Boerhavia coccinea, B. intermedia, B. spicata, a small-flowered form 
called B. torreyana (S.Wats.) Standi, (considered a synonym of B. spicata), and B.wrightiiA. 
Gray. All were observed in Las Cruces, New Mexico, and are vouchered under my collec- 
tion numbers in the New Mexico State University herbarium (NMC). Among these, B. 
coccinea (7867), the only perennial in this study, is a pan-tropical species similar to B. 
diffusa, the former often considered a synonym of the latter (compare, for example, 
Whitehouse [1 996], both species recognized,and Wunderlin [1 998] or Diggs et al. [1 999], 
one species recognized). Boerhavia spicata Cholsy {7866) is an annual which, in its inclu- 
sive sense, includes several synonyms referring to phases more or less different from 
one another (e.g., B torreyana - 7868) but linked by various intermediate forms (Reed 
1 969). Boerhavia intermedia (7869), an annual of arid and semi-arid regions in southern 
North America, is sometimes considered as a variant of the widespread, weedy, B. erecta 
L [B. erecta var. intermedia (M. E. Jones) Kearney & Peebles]. Boerhavia wrightii (7870) is 
part of a small complex of species from North and South America that are fairly distinct 

Boerhavia plants respond to summer rains, flowering primarily in August and Sep- 
tember in southern New Mexico. In 1984 summer rains in the region were "good, "result- 
ing in ample late season growth for both perennials and annuals.Observations on polli- 
nation mechanisms were made daily during an eight day period (31 Aug - 7 Sep). The 
positions of stamens and stigma during the period of anthesis were observed with a 
10x hand lens. Boerhavia coccinea and B. spicata were studied in a small weedy area on 
the NMSU campus where the species were intermixed. The three other annuals were 

Larrea, where they 

Periods of observation. — Plants were observed daily, with observation of i 
lasting for 10 minutes. The observation periods were rotated from one spe 
next, with a few minutes allowed in between for relocation. The first period e 
ing began with a different species. Observations were made at each locatior 
campus) on alternate days. 

Insect visitation.— Records were kept of insect visitors, the duration of vi 
as possible where the insect next visited, and general identity. Records we 

individual subumbellate clusters in B.coccinea and B. intermedia) and visits to entire plants 
under observation. The latter data were not corrected fo> number of visitors relative to 
the number of total terminal inflorescences. In addition to observations, insects were 
also collected either with a net or i f him - I ilk I u d I nm identified to taxon 
as precisely as po il Juinl i I 11 it pmii in were analyzed using JMP 3.0 

cies.An a level of 0.1 was considered significant. 

Wind noHnhilinn. lo estimate the lole ol wind pollination, lour ql\A enn-t ooteo 
microscope slides wm inoii«,ih m u iiill, t if u ink. tl it were located within 
1 cm of inflorescences of a single plant of each species. Slides, each having a surface 
area of 12.5 cm ? wereapproxim >t< ill u <- I i i lit J lit i r< slides were placed 
in populations one day during the entire period from immediately prior to perianth 
opening to closure. 

Exclusion of pollinator-,, moss pollination was prevented by loosely wrapping an 
inflorescence in several layers of fine nylon stocking mesh immediately prior to anthesis. 
This effectively prevents passage of the large and spinulose Boerhavia pollen (pollen 
described in Nowickc I 0701. the neltmp remained on the plant until seed maturation, 
typically 9-10 days. The netting prevented loss if fruits were shed prior to removal of the 
net. At the same time, other inflorescences were marked, exposed to open pollination, 
then bagged in a sir r o I in i i II u, umi ' r tin flowers closed. Fruits 

werethen collected ft ml ph m mrmui it 1 opene 1 ! i immme whethereach con- 
tained a seed. 

Pollen/ ovule tntios sinuoo. ii\> m ,pe. no pi odm e one ovule per fruit, pollen / 
ovule ratios were calculated simply by counting the pollen grains produced by a flower. 

scope slide and the pollen grains were counted using a compound microscope at 1 00x 
magnification. Ten flowers for each species were examined. 

Hybridization— As a generalization, breeding barriers may be weak within autoga- 
mous species beyond the barrier provided by the breeding system itself (e.g., Lewis 1 963; 
Stebbins 1 957), which might account for some of the variation patterns with the B.spicata 
or B.erecta complexes. Woodson and Kidd (1961) suggested that hybridization occurs 
within mixed population of,' e , ( i mrennial) and B.erecta (annual), the putative hy- 
brids representing B. coccinea in the sense of Standley (191 8). To attempt to gain some 
perspective on th f i i nit l i i u i mi f i i s vas made in early 

September, 1984. In the mixed popular u m the Nt M< tote University campus, 

B.coccinea was used i i i i II i r m film 1 t i m pollen donor.Eleven 

ili - ii nuin i in in uinlh il m In l< i . u in i i ul m I i . il m tl on m . n 
anther dehiscence with fine forceps under a dissecting microscope. Stigmas were thor- 
oughly inspected with the dissecting microscope and the few pollen grains already on 
the stigmas were removed b\ pin th m fl I i moist m I dissecting needle. 

inflorescence of B.spicata to B. coccinea and brushing the 
recipient stigmas. The flower cluster was bagged in dout 
mature fruits were collected in 1 days. 

Floral "behavior."— In each of 
only a portion of a day (Fig. 1 

flowers in a single flower cluster. In B. spicata and B. < 
before it began in the other three (the perhaps v 
ronment may have influenced timing and durat 
with the opening of the corolla-like perianth, stamens and styles uncoil and, in the larger- 
flowered species,the final stigma position is slightly beyond the anthers. In the smaller- 
flowered species stigma and anthers are not as well separated initially (Fig. 1). As the 
morning progresses, the filaments and style curl and the anthers haphazardly contact 
the stigma. Self-pollination may occur at this time. Perianth closure begins in late morn- 
ing and progresses rapidly, so that by mid-afternoon the perianth is crumpled, the sta- 
mens and style usually contained within. Only rarely do flowers weakly open a second 
day.These observations fully support those of Chaturvedi (1 989), who reported that the 
widespread perennial, B. diffusa, is autogamous (as known from plants studied in the 
botanic garden, Allahabad University, India). 

Insect visitation. — Insect visitors varied in kind, freguency and duration (Table 1), 
with various bees and flies frequent visitors. This is concordant with Bittrich and Kuhn's 
(1 993) review that Boerhavia flowers fit the profile of bee and fly pollination. In a previ- 
ous observation, however, ants covered with pollen grains were noted entering flowers 
of B. coccinea. On such occasions,ants may also serve as pollen vectors (observation and 
comment on specimen, Mexico, Colima, 1 km SW of Tecoman, El Real, Spellenberg 2955, 
NMQ.The total number of insect visits varied considerably for the five taxa of Boerhavia 
studied (Table 2). Small sample size and considerable variation must be considered when 
viewing data in these tables. Raw data of total insect visit per species per day, summa- 
rized in Table 2, is not normally distributed (Shapiro-Wilk WTest, prob < W <0.0001 ).The 
Wilcoxon / Kruskal-Wallis Ranked SumsTest indicated significant differences in the data 
(% 2 = 9.1 96, df= 4, p = 0.0564). The Tukey-Kramer comparison of all pairs indicated that 
Boerhavia spicata differed most from all other species, significantly from from B. wrightii, 

Based on these preliminary observations, no conspicuously strong floral fidelity was 

observed by visiting insects. An in ec atin i /isits on one species might fly to several 

inflorescences on the same plant, then move to another plant of the same or different 
species,then return to the original plant,and so forth. Therefore, in< 
allow both for autogamy, xenogamy, and hybridization. 

Wind pollination— Very little Boerhavia pollen was trapped c 

slides. Grass and Salsola pollen was frequ 
grains were trapped upon the slides and 
miUei squared per hour was calculated (r 
0.025), B. intermedia (7: 0.035), B. spicata ( 
One slide from i ty/if/7 was rem ed 

i of giainvL|iaim,/hi/am):/u- 

vidual plants of each of five species of Boerhavia.H is equal to the number c 
observation periods.The average number of seconds per visit in given in 
Small Hymenoptera visitors consisted of Bethylidae, Andrenidae {Perditc 
(Dialictus, Lasioglossum, Halictus). 

B.coccinea B. intermedia B.spicata B.torreyana 

Total number of visits for all observation periods (average dur 


Scol iidae (Scolia) 14(5) 4(7) 


bphec iciae 

Calliph \* 

pollen, presumably from the landing of a pollen-laden insect. From this data it seems 
unlikely that wind pollination is significant in Boerhavia. Pollen sexine is spinulose, and 
pollen size is rather large, consistent with that reported for these and other species by 
Nowicke (1 970), and characteristic of insect pollinated plants. 

Pollination.— Pollen load on stigmas (as determined by direct observation of pol- 
len grains on stigmas with a 1 0x hand lens) progressively increased during a 

Hour of day 07-0800 08-0900 

'11 ii t 11 11 | m >i< ih in i u hi uihoind I il n > r i l i 1 1 I I 01 

samples of ft torreyana (Table 3), the species with the smallest flowers, data indicate the 
same trend,though somewhat erratic.This species also received the fewest insect visits dur- 
ing anthesis (Table 2). Four of the five spec ies, ' v < itath < a pi ion, still had a low to 
moderate percentage of stigmas without pollen at the imu ■ of oeiianth closure (Table 3). 

Even though some stigmas were unpollinated at time of flower closure, percent- 
age of filled fruits in each species was high for both open pollinated and pollinator- 
excluded inflorescences. Curling of stamens and the style places pollen on the stigma, 
with the assumption that autogamy results. In each case in the following pairs of data, 
the number of filled seeds precedes the number of unfilled seeds for plants protected 
from pollinators and for plants openly pollinated: in plants protected from pollinators - 
ft coccinea 20/0; ft intermedia 5/0; ft spicata 8/0; B. torreyana 1 9/0; B. wrightii 5/0; in plants 
openly pollinated— B.coccinea 1 M u '■ UU J 1 1 o\ ed; B.spicata 23/0; ft torreyana 
MM, wrightii 13/0. 

- Mi v ovule ration Pollen/ovule ratios lor the live Boerhaviu species examined 
ranged between 28:1 and 102:1. These figures lie between those proposed by Cruden 
(1977) for obligate and facultative autogamy (, lablo-lj.l oi eadi taxon more than 95%of 
the pollen grains stain, M ell in iiion blu iMMi Mnti I uooesting a high level of 

Boerhavia coccinea that had received pollen from ft spicata. All appeared normal. One 
was opened and the seed was normally filled. The other 10 were planted in native soil 

Number of grains on stigma 
Species, hour of day, number of stigmas scored Percent of stigmas with abovi 

The flowers of the five different types of Boerhavia each opened for a few hours in the 
morning, and then closed permanently. Movement of stamens and style apparently as- 
sure self-pollination. These observations conform with those of Chaturvedi (1989) for 
Boerhavia. A similar mechanism was noted by Cruden (1 973) and Hernandez (1 990) for 
some M/rab/7/s,Mahrt and Spellenberg (1995) for Cyphomeris, and Spellenbergand Delson 
(1977) for Ammocodon. Allowing for the exception in the specialized Asclepiadaceae 

2 species of Boerhavi 

Facultative autogamy 162 ±22 

Facultative xenogamy 797 ±87 

Xenogamy 5859 ±937 

(Wyatt et al.2000), pollen/ovule ratios suggest high levels of autogamy (Cruden 1 977). In 
an independent study to determine various sugar concentrations in nectar (unpublished, 
ratios provided on herbarium vouchers at NMC), I extracted nectar from all species;flowers 
of B. coccinea and B.spicata individually produce much more nectar than flowers of the 
other three; these two species had the most insect visitors, B.spicata significantly so. 
Bittrich and Kuhn (1993) review that Boerhavia flowers produce nectar in the narrow 

tions supported by this study. Insect visitation between plants within a species would 
allow for occasional outcrossing as noted for autogamous plants by Lewis (1 963). This 
would also allow for the potential of hybridization, which i\ supported by seed pro- 
diiM'd (but nol qerminatnj) in a wiy small trial ol artificial inl<M--,pecific pollination. Hy- 
bridization may take place occasionally in the field, as suggested by Woodson and Kidd 
(1 961 ).This is especially likely in closely related, little-differentiated populations, as might 
be the case between B. alata and B. intermedia in the Guaymas region in Sonora. Chro- 
mosome numbers known in Boerhavia are fairly high (n = 1 3,20,26,27,58,see Appendix 
I), especially for annuals, which for those known n = 26 or 27. A high n number would be 
an important contributor to a high recombination index,promoting a higher number of 
new gene combinations through segregation and recombination in a limited number 
of generations (Stebbms 1951) than would a low n number. Hybridization followed by 
recombination and repetitive inbreeding would be expected to produce a rather fine- 
grained patchwork of populations homogeneous within and more or less different be- 
tween. This kind of population structure, reviewed by Lewis (1 963) and Stebbins (1 957) 
in discussions of the relationship between autogamy and problems of classification, 
applies in Boerhavia. As noted by Lewis,autogamy per se creates no taxonomic problem, 

outcrossing, many local phenotypically differentiated populations may be temporarily 

and B.spicata (both sensu la to), and such complexes may be treated in the same mannei 
as those where complex patterns of variation result from outcrossing and/or hybridiza- 
tion without notable inbreeding (Lewis 1 963). In such situations, the taxonomist exer- 
cises considerable personal judgment, attempting to communicate in a classification a 
useful taxonomy that corresponds to broad limits on gene flow and/or fidelity to certair 
ecological situations. 


Known chromosome numbers in Boerhavia.My original counts are indicated by voucher 
citations,unless otherwise indicated, collection number is mine;voucher specimens are 
at NMC, with many widely distributed. Other counts were compiled from the literature, 

verted to expected gametic numbers for ease of comparison within the following listing. 
Within a species, all are organized alphabetically by country.Original counts were obtained 
from buds fixed in cold modified Carnoy's solution (4:3:1 - chloroform: ethanofglacial 
acetic acid), and stained and squashed in hydrochloric acid-carmine (Snow 1963). 

Boerhavia coccinea Mill.(perennial).n = 26. Arizona, Maricopa Co., 1 2 mi. N of Phoenix.2527; 
NewMexico,DonaAnaCo.,DonaAnaMts.NofLasCruces,794J;Bolivia(Fernandf O I mm 
Piqueras, 1981 ); Mexico Vera Cruz Paso deOvejas Pil ' + t i ^b New Mexico, Doha 

Boerhavia diffusa L.(perennial) n 3.lndid,(Srivisia a&Misra'l )66) anzania (Gill & Abubakar 

1 i i i H m it 1 t h n i ),ih'i | in i I i 1 roadside weed along 

highway H-93, s end of Waianu Range, 6396] Haleiwa at Waialu Bay, 6406; n = 58, India (Thombre 

nnual).n = 26. New Mexico I '■ in *n - h m - 

Boerhavia linearifolia A. Gray (perennial), n - ca. 26. New Mexico, Chaves Co., 9.2 mi 
Boerhavia mathisiana F.B.Jones (perennial). n = ca.26.Texas,San Patricio Co.,ca.2 mi 

Boerhavia repanda Willd. (perennial), n = 20. India (Gajapathy 1 962); n = 21 . India 
Rao 1963). 

Boerhavia spicata Choisy (annual), n = 26. New Mexico, Doha Ana Co, Las Cruces, 2C 
'6. Mexico, Puebla, 1 mi. W of Acatlan, Pilz&Strother 671;n = 27. New Mexico, Doha Ana i 
:es, 7866; 8291 [latter B. torreyana (S.Wats.) Standi, phase]. 

Boerhavia wrightii A. Gray (annual), n = 27. New Mexico, Doha Ana Co., Las ( i uces, 7, 

eds.The families and genera of vascular plants (ed. by K. Kubit/kT Sprinqm Verlag, 

Berlin. Pp.473-486. 
Carr,G.D. 1 978. Chromosome numbers of Hawaiian flowering plants and the significance 

of cytology in selected taxa. Amer. J. Bot. 65:236-242. 
o i ii l ! K ] C W). A new deviceof self pollination in Boer/iawVjd/T/imaL (Nyctaginaceae). 

Beitr. Biol. Pflanzen 64:55-58. 
Crudfn,R.W. 1973. Reproductive biology of weedy and cultivated A/f/Vafc>///5 (Nyctaginaceae). 

Amer. J. Bot. 60:802-809. 
1977. Pollen-ovule rations: A consei jtiv< nidi 1 i ' i ceding systems in 

flowering plants. Evolution 31:32-46. 
Dices, G.M., B.L. Lipscomb, and R.J. O'Ki-nnon. 1999. Shinners & Mahler's illustrated flora of 

North ( mitral Texas. Sida, 16. 
Fernandez Casas, J. and LFi nAndi I' i 1981. Estudio cariologico de algunas plantas 

Gajapaimy, C. 1962. Chromosome numbers of some south Indian plants. Current Sci. 31: 

Gin, L.S. and A.M. Abubakar, 1975. In: A. Love,ed.lOPB chromosome number reports XLVIII. 

Taxon 24:367-372. 
HiRNANi)i.',H.M.1990.Autopohnr"k i n ' <> ■ , ,i_ i i i imaceae). Acta Bot. 

Mex. 12:25-30. 
Holmgren, P.K., N.H. Hoi mgrfn, and L.C. Barnett. 1 990. Index herbariorum, Part l:The herbaria 

of the World. Regn. Vegetabile 1 20:1-693. 
Lewis, H.1963.Thetaxonomic problem of inbreeders or howto solve any taxonomic prob- 
lem. Regnum Veg. 27:37-44. 
Mabbi ri y, DJ. 1 997.The plant-book, 2 nd ed. Cambridge Univ. Press, Cambridge. 
Mahri, M. and R. Speli enberg. 1995. Taxonomy of Cyphomeris (Nyctaginaceae) based on 

mult m ii m il, i h it t i iht i 1,1 It) 679-697. 

Nowicke, J.W. 1 970. Pollen morphology in the Nyctaginaceae I. Nyctagineae (Mirabileae). 

Grana 10:79-88. 
Oim i,R. 1969. Repi kn I ml i in relation to systematics. Taxon 18:121-133. 
Ph /,G.E. 1 978.Systematics of Mirabilis subgenus Quamoclidion (Nyctaginaceae).Madroho 

Reed, CT. 1969. Nyctaqimn cic. In: ml . I undell, ed. Flora of Texas 2(1)4 51-220. 
Snow, R. 1963. Alcoholic hydrochloric •mid carmii im is a stain for chromosomes in squash 

preparations. Stain. Tech. 38:9-1 3. 
Spellenberg, R. and R.K. Dfi son. 1977. Aspects of reproduction in Chihuahuan Desert 

Nyctaginaceae, pp. 273-287 in R. H. Wauer and D. H. Riskind, eds. Transactions of the 

111 I N I ll | ill Ml Ol If Mill I ihu tU I Ill It 

and Mexico. U.S. Dept. Interior, National Park Service Transactions and Proceedings 

Srivistava, A.K. and K: 

C. Misi 

^a. 1966.Chron 

number in 

Boerhavia diffusa Li 

Cult. 32:31 5. 



:eae. North Arr 

lerican fl 



STEBBiNS l G.L1951.Va 


i in plant 


a University Press, N. N 

1957. Self-fertil 

ization and pc 



in the higher plants 

Naturalist 91:337- 


Tandon, S.L. and G.K. 

Rao. ' 


tudies ir 

i Boerhavia repanda Willd. Cum 


Thrombre, M.V.I 959. ( 

losome numbt 

?rs in sor 


>n flowering plants. S 




me species of Abronia ( 


ceae).Brittonia 19:29 

Whitehousf, C. 1996. 


ginaceae. In R. 

M. Polhil 

1, ed. Flora 

of tropical East Afri< 


am, Brookfield. 

Panama, pt. IV,fasc.4. Pp.5 

probes the rule. Syst. Bot. 25:1 71 -1 80. 

and submitted so that some of its c onclusinns could he referenced in an 
hAin M , in i - ■ Hi n ,mnmv followed wasthat of Reed 

r, changes that may be referenced by way of voucher numbers given in the 
?'-|uri'i- :nii it ' .' h , : ml originally called B.spicata and B. 

>ok.f.) S.Wats., 7868, fourth column of Fig. 1;8297, chromosome number, n = 


imn of Fig. 1, 2080, 7866, chromosome num- 

Delena Tui i . 1 999. Edible and Useful Plants of Texas and the Southwest. (ISBN 0-292- 
781 64-4, pbk.). University of Texas Press, P.O. Box 781 9, Austin,TX 7871 3-781 9, U.S.A. 
(512-471-4032,512-320-0668 fax; orders: 800-252-3206 or 800-687-6046 fax; ' 21 ' p >l -r i 4 2 pp Line drawings, 57 color photos. 
Edible and Useful Plan' i Mi it it ' i a practical guide. In its 

,eioni edition Kit i in f 1 tl i tin I [It i ul l< i in ii ition on plant character- 

istic s, habitat, and range in be used with a supplemental and more detailed flora ofTexas. 

Despite the pief u . in hi m mil i i- n A i il I i , ■ home, Alaska, the book 

undo htedh rittf t,l , 1 

• I MtT'-.,, 

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Jin' in hio|. i ji. il lio-i it , >ll t 

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The book breaks down in 

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H p,n , 1 Ho. . 1 1 \ i oD .or , 


>liy,.ind index. 

Part One, titled"! dihlo and Us. 

useful, hits is roughly a third o' 

the boo-, a 

id offers encyclopedic referent 


- 1- ■ IS f 1 ,1 

una then uses. Part Two, "Teas < 

Part Three, "Edible and 

Poisonous Berries and Other Fleshy Fruits" 

in trdiiP .aluablefortheca 

mper or r 



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g, plant toxins, and then lists lo 

Part Five, "Colorful Dyes with Texas Plants" 

is dotiintelv a strong point of tl 

ais book. F 

list ikscussim; 

t i ilT i i H il ml I < I i i I nnmg on to discuss 

influence dye colors, dyeing techniques and dye recipes. Part Six,"F 
plants for basketmahr i I ul I F l nnaktrni P iM >evu 

Industrial Resources from Texas Plants"closes the book. 

This would be a gre t hook for any 1 is univeishv leveP 
great book to take > in n I m t , tu| i I lu 

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perhaps a selected ethnography of local group, would be sl 
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is hotany <. 

edit.With 50 plusc 

aady and easily ace 


Id guide for identi 

■ in ) md hit n t ;e : 

sons -e. • 




-n><t>i)y Department 
Valdosta State University 
ildosta,GA 31 698-001 5, U.S.A. 

studies show C. sanguinolentu iH • < m 1 3<onomically 

ndistinguishable, and C 

i l r i ir n 'iii > Tfie widesprea 

weed, C. sanguinolentus, 

formerly thought to be restricted to the Eastern Hemisphere, is reporte 

"rented " ^ ^^ d ' Stributl ° n ' and eCOl ° 9y in ^ ^^ 

stern United States are 


Losestudiosdecampomuestran i,u- ,, . ^ '^ localmente 

comuny una mala hierba 

en el este de Louisiana y el sur de Mississippi y que se extiende hasta Alabama y Georgia. Estudios 

morfometricosydeherbanomuestidii | > ^ , • is son taxonomicamente 

indistinguibles, y C.louisianensis se trata como un sinonimo de mala hierba 

como nueva para Norte Ameiu a h-i m | ] uti u» n> i 

istribuciony ecologia en 

el sureste de los Estados. 


Cyperus sanguinolentus Vahl is widely distributed in the Eastern 

Hemisphere, where it 

has been cited as a weed (Holm et al. 1991; Mingyuan & Dehu 

970; Reed 1977;Kuhn 

1 982). It is known from northeastern Africa, the Middle East, India 

Sri Lanka, central Asia, 

southeastern Asia, China, Taiwan, Japan, Korea, the Phi ippines, li 

donesia, Malaysia, and 

Australia (Clarke 1 894; Holm et al. 1 991 ; Kukenthal 1 935-1 936; 

hwi 1965; Mingyuan & 

Dehu 1 970; Kern 1 974; Reed 1 977; Kunn 1 982; Haines & Lye 1 98: 

; Wilson 1993) but has 

not been previously reported from the Western Hemisphere. 

Cyperus sanguinolenlih is highly vaiiable. Kukenthal (1935-1936) segregated five 
varieties and named seven forms, including six under the typical variety. Kukenthal's 
(1935-1936) infraspecific taxonomy of Csanguinolentus is difficult to use, since he pro- 

vi.ied nan has sevs nor paiallel oescaptions ol the l axa. Kern (loop (loaned tour s^Pspe 
eies, including ihe typical one, for Malaysia, and others (e.g., Ohwi 1965; Haines & Lye 
1977) have treated i I in i I n ih i i hi I m| inftaspecific taxono- 

mies of Kukenthal (1935-1936) and Kern (1974). Further research on this widespread 
and variable species throughout its range is needed for a more complete understand- 
ing of its infraspecific variation; however, such is beyond the scope of our study to deter- 
mine the range,clMn i i i i I ti I , tat i M , i in 

Its bifid style and lenticular achene with achonn angle adjacent to rachilla clearly 
place Csanguinolentus into subgenus Pycreus. Clarke (1 894, 1 908) segregated Pycreus as 
a genus and treated the taxon as Pycreus songuinolentus Nees in subgenus Reticulatae 
section Vestitae. Kukenthal (1 935-1 936) adopted a broader definition of Cyperus, incor- 
porating this taxon into ubgonu '■> a eetion \(/A ./(/ of that oenus.The floral scales 
of Csanguinolcntt, n I in dium no n II I i I nooves (or sulci) typi- 
cal of section Sulco ml s it eait pith i 'i blood-red floral scale pigmenta- 
tion. Although some a^ .on n tm i i I a i o loetghebeur 1986, 1989; Adams 
1994;Bruhl 1995) fragment Cyperus and segregate Pycreus at the rank of genus, we have 
followed the more conservative generic taxonomy of Kukenthal (1935-1936) and 

& Lye 1 977;Tucker 1 983, 1 987, 1 994). 

In 1977, Thieret described a ne\ pecies, C". /o /on. hum specimens he col- 
lected at two close sites in Tangipahoa Parish, Louisiana. Thieret (1977) placed C. 
louisianensis in subgc nu , a m t i it nuihni h lumolentus, and pro- 

vided several contrasting chaiacteristu s separating it Horn Csanguinolentus (Table 2) 
andadichotomous K listinuuishii i it fi an relate I" ith m a in species in subge- 
nus Pycreus. Cyperus louisianensis was listed by the Department of Interior, United States 
Fish & Wildlife Seivic r in , in ni unn, i 1 in i nt d species (Anony- 

mous 1993). Until Bryson and Carter (1994) showed it was widespread and weedy in 
southern Mississippi , , , - ^ • i, th oiuht t ^ he a narrow endemic restricted to 

In 1993,the first autho 
to prepare a status survey ho as iana > ^hich provided th. initial financial supp 
for this study.The ma|.n i . i 1 1 i- i 't it a .r ,<o .sea t, i , af a Iditional population: 

Clouisianensis and additional . II r I hat ao ie it tatus as a potenti 

rare plant, and to examine its taxonomic relationship with the Old World weec 
songuinolentus. In this report, we provide a complete record of our field and herbarh 
investigations intuth ti tin unno I i ir lta>unon a I P i hi| of ' . ea ' 
vith i < mguinolentus. 

Kiikenthal (1935-1936) 

). melanocepholus] 

eysmannii (Boeck.) K 

studies.— Systematic intensive field surveys for C. louisianensis populations in south- 
m Louisiana and southern Mississippi were made by the authors during the peri- 
5-21 September 1993 and 14-1 8 October 1993. Subsequently, the authors have 
arch sporadically for this taxon when time and circumstances have al- 
lowed. Thieret's (1977) published account and more recent collections from the type 
locality with additional documentation, kindly provided by Nelwyn Gilmore, Louisiana 
Natural Heritage Program, were used to relocate the holotype locality in Tangipahoa 
Parish, Louisiana. Attempts to re-locate Thieret's paratype locality were unsuccessful, which 
is not surprising since habitat in the vicinity of this site was substantially altered by high- 
way and commercial construction activities.Observations at the holotype locality showed 
the species to be locally abundant along the margin of a shallow artificial pond and 
nearby ditches in the flatwoods, habitat greatly altered by humans. Searches of potential 

habitat began outward from the holotype locality. Habitat descriptions, estimates of 
population size,and voucher specimens were made when populations were found. 

Greenhouse studies.— Transplants and plants of Clouisianensis started from seeds 
were maintained under . r m r ill. I it litmus in a greenhouse at the United States De- 
partment of Agriculture, Agricultural Research Service, Jamte Whitten Delta States Re- 
search Center it Slot n l. i i i , i )\ m m u ih J nt by the second au- 
thor were made in order to understand better the life history and phenology of C. 
louisianensis, especially to determine whether it is annual or perennial. 

In greenhouse experiments, seeds of Clouisianensis were sewn in flats on top of a 6 
cm-deep mixture of a Bosket silt loam soil (Mollic Hapludaf) and sphagnum (50% v/v) in 
the first week of March, June, September, and December in 1994, 1995, and 1996. Trays 
were watered from beneath to prevent seed and soil disturbance. Individual C. 
'<< (inciiM (idliiHi v i m ml ' t« mnsplantmi into . in diameter pots and 
watered from beneath.The greenhouse was maintained at 30 to 35° C day and 25 to 30° 
C night, at 60 to 75% relative humidity, and without supplemental lighting. In 1 994 and 
1995, flowering ami hum i 1 1 i I , m transplanted from several sites 

in I I ii ^ unty, I i ( | ii l i I inmi in I n i I n i I I mil irn U i 

pots and maintained under the same greenhouse corn I it inns m [Tints grown from seeds. 
Data were taken on date of seedling emergence, date of flowering and fruiting, and 
plant longevity. 

Herbarium and morphometric studies.— Specimens I louisianensis and C. 
sanguinolentus were borrowed from selected herbaria (BRIT,GH,MO, NY and US) in order 

sanguinolentus. Quantitative data on the achene characteristics used by Thieret (1977) 
W) distinct ii in lorn an, , Ik m ' >an itiinoh nlu ' k m 1 < fiom lln se ■ pnt iiiipii 
and from our own collections. Five achenes from each of 1 3 specimen 
and 20 specimens of C sanguinolentus were examined. A Bausch & L 
(6X-30X) dissecting microscope with ocular micrometer was used to measure various 
achene dimensions (Table 3, Fig. 1), and ratios (Table 3) derived from these measure- 
ments were used to analyze differences in the plane shapes of achenes used by Thieret 
(Table 2) to distinguish Clouisianensis and C smm,, - n muni I mloase 1 1.21 (Anony- 

mous 1996) was used to analyze these data statistically and to construct scatter dia- 
grams for comparing variation in r lot ianensi ind I n!u Representative 
spikelets from selected specimens were photographed using an Olympus™ SZ40 
stereozoom dissecting microscope eguipped with a Kodak™ DC1 20 zoom digital cam- 
era. Our more numerous recent collections were used with Thieret's (1977) original de- 
sc ription and type materials to prepare a revised description and an illustration. 

Distribution and ecology of Cyperus louisianensis— Since 1993, our field surveys have 


...» -o- < 

frontal view 



g Cyperus louisianensis ami i. 

ACHTH Achene thick -rv- hr-n; 

ACHTH/ACHL Ratio used b\ Tl 

ACHL/ACHW Ratio approxi ,, 


southern Mississippi and isolated populat 
Georgia. Cyperus louisianensis appears to I: 
sippi, and adjacent St. Tammany Parish, 
iedy, often locally abunc 
specially those maintained by it 
lere it appears to be aggressive 

ant, and restricted to periodically disturbed 
owing, such as ditches and edges of artificial 
and freguently forms dense stands often in 
ve weeds (Table 4). 

Greenhouse observations —In greenhouse experiments, C.louisian plant 

emerged from mid- and late May until mid-September each year; plants flowered from 
late August until mid-December, with peak flowering in early September to early Octo- 
ber; and plants subsequently Iniimd > j>< ■ , ', . mvns/s plants emerging later in the 
year were shorter at flower initiation than those emerging earlier, suggesting that C. 
louisianensis is photoperiodic. Most of our field collections were made from September 
16 through December 9 (see specimen citations lot the U.S.A.), with peak fruiting ob- 
served and mature plants collected in mid-October; however, a few fruiting plants were 
observed and/or collected in April and Ma\ foll> in i >i mild '.-.inter (1993-1994). If 
indeed C. louisianensis is photoperiodic, this may account for the few collections of it 
from May until mid-September by us and by other botanists. All C. louisianensis plants 
transferred into the greenhouse from the field and those grown from seeds in green- 
house experiments died shortly after fruiting, showed no evidence of c 

Relationship with Cyperus sanguinolentus— Jhlerei (1 977) was limited by a small 
number of specimens collected from only two sites located less than 1 miles apart.Our 
larger sample from a wider geographical area shows that C. louisianensis is more variable 
than previously thought in the diagnostic achene characters used byThieret (1977) to 
separate it from C. sanguinolentus. Figure 2 shows some of the spikelet variation ob- 
served in herbarium specimens of C sanguinolentus and C. louisianensis. Of the numer- 
ous specimens of C.sanguinolentus we have examined from throughout the Old World, 
certain ones from Japan [e.g., M. Furuse s.n„ 30 Sep 1 959 (GH); M. Furuse s.n., 1 1 Oct 1 960 
(GH); Hutoh 1 1517 (NY); Okamoto NSM 584 (BRIT, NY)] are indistinguishable from C. 
louisianensis in general and in spikelet (Fig. 2) and achene characteristics used byThieret 
(1 977). Our field and herbarium observations show considerable variation in the degree 
of development of pigmentation and sulcus in floral scales. At least some of the variation 
in floral scale pigmentation appears related to photopenod and temperature. Field ob- 
servations in the southeastern United States show that in mid-September the floral scales 
are typically pale with only faint pigmentation, and by mid-October they are deeply 
pigmented (cf. Figs. 2A and 2B). Opposite sides of an individual spikelet may also vary 
greatly in pigmentation (cf. Figs. 2G and 2H). Scatter diagrams (Figs. 3-6) show C. 
C.sanguinolentus are indistinguishable with regard to the critical achene 

Cyperus louisianensis is widely distributed in the coastal plain < 
States and is often locally abundant in habitats subject to periodic artificial disturbat 
and maintenance. Distribution, frequency, and habitat of C.louisianensis are indicative 
a semi-aggressive weed, not of a narrow endemic species. Furthermore, it appears t 
the range of this taxon is apparently expandin j in in >uth< rn nil I lah 
that its dispersal has possibly resulted in part from road construction and maintenai 

Andropogon virginicus L 

Axonopusfissifolius (FUldi'l Kuhlm. 
Bacopacaroliniana (Walt.) Robins. 

Boltonia diffusa Ell. 

/./pp/o nodiflora (L.) G'eere 

Magnolia virginiana L. 

Mitreola sessilifolia (GniH.) G. I )on 
Oxypolis filiformis (W.-iltei) I r it 1 

Phyllanthus urinaria L. 
Polygonum hydropiperoides 
Polypremum procumbent 1 . 
Rhynchospora corniculata (I 


Fig. 2. Spikelet variation in (yperussanguinolenl 
U.S.A., 18 October 1993, Carter 11 562. -C. U.S./ 
7 7577.-F. China, 7sang20665.-G&H. Opposite 

ides of same spikelet, Japan, K. Okamoto NSM 584. Scale bar = 1 mm. 

• . .' 

c louis 

• sang J 

X:V:-:' •. 

»',v ;d*» oa 


' * „ ° § Hs. 


&° \ • 




.05 1.15 1.25 1.35 1.45 1.55 1.65 1.75 1.85 

regard to critical ratios defining 




,ou 1S 
| + s ang [ 


o th ?f iitk il ".Tios defining achene shape; character abbreviations keyed ir 

activities, as suggested for C entrerianus Boeck. (Carter 1990). At Kings Bay Submarine 
Base, where extensive populations were found in Camden County, Georgia, road rights- 
of-way are maintained by work crews traveling from Alabama under contractual service 
agreements with the Department of Defense (pers. comm., R. Wilkerson).Thus, it is pos- 
sible that achenes of Clouisianensis were accidentally dispersed into Georgia from Ala- 
bama, or elsewhere, with the transport of mowing eguipment. 

Thieret (1977) distinguished Clouisianensis from C.sanguinolentus based on its more 
overlapping floral scales and its more elliptical and more flattened achenes (Table 2). 
Our results indicate the New World populations called Clouisianensis are encompassed 
within the total range of variation exhibited by the more variable widespread Old World 
weed, C. sanguinolentus, and are most similar to certain specimens from Japan. Further- 
more, as shown in Figures 3-6, the United States specimens are less variable than those 

from the Old World, which is consistent .- ii' i u hIm puru if I' (Mayr 1 942; Davis & 

Heywood 1 973) and would be expected in a case of accidental long-distance dispersal. 
Although typification of the plethora of synonyms and accepted infraspeciftc names 
under C.sanguinolentus (Table 1 ) is beyond the scope of this study, it appears the United 
States specimens are closest to C.sanguinolentus var. sanguinolentus {fide Kukenthal 1 935- 
1936) or C.sanguinolentus ssp. sanguinolentus {Me Kern 1974). 

The presence of C.sanguinolentus in the southeastern United States is not unex- 
pected for the following reasons. (1) It has been cited as an agricultural weed in the 
Eastern Hemisphere (Mingyuan&Dehu 1970; Kern 1974;Kuhn 1982;Holm etal. 1991). (2) 
Reed (1977) listed it among foreign weeds posing "potential problems in the United 
States." (3) There are numerous other examples of weedy Cyperus spp.and other sedges 
in the southeastern United States introduced from Asia or elsewhere (Carter 1 990;Bryson 
& Carter 1 992; Bryson & Carter 1 994; Carter et al. 1 996; Carter & Bryson 1 996; Bryson et al, 
1996;Bryson et al. 1997; McKenzie et al. 1 998). (4) Krai (1971) reported Fimbristylis spp. 
(Cyperaceae), common in current and former rice-growing areas of the United States, 
that were likely introduced from Asia with rice {Oryza sativa L) agriculture. (5) Histori- 

dant (Anonymous 1959, 1982). 

Cyperus louisianensis is much more widespread than previously thought (Thieret 1 977; 
Bryson & Carter 1994). It is locally common in southern Mississippi and southeastern 
Louisiana, and satellite populations have been found in southern Alabama and south- 
eastern Georgia. Its habitat and freguency are characteristic of a weed,and its range and 
freguency are likely to increase, particularly in the outer Coastal Plain of the southeast- 
Specimens of C. louisianensis from the United States are morphologically indistin- 
guishable from certain Old World specimens of C.sanguinolentus. Thus, we think the two 

k. Habit, base of plant (Bryson 14608 &MacDonald).-B. Inflorescences (left, B 

n 13276).-L Achene with stigma, style, and cross section outlined (Bryson 13276).-0. 
pikelet (Bryson 7i276).-F.Stem section (Bryson 13276). 

inty ofCyperussanguinolentus in the southeastern United States. 

2b. Stamens 3; style 

);floral scales usually at least marginally suffused with re 

divided nearly to base;style branches conspicuously exs 
cale, exposed portions about as long as floral scale; f 
of northeastern and upper mid-western United States. _ 

han half iLs length; style branches n 

long as floral scale;plants 

States. C.sanguinolentus 

CyperussanguinolentusVahl, Enum.PI. 2:351. 1805. Pyaeu^ <• i - - om ikkfJ. > Linn * > 
l 0^18^' I ilfj[|*ii | N ' H .1 ■ I i •[ M i , • ) 10 ft, Oct 1894, Gamble 

/5//7(L)[typ.cons.prop.,Kukkonen 1995], 
Cyperus louisianensis J.W.ThiereL, Proc. Louisiana Acad. Sci. 40:23. 1 977,Typf: U.S.A. Louisi- 
ANA.Tangipahoa Parish:ca.7 mi E of Ponchatoula, along road to Lee's Landing, 1 Oct 
1 972, Thieret 33585 (Holotype: GH!; Isotypes: DUKE, KNK, LAF, NC, OS). Paratypes: U.S.A. 
I i iTangipil i L .ii h i in tk i I * 'i M uh i ' ii II- highwayandroad 
to Lee's Landing, ca. 7 mi I of Ponchatoula, 18 Oct 1970, Thieret 32609 (DUKE, GH!, 

Annual herb,appt hii i i I m i 1 | I ^ |i i n ' iecumbent vegeta- 

tive lateral branches from lower nodes. Stems (6.5-)12-38(-60) cm long, 0.7-2.0 mm 
wide, trigonous. Leaves (1 -)3-7; bases sheathing; blades linear, (3-)5-1 1 (-1 6.5) cm long, 
(1-)2-3.5 mm wide. Primary inflorescence bracts 2-3(-4), linear; longest (1-)3-12(-19) 

cm long, 1-3 mm widt nfkm- u> teimnii mo . mn. u apitate, or with 1-3 

pedunculate rays to 4 cm It 10(-15) mm long, 

2.2-3.0 mm wide, with 8-32 floral scales. Floral scales mostly closely imbricate, membra- 
nous, ovate, 1.8-2.7 mm long, carinatc; kecd qrrtMi, 3 5 nerved; sides variable in color, 
usually variegated whitish, reddish brown to sanguineus, each with a narrowly elliptic 
translucent sulcus d> -id- g in-i - lihfuin, itani ul n tenerally becoming 
more conspicuous late season. Stamens 3;anthers 0.3-0.6 mm long. Style bifid one-third 
to one-half its length stigmas exseited <\t hem lent inula i In. onvex, 1.0-1.4 mm long, 
0.6-0.9 mm wide, 0.3-0.5 mm tin. I elliptii loobovnk uall isymmetrical near apex 
al i i id< adj icent u rachiila ,uda e retk ul it. irayish i i vn to brown. Fig 

>'' 'otio' j"/'s,'o'- '.ik I oimi'ii'., . - , <j m tropical and subtropical re- 
gions of the Eastern Hemisphere; central and eastern A 
donesia, Malaysia, Philippines, Australia, and eastern A 
coastal plain of the southeastern United States of Nort 

Georgia (rig. 8). In tin utln t n i I ink ottei ko ii\ uninon and weedy in 

Phenology. — In tin n, ■ i k ru ' k t> d it- tl • - in i it I tit, tin i In m - | 

/SC). Mobile Co.: Mobile, Battleship Part 

c, Hwy. US 90, 30 May 1 994,Mears 94-25 (ctb, VSC); 1 2 

»5, Carter /2705(VSC).GEORGI A. Camdt 

?n Co.: Kings Bay Submarine Base,0.2 mi E jet. U.S.S. 

. Stimson Dr. and James Madison Rd., U 

.S.S.Henry I. Stimson Dr., 11 Oct 1996, Carter 1 3873 

ist Njct.U.S.S.Benjamin Franklin Rd.and 

U.S.S. Georgia Ave., U.S.S. Georgia Ave., 25 Oct 1 996, 

3939 (VSC); ca. 1 00 m N jet. U.5.S. Benjan 

lin Franklin Rd.and U.S.S.James Madison Rd„ U.S.S. 


Carter i 

James Madison Rd., 25 Oct 1 996, Carter 13940 (VSC); ca. 200 m S jet. U.S.S. Henry L. Stimson Dr. and 
U.S.S. Kamehameha Ave., U.S.S. Kamehameha Ave., 25 Oct 1 996, Carter 1394 1 (VSC); 0.09 mi N jet. 
U.S.S. Daniel Webster Rd. and U.S.S. Benjamin Franklin Rd., U.S.S. Daniel Webster Rd., 25 Oct 1 996, 
Carter 13954 (VSC); ca. 300 m E Franklin Gate, S side U.S.S. Benjamin Franklin Rd.,4Dec 1996,Carfer 
13962 (VSC). LOUISIANA. St. Tammany Parish: Goodbee, 1 2 Oct 1 960, Hebert 377 (MISS); Hwy. US 
1 90,0.35 mi E jet. Hwy. US 1 90 and LA 1 077 in Goodbee, 1 8 Sep 1 993,Carter 1 1367 (VSC); Slidell, Hwy. 
US 1 90E, 0.84 mi W jet. Hwy. US 190E and 1-1 0,1 5 Oct 1993, Carter 11490 (VSC); Slidell, ICC Railroad 
right I u i) I I miHi i II LA 433 and US 11, 15 Oct 1993, Carter/ 7505 (VSC); Slidell, Hwy. US 
1 90, 0.1 mi W jet. Hwys. US 190 and US 11, 16 Oct 1993, Carter 775 1 ' " lidellH n l< ' 
mi W jet. Hwys. US 1 90 and US 1 1 , 1 6 Oct 1 993, Carter 1 1540 (VSC); Lacombe, Hwy. US 1 90 at Tran- 
quility Road, 16 Oct 1993, Carter 1 1541 (VSC); Slidell, Hwy. US 1 1,250 ft. S jet. I Iwy. us n lJMl K .nnllo 
Avenue, 17 Oct 1993, Carter 11558&Bryson (VSC), Brysonl3218& Carter (ctb, VSC); Slidell, Hwy. 1-1 0,S 
jet. Hwys. I-10 and US 190, 17 Oct 1993, Carter 7 756 7 &Sryson(VSQ lid II 10.n 

533, 17 Oct. 1 994,8rysor) 14565&MacDonald (ctb, VSQ.Tangipahoa Parish: ca.7 mi SEPonchatoula, 
ca. 1 mi N Lees Landing, 1 6 Nov 1 989,Gilmore 3977 (VSC), Gilmore 3978 (ctb, VSC); 1 9 Sep 1 993, Carter 
1 1374 (VSC) Ponchato I II I i) n I j t I \ . II till re. t 17 Sep 1993,Carfer 

1 1355 (VSC); Ponchatoula, Hwy. LA 22, W jet. Hwy. LA 22 (E. Pine Street) and West Street, 1 8 Sep 1 993, 
Carter 1 1372 (VSC). MISSISSIPPI. Hancock Co.: Hwy. MS 43, 0.44 mi N jet. Hwy. US 90 and MS 43 in 
Waveiand, 16 Sep N f) i irt \2\ C) 08 mi N jct.US90 and MS 43 in Waveiand, 16 Oct 1993, 

Bryson 13166& Carter (ctb, SWSL,VSC), 1 7 May 1 994, Bryson 13535 (ctb, VSC); Hwy. MS 43, 5.25 mi E jet. 
Hwys. MS 43 and 1-59 in Picayune, 1 8 Oct 1 993, Carter 1 1 567 & Bryson (VSC), Bryson 13265 & Carter 
(ctb, SWSUVSC); 9.0 mi NW jet. Hwys. 43 and 603 in Kiln, Hwy. MS 43, pipeline crossing, 1 8 Oct 1 993, 
Carter 11 568 & Bryson (VSC); 17 May 1994, Bryson 13542 (ctb, VSC); 21 Oct 1997, Bryson 16217 (ctb, 
SWSL,VSC);NofKiln,0.6mi.Sjct.ofHwys.MS43andMS603,18Oct 1993,/3ryson 1 3267 & Carter (ctb, 
SWSLi hln H M \ '"hi t I MS 43 and MS 603, 18 Oct 1993, Carter 1 1569 & Bryson 
(VSC); Kiln, Hwy. MS 43, vicinity Shifalo Baptist Church and Kiln Post Office, 1 8 Oct 1 993, Carter 1 1570 
&Bryson (VSC), Bryson & Carter 13268 (ctb, SWSL, VSC); 21 Oct 1 997, firyson 762/6 (ctb, VSC); jet. of 
Hwys. MS 43 and l-l between Kiln and Waveiand, 1 8 Oct 1 993, Carter 1 1571 & Bryson (VSC); SE jet. 
Hwys. LI and MS 43, 1 8 Oct 1 993, Bryson & Carter 13271 (ctb, SWSL, VSC), 1 7 May 1 994,8ry5on 13534 
(ctb, VSC); E of Picayune, 5.8 mi E jet. Hwys. 1-59 and MS 43, 1 6 Oct 1 994, Bryson 14537 &MacDonald 
(ctb, SWSUVSC); Mississippi Welcome Center, SE jet. Hwys. 1-10 and MS 607, 17 Oct 1994, Bryson 
14567 & MacDonald (a\ ' I ( i hln, NW jet. Hwy.MS 43 and Kiln-Delisle Road, 17 Oct 1994, 
Bryson 14597 & MacDonald (ctb, SWSL, VSC); Waveiand, Nicholson Avenue, 0.2 mi S jet. Hwys. US 90 
and MS 43, 1 8 Oct 1 994, Bryson 14608 & MacDonald (ctb, SWSL, VSC); Waveiand, Central Avenue, 
between Central Avenue and RRjustW Washington Street, 1 8 Oct 1 994,Bryson 1 461 0& MacDonald 
(ctb, SWSL, VSC); Waveiand, NW jet. Hwys. US 90 and MS 43, 7 Dec 1 994, Bryson 14709 (ctb, SWSL, 
VSC); Waveiand, NW jet. Hwys. US 90 and MS 43, 21 Oct 1 997, Bryson 162 14 (ctb, VSC), 20 Nov 1 998, 
Bryson 16939&Sudbrink (ctb, SWSL, VSC); Waveiand, 1.2 mi S Hwy. US 90 on Nicholson Avenue, then 
1.6 mi Eon Central Avenue,21 Oct 1997,Sryson 762 75 (ctb, SWSUVSC). Harrison Co.:Orange Grove 
Community Center Park,W Hwy. US 49, 0.3 mi N jet. Hwys. US 49 and 1-1 0,1 6 Oct 1 993, Bryson 13164 
& Carter (ctb, SWSL, VSC), Carter 11544& Bryson (VSC); Orange Grove, Hwy. US 49, 1 .1 mi S jet. Hwy. US 
49 and O'Neal Road, 1 8 Oct 1 993, Bryson 13276 (ctb, SWSL, VSC); N Gulfport, Harrison Drive, 0.3 mi W 
ict. Harrison Drive and Ml. King Jr. Blvd., 18 Oct 1993, Carter 1 1574 (VSC);Popps Ferry Road, 3.32 mi II |i t ill I il nil O , t , llU , n 7(VSC),NW 

jet. Hwys. 1-10 and US 19, 18 Oct I 1 "' i il W I \ u I . i I ( k h SE jet. Klondyke 

and Commission Road, 18 Oct 1994, Bryson 14606 & MacDonald (ctb, SWSL, VSC). Jackson Co.: 
Pascagoula,SE jet. Washinc uon Ave, and louisc M.,vk. Bayou Casotte,T8S R5W517, 16 Sep 1991, 
Bryson 11 032 (ctb N\ N i I ef i t <n> i I « firm Road 2 mi N 

jet. Old Fort Bayou Road and Rosefarm Road, 1 8 (A I '-<- ,m - ' ^ . { , , „ , „, 
miWjct.OldFortbi\oul r.nhl iM i ,,11 t Mo / /580 (VSC); vicinity 

St. Martin, Old Fort Bayou Road, 0,19 mi I it 'til r M i I il 1, Ii > \ Jacket Drive, 18 Oct 
1 993, Carter J /58/ (V'.c h i iouIj M i m I > 1 / / * ,', ,/ 1 u lb,VSC);Moss Point, 

SEjct.Hwys.HOandM In tl 1 ' il I V O I atimo 1 2 mi 

N jet. I Iwys. II and M n()n I ' mi N linker mil ( ook Road alonu lm ker Road, 16 Oct 1994, 
Bryson! 4559 & Ma, IX n Ml \\ I \ • < I , , , ui t n 1 1 m - I mi Wjct of Hwys.US 90 
and MS 63, 6 Nov PM , , ,1 LiNOum pnngs, Hwy. 1-10, E mi 

marker 50, 1 9 Oct 1 994,Bryson 14636 (ctb,SWSL,VSC). Pearl River Co.: Picayune, Hwy. MS 43,0.50 mi 
Wjct.Hwys.MS43and 1 u 1' a < son 16874 & Sudbrmk 

(ctb, VSC); Picayune, Iron nv t ,d , H.\ ' <• m N M - M 1 and I 59, 18 Oct 1993, 

Bryson 13222 & Car k < i o L on , > ( >,\ \ i m, Bryson 16900 & 

Sudbrmk (ctb, SWSL VM No M n « , > il | i ) Picayune, N side of 

I 1 1 ) n vi I .11 v \l 1 i II I 1 ) v k uttr (ctb SWSL, VSC); 

Picayune, SW jet. Hwys. 1-59 and MS 43 S, 19 Oct 1 994, Bryson 14634 (ctb, SWSL, VSC); Picayune, 21 
Oct 1 997, Bryson h ■ • . ■ Stone Co.: Wiggins, 

I rlC l 1 ' in. I ri i \ r 1" < - , , , Mctb.VSC). 

EURASIA. RUSSIA, ii I I n ' h I \ I |h, t p 1929, Baianova 647 

(NY). BHARAT (INDIA). h< n ml > u, lh nlntri 1 r ml n. ar Dandiganahalli, 

Ambala, 2-10-196 >ahlgam, 16 Aug 

1920, Stewart &Stewan I r n i n I ) tl I ik i Nnl '1 u o / (NY) Kulu, Rotang 

i'.r/,'M'P I « || PAKISTAN. Bain Ian < a 1 5 mil Skmdu all (a 7500 ft, 26 Aug 1955, 

I/Vebsfer6585 (GH) sk aid u i hi k kill m , ' k , n n i mqa Valley and road to 

NangaParbat,bek NEPAL. Kali Gandaki, 

) Aug 1954, Stainton, 
Sykes& Williams 7591 (NY);Bongakhani,22 Aug H5 1 ' f ., ,o,j (BRIT,NY);Aruna 

Valley, Sedua, NW of k, n ( 1 fj i hi ii \pr 1953, Gardner 141 

(NY); Argam, near Pokloni, 1 1 1 4 ,~, -.1' ,<<• N , hjarkut District, Maina 

gaon,12Aug 1979,Ro7far)onc/ara&/?oy45S5(NY),Rukun,M it ion m- H nop 1982, Raphandara 
& Malla 6535 (NY); Sindjuli District, Patlebhanjyanq Din I > . j. o hajbhandara 3343 (NY); 
Dolakha District, hamuli iq ir r > Hum In Jul I" / n , o , , , I j\ ) CEYLON. Amparia 

District, Senanaikc > im i Ir hi i I I I M - I i I m i ") L»e 1 L U 

Comanor690 (NY); ( . nlial I'm i nC e K inch I > ,Uu i ca r . mi SE Gampola, 24 Oct 1 974, Davidse & 
Sumithraarachchi 7924 (NY);Sabaraqar lit |[ i ivaya,22 Oct 1974, 

Heart &Cooray 13460 (\ {) mpoai Distr i onanaik. n hn Padagoda, 6 Feb 1971, Koyama & 

Balakrishnan 13983 (NY). CHINA. Prov. H i m h h Hi khov Hunan Sheng, 

XinningXian Ziyun Inn \ 1 c4 | H j h 1885-88, Henry 29(97 

(GH);Prov. Kiangsi lini ir i h ii lnn\i(it\ 1 >u,| I - , „ I , i | , u v Sikanq 1935 }r> 

II inj(t44 (i. H) I in i I I mil i n \u i 1 I i I i I uji hi j\ in Lot Longxi, 

p 1 98£ / ' ' v?4t (MO NY);Kwnicl i /( NY), N 

Prov.Chong Uen Shan ihmi i i j^oll >< -,",., Jucloux 252 (NY); 

Hong Kong, Lantao Island, Tungchung,Taishui-hang, 29 Sep 1940, Taam 1819 (NY). TAIWAN. 
Niitagagun, Hosya, Kusunokizinzya, 14 Oct 1 935, S. Suzuki s.n. (MO); Little Quemoy, 16 Sep 1961, 
Chuang 4469 (GH);Botel Tobago, 1 6 Ai < . \t ) JAPAN. Hondo, Yamanakako in Kai, 

3 Aug 1 952, K. Okamoto NSM 584 (BRIT, NY); Prov. Tootoomi, Pref. Shidzuoka, Hondo, 30 Sep 1 959, 
MiyoshiFuruses.n. (GH),11 Oct 1 960, M/yo b • iru i ' (GH) Prov. Kadzusa,Pref.Chiba, Hondo, 5 Sep 
]962,MiyoshiFuruses.n. (GH);Prov.Yam,v,hiio: Ml. I lin/an r Shirakawamichi,9 Sep ]93],Tagawa617 
(NY);Pref.Mie,Ogurusu, Kiwa-cho,Minamimuro-gun, 17 Nov ]977,Mimoro,Tsugaru & Deguchi 4289 
(MO); Pref. Settsu, Ashiya, 1 Sep 1 954, Hutoh 1 1517 (NY). 

SOUTHEASTERN ASIA. VIETNAM.Tonkin,Chapa,Pefe/o? 6099 (NY).THAILAND.Phetchabun, 
Phu Miang, 2.1 0.1 967, Schimizu et al.T.11391 (NY); Chiang Mai, Doi Intanoid, 1 3 Sep 1 974, Larsen & 
Larsen 34512 (NY); Maehongson, Khun Yuam, 7 Sep 1 974, Larsen & Larsen 34254 (NY). INDONESIA. 
Lesser Sunda Islands, Flores, near Keli Moetoe, Kampong Ndoearia, 8-6-1938, Jaag 180 1 (GH); Java, 
Mt Gedeh.Tjibodas, 30 Apr 1950, Kern 7998 (GH). PHILIPPINES. Northern Luzon, Prov. Benguet, 
Trinidad 28 Sep 1904 1' - ' > I Nt 

AUSTRALIA. QUEENSLAND. Moreton District, Brisbane, 5 Aug 1 937, Blake 12965 (GH). 

AFRICA. Eritrea. Ocule Cusai, 1 6 Sep 1 902, Pappi 1799 (NY). 

We gratefully acknowledge the following individuals and agencies:Cary Norquist,United 
States Fish & Wildlife Service, for providing financial support for the first author's field 
work in Louisiana and Mississippi during 1 993; Nelwyn Gilmore, Louisiana Natural Heri- 
tage Program, for providing data and specimens she collected at the holotype locality; 
Georgia Natural Heritage Program for funding the first author's field work during 1 996 at 
Kings Bay Submarine Base, Camden County,Georgia;Ron Wilkerson for assistance at Kings 
Bay Submarine Base and providing information on road maintenance practices there; 
J.R. Burkhalter and Randy L. Mears for sending Alabama specimens to the first author for 
determination; John R. MacDonald, Don Sudbrink, and Shaharra Usnick for assistance 
with field collections; Serial Kenerson for providing rice production statistics for Hancock 
County,Mississippi;RussGoddardforassistance with and lending of photomicrographic 
equipment; Gordon C.Tucker for providing helpful comments on the manuscript; and 
the following herbaria for lending specimens without which completion of this project 
would not have been possible: BRIT, GH, MO, NY and US. 

Adams, CD. 1994. Cyperaceae. In: G. Davidse, M.S. Sousa and A.O. Chater, eds. Flora 

Mesoamericana,Vol. 6: Alismataceae a Cyperaceae. Universidad Nac 

de Mexico, Cuidad Universitaria.Pp. 262-485 
Anonymous. 1959. United States Census of Agriculture, Vol. 1, Part 33, Mississippi. U 

States Department of Agriculture. 
Anonymous. 1 982. United States Census of Agriculture, Part 24, Mississippi. United S 

Department of Agriculture. 
Anonymous. 1 993. Endangered and threatened wildlife and plants; review of plant ta; 

listing as endangered or threatened species; Department of Interior, Fish and Wi 

Service, 50 CFR Part 1 7. Federal Register 58 (1 88): 5 1 1 44-51 1 90. 

A i,r, i 1'JQo Minitab reference manual: release 1 1 for Windows™ Minitab lnc.,5tate 

i oilrcjo, PA. 
Bruhl, J. 1 995. Sedge genera of the world: relationships and a new classification of the 

Cyperaceae. Aust. Syst. Bot. 8:1 25-305. 
Bryson, C.T. and R. Carter. 1 992. Notes c 

with records of six species new to t 
Bryson, C.T. and R. Carter. 1 994. Notes c 

sissippi with records of eight specie 

Bryson, C.T., R. Carter, LB. McCarty, and F.H.Yelvert >n. 199 .Kyllinga,a genus of neglected 

weeds in the continental United States. Weed Technology 1 1 :838-842. 
Bryson, C.T, J. R. Ma [) m-.i,,R L'hii .ind Mj W. 1996 Noteworthy Carex, Cyperus, 

I leo( haris, Kyllinga, and Oxycaryum (Cyperaceae) from Alabama, Arkansas, Georgia, 

Louisiana, Mississippi, North Carolina, Tennessee, and Texas. Sida 17:501-518. 
Carter, R. 1 990.Cyperus entrerianus (Cyperaceae),an overlooked species in temperate North 

Carter, R. and C.T. Bryson. 1 996. Cyperus entrerianus: A little known aggressive sedge in the 

southeastern United States. Weed Technology 10:232-235. 
Carter, R„ R.L. Mears, K.C. Burks, and C.T. Bryson. 1996. A report of four exotic Cyperus 

(Cyperaceae) species new to Florida, U.S.A. Sida 1 7:275-281 . 
Clarke, C.B. 1 894. Cyperaceae. In: J.D. Hooker, ed.The flora of British India, Vol. VI, Part XIX. L. 

Reeve & Co., London. Pp. 585-672. 
Ci arki , C.B. 1 908. New genera and species of Cyperaceae. Kew Bull. Add. Ser. 8:1 -1 96. 
Corcoran, Ml. 1 941 .A revision of the subgenus Pycreus in North and South America. Catho- 


r. Biol. Ser. 37. 

CiOl l<..,HIBIUR,P. V 

986. Genera Cyperacearum:een 



?n fylogenese van de Cyperacea 




1989. Studies in Cyperaceae 9 

'. Problems in t 


taxonomy of Cyperus L. Bull. Soc 


■ lectotypification 
Soc. Roy. Bot. Belg. 122:103-114. 
Haines, R.W. and K.A. Lye. 1 983.The sedges and rushes of east Africa. East African Nat 

History Society, Nairobi. 
Hoi m, L.G., J.V. Pancho, J. P. Herberger, and Dl. Pi ucknett. 1 991 . A geographical atlas of w 

weeds. Krieger Publishing Company, Malabar, Florida. 
KfrnJ.H. 1974. Cyperaceae. In: C.G.G.J. van Steenis, od. I Iota Malesiana, Series I, Vol. 

NoordhoffLeyden. Pp. 435-753. 
KoyamaJ. 1985. Cyperaceae. In: M.D.Dassanayake, and F.R.Fosberg,eds. Flora of Ceylon 

S. Amerind Publishing Co. Pvt. Ltd., New Delhi. Pp. 125-405. 
Krai , R. 1 971 .A treatment of Abildgaardia.Bulbostylis and Fimbristyli 

Ki ii in, II I 982. Cyperaceae. In: Hafliger, E„ et al., Monocot Weeds 3. 

KukenthauG. 1935-1 936. Cyperus. In: A. Engler and LDiel,eds. Das Pflanzenreich IV.20 (Heft 
101). Pp. 1-671 . 

Kukkonen, 1. 1 995. Two proposals to conserve species names in Cyperaceae.Taxon 44:625- 

Mayr, E. 1 942.Systematics and the origin of species. Columbia University Press, New York. 
McKenzie, P.M., B. Jacobs, CT.Bryson, G.C.Tucker, and R.Carter. 1998. Cyperus fuscus (Cyperaceae), 

new to Missouri and Nevada, with comments on its occurrence in North America. 

Mingyuan, X. and M. Dehu. 1 970. Farmland weeds in China: a collection of coloured illus- 
trated plates. Agricultural Publishing House, Beijing, China. 

Ohwi, J. 1965. Flora of Japan. Smithsonian Institution, Washington, D.C. 

Reed, C.F. 1977. Economically important foreign weeds. Agriculture Handbook No. 498. 
United States Department of Agriculture, Washington, D.C. 

Stearn,WT. 1 992. Botanical Latin. 4th Ed. David and Charles Publishers. Brunei House. New- 
ton Abbot, Devon, England. 

ThieretJ.W. 1977. Cyperus louisianensis (Cyperaceae), a new species from southern Louisi- 
ana. Proc. Louisiana Acad. Sci. 40:23-26. 

Tucker, G.C. 1 983. The taxonomy of Cyperus (Cyperaceae) in Costa Rica and Panama. Syst. 
Bot. Monog.2:l-85. 

Tucker, G.C. 1 987.The genera of Cyperaceae in the southeastern United States. J. Arnold 
Arbor. 68:361 -445. 

Tucker,G.C. 1 994. Revision of the Mexican species of Cyperus (Cyperaceae). Syst. Bot.Monog. 

en, ed. Flora of New South Wales, Vol. 4. New 


PEibR Thomas. 2000. Trees: Their Natural History. (ISBN 0-521-45963-X,pbk.) Cambridge 
University Press, 1 1 Midland Avenue, Port Chester, NY 1 0573-4930, U.S.A. $24.95 
pbk., 286 pp. Line drawings. 

From the first senten. i large woody thing that 

both a sense of humor and general enthusiasm about trees. Peter I hnma\ lecturer in environmen- 
tal ion. fsat Keel Hm r n Ml in i i -oh n i mnti n mi through myriads of jour- 
nals and books from all i I r M til f Nht tk 
vital to healthy ecosystems, but also unparalleled in the range of materials they provide for human 

in < i ti It qui linn ) I lov io I < wot I I low an ihe\ i i im I Vnd lm tin th |to\ 
and reproduce? And in I I I t t i 

the natiiial histoivol tiees coverinq then biology and ecology. 

Beginning with a preface and an overview ol trees then I >lk> en h> eight chapters that 
more specifically address the parts of trees and how they live and die. Chapter Two covers the 
leaves, Chapter Three the mink and blanches, Ch.iptoi I our the roots, Chapter Five flowers, fruits, 
and seeds, Chapter Six," I he growing ttee/'t haptet Seven, l he sh<i|.eol I tees; Chapter Eight/New 

treesfromold, "and Chapter Nine,"Health, damage at d ie nh II C then concluded bya list 

of further reading and mm I Hi m hit ph t imp m i black and white line 

drawings are abundant tin uh n mi i iipiqutt. i ih ql i inq feature is the list of 

further reading that follows each chapter. These lists are chapter specific and contain books and 
journal references no! listed in the lisi ol ititthet teadinq at the end of the book. 

The author suite. I n, hi „ to dm u ,, thm tiand lint mi ition to create a readable 
book that would answer common guestions about trees,set right a number of myths and open up 
the remarkable world of how trees work, g row, repro b eand dii Vhil ntren for lay audiences, 
the book is substantially scientific and the references at the end ol eat h chapter offer the reader 
more specific academi I v u| in unit n i il i n \ tiki I 

questions they've vvoncieted about j ot vears — he\ in I i ir<;vv, Botom, ;i Bnseorch Institute of Texas, 




Richard Carter 

1 ' 'og> Department 

Valdosta State University 

V,ikio\Ui.(.r\ lir.*S-00i.\U.\A. 

i previously reported fi 

i Dade County, Florid 


e Cyperus en los Estados I 

In late 1999, the second author discovered a population of an unknown Cyperus species 
in Dade County, Florida. Specimens were sent to the first author for determination and 
were identified by him as Chyalinus ihlj • a nus has a wide paleotropical dis- 

tribution, ranging from eastern Africa, Madagascar, Mauritius, India, Sri Lanka, tropical 
Australia (Queensland),and Malaysia (Kukenthal 1 935-1 936; Kern 1 974; Haines & Lye 1 983; 
Koyama 1985). This remarkable little sedge has not been previously reported from the 
Western Hemisphere. 

The taxonomic relationships of Chyalinus are obscure,and its nomenclature is com- 
plex. In addition to Cyperus,the species has been treated in the segregate genera Pycreus, 
Kyllinga, and has also been placed in various subgenera of Cyperus, i.e., 
subg. Kyllinga (Kern 1974), subg. Afar/sa/s (Kukenthal 1935-1 936), subg. Pycreus (Clarke 
1 884), and subg. Queenslandiella (Govindarajalu 1 975; Haines & Lye 1 983). Its lenticular 
achene, bifid style, compressed spikelets with multiple flowers and fruits, and open 
anthelate inflorescence suggest a relationship with subgenus Pycreus. However, persis- 
tent scales and disarticulating spikelets defy placement there and indicate an affinity 
with Kyllinga or Mariscus. Because its treatment as a Pycreus, Kyllinga, or Mariscus is prob- 

i segregate it in the monotyf>ic o. t , i < <■ ■ ■ h '.7, however, con- 
uire the segregation of other genera from Cyperus, which would up- 
b. Based upon a studs is it h nt . mdarajalu (1975) placed 
C. hyalinus in monotypic subg. Queenstandiella of Cyperus. Until there is uneguivocal 
molecular evidence to the contuo j tkmf il nlk n tin i >perus similar to the 
concept of Haine mil i 1 i i i u< n i inq current nomenclature. Thus, 

we neat this spot ies in ( i/pcoo subg. Oueenslandiella. 

Our objectives herein are to report C. hyalinus new to Florida, U.S.A., and the West- 
ern Hemisphere and to provide 1 a dichotomous key, technical description, notes, and 
photographs to facilitate its identification in the United States. 

■"' 'If 'f.l. 



Spikelet rachilla remaining attached to rachis.not bas, 

ally articulated; floral scales 

and achenes disarticu ating from base to apex of rachi 

.'. o ..•[•■-. iiu no, k... he-es trigonous. 

i. Spi<elels various \ a'uncjed.but not in diqitatoc lu 

v Spikelets in digitate < !usters;plants of hydric to me 

?sic habitats;kranzanatomy 

absent. subg. Anosporum C.B 

( larke | subg /V nosier T ; e B lail i | 

2. Style branches 2;achenes lenticular (rarely turgid ar 

C Spikeleis laterally ; ompressed;achene angle adjacent 

to rachilla. subo.Pycreus 

lilla. subg.Juncellus(Ghseb) 


Spikelet, floral scale, and achene articulation not as abc 

5. Style branches 3;ac ;h-iu -, trigonous.-spikelet basally 

as ,i unit Willi floral scales and achenes siill attac hex 

J or spikelet breaking apart 

tiansversely into 1 -fruited segments. 

o Spikelet l)asallvaitieulated, deciduous as a unit wi 

stiil attac hi'tl to rachilla. 

6. Spikelet breaking apart transversely into 1-fruitec 

I segments. subg. Diclidium 

B.Clarke [=subg Torulin, n De , Kiik.J 

5. Style branches 2; achenes lenticular to plano-comt 

tieulated, cieciciuous as a unit with floral scales anc 


3-compressed. subg. Queenstandiella (Domin) Govind. 

Cyperus subg. Queenslandiella (Domin) Govind., Reinwardtia 9:1 94. 1 c 
Inflorescence an open antheltt of m tiv pt lunculate spikes. Spikelets 

Hoe ei and u hi m I i il ul >t I t n in i int ic 1 II >i : il> u i . 

(chlorocyperoid) anatomy. Subgenus monotypic. 


Bot,85:41 6. 1 91 5.Mariscopsis suaveolens Cherm. Bull. Mus. Hist. Nat. (Paris) 25:60. 1 919. Pycreus 

hya//nus(Vahl)Domin,Biblioth.Bot.85:417.1915.M<// >pvs/i ilmus* iiil) F Ballard, Bull. Misc. 

Inform. Kew 9:458.1 932. Queenslandiella hyalina (Vahl) F.Ballard in Hook. Icon. PI. 33:t. 3208. 

1 933. Kyllinga hyalina (Vahl) T. Koyama, J. Jap. Bot. 51:313.1 976. 
Loosely cespitose aromatic annual herb. Roots fibrous, brown. Stems glabrous, trigonous, 
3-14 cm x 1-2 mm. Leaves 3-7, basal; bases sheathing; blades 4-15 cm x 2-5 mm. Pri- 
mary inflorescence bracts 4-8, mostly exceeding rays, longest to 12 cm long, 2-4 mm 
wide. Inflorescence anthelate;rays 3-8, longest 2.5-4 cm long; spikes simple (rarely with 
short basal branch), mostly pedunculate, oblong-ovate, (7-)1 2-20 mm x 8-15 mm, with 
(5-) 1 2-1 7 mostly divaricate spikelets; rachis grooved, winged. Bracteoles narrowly trian- 
gular to aristate, 0.4-2.2 mm long, membranous. Spikelet prophylls rounded to acute, 
0.7-1.4 mm long, membranous. Spikelets laterally compressed, narrowly ovate toelliptic, 
4.1-5.7 x 1.9-2.2 mm, deciduous; rachilla wing ca. 0.5 mm wide, membranous. Floral 
scales 4-7 [3-4 fertile], imbricate, broadly ovate, 2.1-2.4 mm long, mucronate, membra- 
nous; keel green, scabrid; wings yellowish to whitish to pale green; lateral nerves 6-8. 
Stamens 2; anthers narrowly oblong, 0.4-0.5 mm long. Style bifid, divided ca. 3/4 of its 
length. Achene brown, broadly oblong to suborbicular, 1 .0-1 .4 x 1 .0-1 .1 mm, piano- 
compressed, 0.3 mm thick,gibbous, truncate-retuse, minutely puncticulate. Fig. 1. 
Phenology— In the United States, flowering late July through November. 
Distribution— \n the Old World, ranging from eastern Africa, Madagascar, Mauritius, 
India, Sri Lanka, tropical Australia (Queensland),and Malaysia (Kukenthal ibid.; Kern 1974; 
Haines&Lye 1983; Koyama 1985). Herein reported new to the Western Hemisphere, where 
so far it is restricted to southern Florida, U.S.A. Fig. 2. 

Voucher specimens. U.S.A. FLORIDA. Dade Co.: E side of Miami International Airport, just N of 
Perimeter Rd, road shoulder, sandy soil, 26 Oct 1 999, Randy Mears s.n. (EIU, FLAS, MICH, MO, US, USE, 

Cyperus hyalinus (Fit] I 11 readil\ listinguished from all other congeners by the follow- 
ing combination of characteristics: broadly oblong, truncate-retuse, piano-compressed 
achene; bifid style; 3-4-flowered, deciduous, flattened spikelets; membranous, yellowish 
to pale greenish, 6-nerved, mucronate floral scales with scabrid keel;and open anthelate 
inflorescence. Morever, dried specimens exhibit the distinctive odor of fenugreek 
{Trigonella foenum-graecum L), previously noted by various authors (e.g., Kern 1974; 
Govindarajalu 1975; Bruhl 1995) and also characteristic of Cfuscus L, C. setigerus Torn & 
Hook, and C sguarrosus L. (see McKenzie et al. 1 998). 

most recent in a series of exotic Cyperus spp. reported new to the United States (Carter 

■ .^3 X 


1990; Carter et al. 1996; Carter & Bryson 2000).The broad dispersal of such species is not 
surprising given their weedy nature, their copious production of small fruits, and the 
current ease and frequency of rapid, long-distance transportation of humans and cargo. 
Its rarity and proximity to the Miami International Airport suggest a recent introduction 
of C.hyalinus via shipment of air-freight. 

In addition to the original population growing along an open, sandy road shoulder, 
the second author has discovered another one about one-half mile away along a rail- 

Bidensalba (L.) DC. ,Cenchrus incertus M.A.Curtis, Chamaesyce hirta (L.) M\\\sp.,C.hyssopifolia 
(L) Small, C.maculata (L.) Small, Dactyloctenium aegyptium (L.) Willd.ex Asch.&Schweinf, 
Polypremum procumbens L,Setaria parviflora (Poir.) Kerguelen,S/c/oe///off/7Torr.& A.Gray, 
and Tridax procumbens L. 

The occurrence of C. hyalinus in Australia, Madagascar, Mauritius, and Zanzibar is 
sporadic (Ballard 1932, 1933), and it does not appear to exhibit aggressive or invasive 
properties in southern Florida. Thus, currently C.hyalinus would not seem to threaten 
native biota in the United States, and its tropical distribution in the Old World suggests 
establishment is unlikely in more temperate regions of North America. However, it should 
be monitored and additional populations sought in southern Florida, especially in light 
of its description as "a weed of sandy soils, near sea level" in eastern Africa (Haines & Lye 

Ballard, F. 1932.LXIII.~The genus Mariscopsis. Bull. Misc. Inform. Kew 9:457-458. 
Ballard, F. 1 933. Queenslandiella hyalina (Vahl) Ballard. Hooker's Icon. PI. 33:t. 3208. 
Bruhl, J. 1 995. Sedge genera of the world: relationships and a new classification of the 

Cyperaceae. Aust. Syst. Bot. 8:1 25-305. 
Carter, R. 1 990.Cyperus entrerionus (Cyperaceae),an overlooked species in temperate North 

America. Sida 69-77. 

Sida 19:325-343. 

(Cyperaceae) species new to Florida, U.S.A. Sida 17:275-281. 
Clarke, C.B. 1884. On the Indian species of Cyperus.J. Linn. Soc, Bot. 21:1 -202. 
Govindarajalu, E. 1975. Studies in Cyperaceae. XIV. Endomorphic evidences for placing 

Cyperus hyalinus under the new subgenus Queenslandiella. Reinwardtia 9:1 87-1 95. 
Haines, R.W. and K.A. Lye. 1 983. The sedges and rushes of east Africa. East African Natural 

History Society, Nairobi. 
Kern, J. H. 1974. Cyperaceae 1. In: C.G.G.J. van Steenis,ed. Flora Malesiana,Vol.7.Noordhoff, 

Leyden. Pp. 435-753. 
Koyama,T. 1985. Cyperaceae. In: M.D.Dassanayake and F.R.Fosberg,eds. Flora of Ceylon, Vol. 

5. Amerind Publishing Co. Pvt. Ltd., New Delhi. Pp. 1 25-405. 

'<:'::> I ' -. i ' , It M<il i utL [ 'i. i . I ! i I 1 in enreich IV.20 (Heft 

101). Pp. 1-671. 
It KtN/ih, P.M., B. Jacobs, C.T.Bm n,G.( I m n,andR.C n i 1 998. Cyperus fuscus (Cyperaceae), 
new to Missouri and Nevada, with comments on its occurrence in North America. 
Sida 18:325-333. 


A. Eduardo Estrada C. and Alfonso Martinez M. 



nil- I mi n mini ih- c entf il fj.iti I ih i 1 n Oi hu 

" in )i K^fe ^ 1 .I i ndeae7/12.0ak 
f i nt j ',i with the highest numbei t i 

i i i I =i r i J i ,)olea,Desmodiu 

:ion central del estado de Chihuahua registro 42 gem 

e generos/numero de especies p 
Jeae 8/28; Caesalpinioideae 7/1 ; 
3c/a (11), Phaseolus (7), Desmodiu 


tituted by three subfamilies of plants Mimosoideae, Caesalpinioideae 
i they are the second most diversified group of plants in Mexico after 
2 (Sousa &Delgado 1993). Legumes are found in all plant communities of the 
country, and are one of the dominant groups of plants in north Mexico.They are found 
eon and Chihuahua (pers.obs.). 
:ated at 28° 1 5-29° 05' N, 1 05° 07'- 1 07° 35' W, 
d includes the eight municipios,Aldama,Aquiles Serdan, Chihuahua, Coyame,General 
as, Julimes, Meoqui and Riva Palacio on a surface of 7500 km 2 (Fig. 1 ).The study area 
:ludes two physiographic provinces (Anonimo 1987), the eastern part of the Sierra 
idre Occidental and Sierras y Llanurasdel Norte, which has two subprovinces,Bolson 
Mapimi and the Sierras Plegadas del Norte. Most rocks are sedimentary or volcanic 
gin from the Quaternary Period. Most mountains in the area are acidic intrusive igne- 
s rocks from Tertiary Period and metamorphic rocks (limestone) outcrops from the 
?sozoic Era, and from the Lower Cretaceous Period (Anonimo 1981). The three main 
mate types for the area are: (1 ) very dry semi-warm, characterized by summer rains, 
2 winter rainfall less that 5% of total, the winter cool— characteristic of western part of 


J-S-oriented 30-40 km wide strip in c 

i:M) 1 

of total, summer warm— found on high plains and small mountains ranges at 1 600- 
2400 m elev. These climatic types are mapped as BWhw(w), BSokw(w) and BS,kw, re- 
■ |)t i HvH\ in Garcia (1973). 

There are three main vegetative communities,grasslands,shrublands and oak-pine 
forest (Rzedowski 1978). Bouteloua, Sporobolus, and Hilaria are the dominant genera in 
the grassland landscape, Bouteloua gracilis Lag, B. curtipendula (Michx.) Tom, B. eriopoda 
(Torr.) Tom, B. hirsuta Lag., Sporobolus airoides (Torr.) Tom, Hilaria mutica (Buckl.) Benth, 
tragrostis intermedia Hitchc, E.lehmaniana Nees, Enneapogon desvauxii Beauv, and Lycurus 
phleoiodes H.B.K,. are the dominant grassland species found. Shrubland communities 
have mostly low elements,seldom over 2.5 m tall, forming subthom shrublands (Anonimo 
1981), where Larrea tridentata (Sess. & Moc. ex DC.) Cav, Flourensia cernua DC, Acacia 
neovernicosa Isely, A. constricta Benth., A. schaffneri (Wats.) Herm., Lycium berlandieri Dun, 
i spinosa Zucc, Condalia ericoides (A. Gray) M.C Johnsl, Yucca elata (Engelm.) 
i spp., Mimosa spp.,CeltispallidaJorr.,Parthenium argentatum Gray and P. 

incanum Kunth are the predominant species (Anonimo 1978). Oak-Pine forests are lo- 
cated in mountain areas, north of Cd. Chihuahua, especially Sierra El Nido and Parque 
Nacional Cumbres de Majalca.and 30-40 km west of Cd. Chihuahua in direction to Cd. 
Cuauhtemoc. Quercus-Juniperus and Pinus-Quercus associations are present; most com- 
mon oak species are QuercuschihuahuensisJre\.,Q.grisea Liebm.and Q.emory/Torr.,and 
in the lower hills and plains, where the woodlands contact the grasslands, Juniperus 
monosperma (Engelm.) more frequent, often forming savanaah. Higher and cooler 
areas of the mountains have Quercus hypoleucoides A. Camus, Q.arizonica Sarg., Q.rugosa 
Nee, 0. grisea Liebm., Q. depressipes Trek, Pinus cembroides Zucc, P. engelmanii Can., P. 
chihuohuana Engelm. and Juniperus deppeana Steud. Southeast of Cd. Chihuahua Prosopis 
glandulosa Torr. var. torreyana (L Benson) I.M. Johnst., forms dense shrubby areas 
(mezquital), especially along roadsides and disturbed areas from Cd. Chihuahua to 
Estacion Horcasitas, 25 km SW form Cd. Chihuahua (Anonimo 1 981 ). Pressed and dried 
vouchers are stored in the herbarium CFNk; incomplete sets are deposited at ANSM, 
BRIT, MEXU, NMC and TEX (Holmgren & Holmgren 1 990). 

During 1 994-1 998 numerous routes through all plant communities of the central part 
of the state of Chihuahua were sampled. On each site where legumes were collected, 
main vegetation type, geographic coordinates, nearest towns, altitude and slope were 
recorded. Nine hundred and fifty samples of legumes were collected in this study. 

I I 

Only a partial geographical representation of legumes exist for Mexico. Regional studies 
have in particular contributed to the knowledge of these plants. Of particular merit are, 
for southern Mexico,Standley and Steyermark (1 946), and Woodson et al.(1 980);for cen- 
tral, Matuda (1 980),and Rzedowskt and Rzedowski (1 979);for southwest, McVaugh (1 987); 
for northeastern, Correll and Johnston (1 970), Isely (1981), Estrada and Marroquin (1991), 
Carranza and Villarreal (1 997), and Ramos (1 999);for northwestern, Munz (1 959), Kearney 
and Peebles (1 960), Shreve and Wiggins (1964), Isely (1 981 ),Spellenberg et al. (1996), and 
Estrada et al. (1 997).These studies show distribution, diversityand ecology of legumes in 
this country, while this research focuses on additional knowledge on presence,distribu- 
tion and ecological aspects of the legumes in northern Mexico. 

In this study, 42 genera, 1 17 species and 36 varieties of legumes from the central 
part of the state of Chihuahua were recorded. Lotoideae has the highest number of 
genera and species with 27 and 77 respectively, followed by Mimosoideae with 8 and 28 
and Caesalpinioideae,with 7 and 12. Appendix 1 shows the taxa by family, and vegeta- 
tion type(s) where they were more frequently found. Dalea, Acacia, Desmodium and 
Phaseolus are the most diverse genera, each with 5 or more species. These genera have 
species in the three main plant communities. Dalea is the genus with more species (1 9), 
nine of which occur in pine-oak forest, eight are dominant in grasslands, while six are 

under 1 700 m elevation, except for A. ana/im " ; t'nt a . i mountainous areas 

dominant species in matorral communities. Five out of seven Phaseolus species are ex- 
clusive to pine oak-forest in the cooler parts of the region (Sierra El Nido and Majalca), 
above 1800 m elevation. All of Desmodi vm species occur in oak or pine forest, only D. 
neomexicanum occurs in the three dominant communities. None of the Astragalus spe- 
cies occurs in forest areas, while all of them occur in grassland communities. No species 
of Caesalpinoideae occur in pine or oak forest, the five species of Senna were seldom 
found in low numbers in matorral and grassland. Most /Vf/mosa species were often asso- 
ciated with oak forests, in particular those formed by Quercus emoryi and Q. grisea, and 
matorral. In both communities, M.aculeaticarpa is the most abundant. 

Highest number of legume species were recorded on oak-pine forests (33) and 
Bouteloua grasslands (31). Oak Forest and Pine Forest, each have a similar number of 
legume species,21 and 20 respectively. In shrublands where Acacia is dominant,25 spe- 
cies of legumes were recorded, three times the number of species recorded in Larrea 
and Flourensia shrubland.Within different grassland association,the Sporobolus and Hilana 
types are areas with lowest legumes, with 9 and 6 species respectively, although they 
shelter some character is! it species Irom these plant com m unities such as Hoffmanseggia 
spp., Astragalus spp.and Peteria scoparia. Several species are found mainly in disturbed 
and overgrazed areas, they are Acacia farnesiana, Mimosa aculeaticarpa var. biuncifera, 
Prosopis glandulosa var. torreyana, Acacia schaffneri var. bravoensis, Crotalana pumila, 

^ central plains, especially in the area between ( hihua 

in abandoned agricultural lands,clos 

Eight legume species (7.5%) are introduced , i> , , , ■■ > .•>,,,,„ ( , u m 

Albi/ia ji till >m s/7 1, Media igo saliva; Medicago lupulina, Lablab purpureas, Robinia pseudoacacia, 
and Wisteria sineneis.the latter three are cultivated as ornamental plants. Eighty eight species 
are native of northern Mexico, some of them occur in southern U.S. From the approximately 
1 35 genera and 1 724 legume species present in Mexico (Sousa & Delgado 1 993), 3 1 .1% and 
7.1 % of them respectively are present in the study area. Grasslands and shrublands cover 
95% of the studied area, but they have lower legume diversity than oak-pine forest, which 
represent only 5% of the surface. Total taxa recorded are distributed in a 7500 km 2 surface 
(lower than a tenth part of the state territory) on three main vegetal communities from the 
seven recognized forthestate of Chihuahua iAnommo i978).A very different and heteroge- 
neous relief altitude, climate and vegeta 
Chihuahua, reveal a constant legume spec 

oak, pine, oak-pine forests and subtropical shrublands. Spellenberg et al. (1 996) list 30 
genera and 65 legume species from Parque Nacional Cascada de Basaseachi; Estrada et 
al. (1997) list 21 genera and 53 species in Babicora Lake.Table 1 shows the affinities of 

presence of 14 genera in central Chihuhahua, absent from Basaseachi and Babicora 
{Desmanthus, Peteria, Painteria, Albizia, Prosopis, Zopoteca, Hoffmanseggia, Pomaria, 
Parkinsonia,Lablab,Melilotus,Nissolia,Pediomelum,Robinia, and Wisteria), and seven gen- 
era in Basaseachi, absent from Babicora and Central Chihuahua, these are Conzattio, 
Pithecellobium, Lysiloma, typical of moist warm environments and Erythrina, Lathyrus, 
Manna and Minkelersia. In Babicora no Caesalpinioideae occurs, while in Basaseachi, three 
species werefound,all of which had tropical affinity.Da/eo is conspicuously more abun- 
dant in the center of the state than in the other two areas, fourteen of the central species 
are absent in Babicora and Basaseachi. All of these are from semiarid climates. Desmodium 
is more diverse in Basaseachi, seven of its species, that are absent in the other two areas, 
are more common in moist environments. Perhaps the greater species diversity of le- 

Genera with more 
Acacia species 
Desmodium species 

Subfamilies, genera and species of legumes are recorded for the central part of the state 
of Chihuahua and plant communities where most freguently found. HG, Hilaria Grass- 
land; SG, Sporobolus grassland; BG, Bouteloua grassland; SL, Shrubland {Larrea as domi- 
nant); SF, Shrubland (Flourensia as dominant); SA, Shrubland (Acacia as dominant); SP, 
Shrubland {Parthenium as dominant);OF,Oakforest;OPF, Oak-Pine forest; PF, Pine Forest; 
D, Disturbed Areas; C, Cultivated. 




?r/ Benth. [SA] 


?nm.) Irwin & Barneby [BG, 

x>k.) Benth. [SG,C] 

Acacia wrightii Benth. [SA] 

Albiziajulibrissin Durazz. [C] 

>> ift-vfoGray [SA,S 

dllianJia runphvlla [iriith. va 

Barneby [SL,SF,BG] 
nna wislizeni (Gray) lrw!n 
wislizeni [SL,SF,SA] 

1 iih n num///s [OF, OPF] 

'",,.,- ,n ,r , , i n- 

Desmanthus cooleyi (Eat.) Trel. [BG, SA, SL, OF] 


' i BG,SA,SF] 

- .',' Ml ,1 ' i ' ' , I.MMli 

Rvdb.) <]( si [SG,HG,BG] 

Barneby [D,SA, OF] 

1 -no aJ\ <u, ,r t , ( , Benth [BG, OF, OPF] 

Barneby [BG.SG] 

' " '-'. 'ii i I'-'iiMi ,n - > „jp , ," /ii; \ 

\sti m,j /[( , \.[) it , , a n i | .mail) 


Barneby [BG,HG,SA] 

moryona [OF, SA] 

Astragalus (tntnjIaWats. [BG, D, SA] 

' ' ^ ," ,' > it n vjnj/e, [sa, OF] 

Astragalus guinqueflorus S.Wats. [BG, HG] 

r n ,h ii fe\ana[SA,SF] 

Astragalus wootonu Sheld. [D, BG, SA, SF] 

Cologania angustifolla H.B.K. [OPF, OPF] 


Cologania obovata Schlecht. [PF, PF] 

• • < < t <• , 1 '1 nton&Rose[BG,OPF] 

uist , ,' > , ] i in . it '-,n >m\ 

Prosopis glandulosalorr. var. torreyana (L.Benson) 

Coursetia glabt 

Dalea foliolosa (Ait.) Barneby var. foliolosa [01 

Dalea formosa Torr. [SL, SA, SF, SP] 

Dalea gray! (Vail) L.O Williams [OPF, OF, PF] 

Dalea humilis G.Don.[Pf,OPF\ 

Dalea jamesii (Torrey) T. & G. [BG] 

Dalea lachnostachya Gray [SL, SA] 

Dalea leponna (Alton) Bullock [BG,OPF,OF] 

Dalea I u tea (Cav.)\ 
Dalea nana Torr. va 



/illci. var. /urea [SA, OPF] 
(Gray) Cory var. neomexicana 
pogonathera [BG, 

SA, SF, SP, SG, HG] 

Dalea prostrata Ortega [BG] 

Dalea versicolor Zucc. var. glabrescens (Rydb.) 

Barneby [OPF, PF] 
Dalea versicolor Zucc. var.sess/7/s (Rydb.) Barneby 

Dalea viridiflora Wats. [OPF] 
Dalea wrightii Gray [BG] 
Desmodium angustifolium DC. [OF] 
Desmodium batocaulon Gray [OPF,PF] 
Desmodium hartwegianum Hemsl. var. 

hartwegianum [PF, OPF] 
, -., „ n t - v, , ^GrayfS^OFBG] 
Desmodium grahamii Gray [OF, BG] 
Desmodium retinens Schlecht. [OPF] 

Peebles [OF, OPF] 
'tusplebeius (Brandeg.) Barneby [PF] 
yj/nt/s aff. delicatulus Sprague & Riley [PF] 
•pinus ehrenbergii Schlecht. [OF, OPF] 

Melilotus indicus (L) All. [D] 
Melilotus officinalis L. [D] 
Nissoiiapnnglei Rose [SF,SA,SL,D] 
A//550//Q w/s/;zen/ (Gray) Gray [SF, BG] 
Pediomelum palmen (Ock.) Grimes [SG, BG] 
Peteria scoparia Gray [BG, SG, HG] 
Phaseolus acutifolius Gray [SA, OF] 
Phaseolus angustissimus Gray [BG] 
Phaseolus grayonus Woot.& Standi. [OF,OPF,PF] 
Phaseolus leptostachyus Benth. [OF, OPF, PF] 
Pha5eo/u5 porw/us Greene [OF, OPF, PF] 
Phaseolus ritensis Jones [OPF, OF] 

Rhynchosia macrocarpa E 

Rhynchosia senna Gil! ex 


i [SA,OF,OPF] 

-pureus (L.) Sweet [C] 

We thank Alfonso Delgado a 
Richard Spellenberg, Emily Lc 
English translation and critic, 


phrosia '< nclla Gray [SA] 
/fo//um '»u/)//e H B K var.amabile [OPF, PF] 

(Greene) Barneby [OPF, PF] 

c/o ludoviciana Nutt. ssp. ludoviciana [OPF, PF] 

I i <h L m i / ' IF ;//, he/to [OPF, PF] 

Zom/o gemella (Willd.) Vog. [SA, OF, OPF] 
Zorn/a reticulata Sm. [OF, OPF] 

d Fortunato Garza for comments on the manuscript and 
t, Barney Lipscomb and Enrigue Jurado for reviewing the 
reviews. Miguel Angel Gonzalez provided the map. 



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Brittonia 14:72-90. 
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uckow, M. 1993. Monograph of Desmanthus (Leguminosae-Mimosoideae). Syst. Bot. 

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de especes des generes Phaseolus et Vigna (Papilionaceae) sur la base de donn'ees 
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Iohlenbrock, R.H. 1961 .A monograph of the leguminous genus Zorn/a. Webbia 16:1-141. 

Iohlenbrock, R.H. 1 962.Tribe Hedysareae,subtribe Stylosanthinae (Leguminosae) of Cen- 
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Piper, B. C. 1 926. Studies in American Phaseolineae. Contr. U.S. Natl. Herb. 22:663-701 . 

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Rudd, V.E. 1 956. A revision of the genus Nissolia. Contr. U.S. Natl. Herb. 32:1 73-206. 

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Ray Neyland and Billie J. Hoffman 

Department of B, 

McNeese State University 
Lake Charles, LA 70609, USA. 

Mark Mayfield and Lowell E. Urbatsch 

Department of Plant Biology 
Baton Rouge, LA, 70803, U.S.A. 
siana, Calcasieu Parish is composed of five major vegetation regions, 
defined by their respective vascuiar plant composition and physiography. A field 
:ular flora of the appr -in it 1 .|i,i hCmeters that make up the Parish 

siana herba 

ha wen 

2 examined 




1,147 specific; 


> ]<"> i''. 

entiif.n pi 


ig 147 famil 

ies found - 

lu.UM III 

s surv< 

,, Inwln 

?n compiled in an an 


catalogue t 

umber for < 

and a designation 


\n additional 134 

sorted ly 

collected in 

leu Parish th 

our visits to 

other 1 

nerbaria are 

also listed 


3 texar 

)a is report? 

;d new for Lot 


Kev Words: V 

Plant Survey, Calcasiet 

j Parish, Lc 



de vegetacion principals, Lstas r< >mposicion flonstica y su tisiogratia. 

Una prospecciondecampode la flora 1 ssculard* li i >ro imadamente 2,844 kilometroscuadrados 
que componen la parroquia fue realizada desde agosto de 1 995 a octubre de 2000. Adicionalmente, 

estudio. Los nombres de las 1,147 entradas especificas ul i rfi i u< tepresentan las 147 

familias encontradas durante esta prospeccion se han compliado en catalogo comentado que 
incluye nombre del colector y numero de la recoleccion de un especimen respresentativo y una 
cesignacion de cada especie sobre si se considera nativa o introducida. Se incluyen en el catalogo 
las regionesde vegetacion en que vive cadaespecic ielistanl nbien i aadicionalescitados 

como colectados en la parroquia de Calcasieu que no se encontraron durante la mvestigacion de 

Located in southwestern Louisiana, Calcasieu Parish lies adjacent to Beauregard Parish 
to the north, Jefferson Davis Parish to the east, Cameron Parish to the south and the 
State ofTexas to the west (Fig. D.Summers in the Parish are hot and humid;winters are 
warm but are occasionally interrupted by freezing t< 

Midkiff 1988). Annual f 1988). The Parish is 

occasionally impacted by hurricanes. 

Elevation in Calcasieu Parish ranges from sea level to 29 meters (Roy & Midkiff 1 988). 
The Calcasieu and Houston rivers drain the central and eastern portions of the Parish, 
and the Sabine River (Hardner 1960) drains the western portion. The Sabine Diversion 
Channel runs west to east near the center of the Parish and diverts water from the Sabine 
into the Calcasieu River. The Intracoastal Waterway is situated in the extreme southern 
part uf the !\)iish and tuns from Calcasieu I ake to mo Sabine River Barge taafiii. is ox I en 
sive in the Intracoastal and to a point north of Lake Charles on the Calcasieu River (Jones 
et al. 1 954; Roy & Midkiff 1 988). A deep-water port is located in Lake Charles (Jones et al. 

j Pari 


TTie recent deposits generally cc 
jthern edge of the parish and ii 
). The Pleistocene deposits cons 
ing the past four glaciation pe 

)hn< I iii I amp and the remaining area 

is urban (Roy & Midkiff 1 988). Land use is primarily devoted to producing timber, raising 
cattle,and growing rice and soybeans. Large petroleum-related industries are present in 
the Parish and are concentrated around the area of West Lake. 

The purpose of this study is to survey Calcasieu Parish for all vascular plants that are 
native or introductions that appear to be naturalized and to define and map the vegeta- 
tion regions the Parish. Cultivars and introductions that do not appear to be naturalized 

Approximately 2,000 plant collections from Calcasieu Parish were made between Au- 
gust, 1995 and October 2000. Voucher specimens were prepared using standard her- 
barium practices and are housed in the McNeese State University (MCN) herbarium. Speci- 
mens from the following Louisiana herbaria were examined to complete this survey. 
These herbaria include: University of Louisiana at Lafayette (LAF); Louisiana State Univer- 
sity (LSU), University of Louisiana at Monroe (NLU) and Tulane University (NO). 

Specimens were identified primarily through the use of the following references: 
Allen (1 992);Correll & Correll (1 941 ); Correll & Johnston (1 970);Cronquist (1 980); Duncan 
(1 975); FNA (1 993, 1 997); Godfrey (1 988); Godfrey & Wooten (1 979, 1 981 ); Gould (1 975); 
Radford et al. (1 968) and Small (1 933). For divisions Filicophyta (ferns), Lycopodiophyta 
(lycopods), Equisetophyta (horestails), and Coniferophyta (conifers), nomenclature fol- 
lows FNA (1993). For Magnoliophyta (angiosperms) subclasses Magnoliidae and 
Hamamelidae, nomenclature follows FNA (1 997). For all remaining subclasses, nomen- 
clature follows Kartesz (1 999). Appropriate experts were consulted to help identify prob- 
lematic taxa. These experts include Charles Allen, (Poaceae; NLU); Doug Goldman 
{Calopogon); )ohr\ Hays (Agalinis; University of Louisiana, Monroe); Phil Hyatt (Ca rex; United 
States Forestry Service, Pineville, LA); David Moore (various rare taxa; United States For- 
estry Service, Pineville, LA); Guy Nesom (Asteraceae; North Carolina Botanical Garden); 
Latimore Smith (various rare taxa; National Heritage Program, Louisiana Department of 
Wildlife & Fisheries, Baton Rouge, LA). 

From an analysis of plant collection data and recent soil survey maps (Roy & Midkiff 
1 988), vegetation regions within Calcasieu Parish were defined and mapped. 

Collections made by the authors form the majority of the annotated lis 

families and 530 genera. Twenty-nine ferns, fycopods an 
1 3% of the Calcasieu Parish flora and six gymnosperms aco 
aining flora consists of angiosperms. About 1 5% of the Calc 


unconfirmed lists are added, the total number of native and naturalized taxa attribut- 
able to Calcasieu Parish is 1,281. 

Three-hundred and two species, not previously recorded in the Parish, were discov- 
ered during this study. Euphorbia texana a new record for Louisiana. Five veg- 
etation regions were identified and mapped for the Parish (Fig. 1 ). 

Calcasieu Parish is noteworthy for the extreme transition from pinelands in the north to 
prairie and brackish marsh in the south. Bottomland hardwoods and swamps are associ- 
ii' I ii h iln o li i k ti . I n in lull hu ton Rivers. I « , |i i m ., tee ions mi I. u u 
Parish are recognized (Fig. 1) and are discussed in terms of soil characteristics, current 
uses,and dominant species composition. 

Coastal Prairie— -This region is characterized by soil that ranges from loamy to clayey, 
is medium in fertility and is used primarily for urban development, cropland and 
pastureland (Roy & Midkiff 1 988). The main crops are soybeans and rice (Roy & Midkiff 
1 988). Most of the rice farming is done on the prairie because of its low altitude, low 
relief and the impervious nature of its subsoil (Jones et al. 1 954). Extensive farming and 
urbanization has heavily impacted the prairie region. Dominant trees in the better-drained 
soilsofhigherelevatiu II i i i i Pmustaeda,Quercus 

alba, and Nyssa sylvatica. Trees in poorly drained soils of lower elevations include Quercus 
niiini,( literals pht Iky inrl / latuniy o< cick ntalis lletbace< m plants typically are weedy 
and include many species of grasses and sedges. 

Coastal Marsh.— This region is very low in relief and rareiy rises more than 1 .5 meters 
(Jones et al. 1954). Soil 1 , in the < oast a I rnaish region range from soft organic to firm min- 
eral clay.These soils are poorly drained, subject to flooding and used primarily for wildlife 
habitat, recreation and rangeland (Roy & Midkiff 1988). The coastal marsh region is in- 
habited mostly by grasses and sedges, and is typically devoid of trees (Jones et a 1 . 1954); 
1 1 ' i ' / i I i ii m in along roadsides. 

Marshes are either fresh or brackish. In general, marshes become more saline the closer 

Approximately 9,906 hectares of brackish marsh occur in the Parish. Salinity ranues 

i the Calcasieu River may be saline up to a point just north of Lake 
Charles where the river is dammed. Common brackish-marsh herbaceous plants include: 
Spartina alterniflora,Distichlisspicata,Juncus sp.and Scirpus sp. Approximately 7,020 hect- 
ares of fresh-water marsh occurs in the Parish. Common fresh-water herbaceous plants 
\nc\ude:Alternanthera philoxeroides,Eichhornia crassipes, Typha latifolia,Pontederia cordata, 

Swampland. — This region makes up about 4% of the parish. Soils are level, fluid, and 
loamy or clayey throughout (Roy & Midkiff 1 988). Swamps are primarily used for wildlife 
habitat and for recreation. Major swamplands are associated with the Calcasieu River in 
the northeastern part of the Parish and with the Sabine River in western Calcasieu. Domi- 
nant trees include Nyssa aquatica, and Taxodium distichum. 

Longleaf Pineland — -This region was clear cut around 1 900 and now serves prima- 
rily as forest range and cropland (Roy & Midkiff 1 988). However, a few relatively undis- 

nizable communities. We recognize four major communities within this region: hillside 
seepage bogs, semi-evergreen broadleaf acid seep forests, sodic flatwoods and acid 

A single hillside seepage bog occurs in Calcasieu Parish northeast of DeQuincy.This 
bog appears to be the headwaters of a small unnamed stream. A semi-evergreen broa- 
dleaf acid seep forest occurs along the sandy ravine adjacent to the bog. Distinctive 
herbaceous species in the bog include Rhynchospora oligantha, Sarracenia alata, and 

An area of sodic flatwoods (sensu Smith 1 996) occur in the western part of Calcasieu's 
longleaf pineland vegetation region. These flatwoods are typically saturated during the 
winterand spring but may become very dry during summerdroughts. Herbaceous spe- 
cies are often prairie like and include Spanina spartinae,Chaetopappa asteroides,Evolvulus 
sericeus, and Liatris punctata.Jhe understory woody vegetation is stunted. 

A few recognizable acid flatwoods (sensu Smith 1 996) occupy the central part of 
the longleaf pine region within Calcasieu Parish.This community is level to gently rolling 
with small elevated areas termed "pimple mounds" by Holland et al. (1 952). Under natural 
conditions and frequent fires, these savannahs support a sparse canopy of longleaf pine 
with few other tree species (Bridges & Orzell 1 989). Distinctive herbaceous species in these 
savannahs include Stylisma aquatica, and Platanthera nivea.Jhe planting of pine planta- 
tions and protecting from fire have nearly eliminated these communities in the Parish. 

Bottomland Forests —This region occurs along the narrow flood plains of the 
Calcasieu River and its tributaries and along the Sabine River near theTexas border.Soils 
in this region are level, poorly drained, and loamy throughout (Roy & Midkiff 1988). Domi- 
nant trees that occur in bottomland forests include Liquidambar styraciflua,Quercus alba, 
Q, falcata, Carya sp„ Platanus occidentalisjaxodium distichum, Pinus taeda, and Fraxinus 

Specific and subspecific entries are catalogued within their respective divisions. An- 
giosperms are further divided into the classes Liliopsida (monocots) and Magnoliopsida 
(dicots).Taxa are listed alphabetically by family,genus,and species within each category. 
The format used is species name,authority,reference,and whetherthe species is consid- 
ered native or introduced. Taxa that are designated by an asterisk (*) following the au- 
thor citation are introductions that appear to be naturalized. Authorities are abbreviated 
according to Brummitt and Powell (1 992). A specimen reference includes the name of 
the collector, the collection number and the herbarium where the vouchered specimen 
is located. Frequently cited collectors are abbreviated as: A = Allen; B = Brown;C = Cocks; 
N = Neyland; M = Mayfield;T = Thieretjh = Thomas. Unless otherwise indicated, speci- 
mens from the abbreviated collectors are housed in the following herbaria: A = NLU;B = 
LSU;C = NO; N - MCN; M = LSU;T = LAF ; Th = NLU.One specimen each from Duke 
University (DUKE), New York Botanical Garden (NY),Missouri Botanical Garden (MO),United 
States National Herbarium (US) and Vanderbilt University (VDB) are referenced in the 
annotated list. Following the collection data is an abbreviation for the vegetation region 
where each species typically occurs: prairie (Pr); fresh marsh (FM); saline marsh (SM); 
swampland (S); longleaf pineland (Pi); bottomland forests (B). Although not a vegetation 
region, disturbed areas such as pastures, fallow fields, ditches, urban areas and roadsides 
are collectively abbreviated (D). Additionally, parasites are designated by (P), lithophytes 
by (L) and epiphytes by (E). Finally, taxa that have been designated as critically imperiled 
within Louisiana (SI), imperiled due to then rarity or vulnerability to extirpation (S2), rare 
and local throughout the state or found locally in a restricted region of the state (S3), 
reported from Louisiana but without conclusive evidence (SR), or of historical occur- 
rence but no records within the pasi ?Q years (SH) are designated. I he Natural Heritage 
Program of the Louisiana Department ofWildlife and Fisheries determined these rankings. 




, - ' , - ' r- ,'t I , ,'■ . it nqrlr 
& Taylor N 564 B f- (Clute) Heller, Maples 

(MCN) B, D, Pi 

FILICOPHYTA dryopteridaceae 

Ze^atluroninBSPMap^m^ ^^^^^^1^ 
Aspleniumplatyneuron (L.) B.S.R, Maples 96 (M( N) /( $ 2% (M(=N) B 


V ,-•'.,•- ..illJJi b,FM,S LYGODIACEAE 

BLECHNACEAE Lygodiumjnn. < . - Ihunl • , • B,F 



Botrychium biternatum (S; 

Maples 106 {MCN) B 
Botrychium dissectum Spr 
Ophioglossum crotalopho 

Ophioglossum nudicaule I 
Ophioglossum petiolatum 




s (Schlect 

1402 S 


(Walt.) Steud.,N 7353 B,D 
:.,N1 334 D,Pr, 
7Torr.,/\ 75722 D, Pi, Pr,D 




Macrothelypteris torresiana (Gaudich.) Ching,* 

Maples 213 (MCN) B 
Phegopteris hexagonoptera (Michx.) Fee, A/384 B 
Thelypteris dentata (Forssk.) E.P. St. John, N J 146 

Thelypteris hispidula (Dene) Reed var. versicolor 

(R.St. John) Le!linger,N269B,D,FM,S 
Thelypteris kunthii (Desv.) Morton, 7 / 3387 B 



Map/es 817 (MCN) D 

s (Hook.) 


Froelichia grc 


Rhus copallina L, W 609 D,P 

FM, Pi, Pr 

Asimina parviflora (Michx.) Dunal, T29432 B 

Cenfe//a erecta (L. f.) Fern., N 666 D, Pi, Pr 
Chaerophyllum tainturieri Hook., W 448 D, Pr 
Ciclospermum leptophyllum (Pers.) Sprague i 

Britton & Wilson,* N 530 B, D, FM 
Cicuta maculata L. var. maculata, N 267 D, Bf 



Jutt. ex DC, N 655 D, Pi 
v1ichx.,W 736 Pi, Pr 
sComm.exLam.,W 735, 

f n v hi :. - au,.' • - \ i \ /4D,Pi 

Math. & Constance,/, /6406D,Pr Aster subulatus var.ligulatus Shinners, N 435 B,D, 

,h i i rr >n V 784 Pi Pi 

i ipillaceum (Michx.) Raf., N 652 D, ^'< ' '< ' 1 J BM 

BM, FM, S Baccharis halimifolia L, N 589 BM, D, Pr 

Ptilimnium costatum (Ell.) Raf., 727955 B, Pi, S Mi !■ button N m D, FM,Pr, 

Son/cu/a canadensis L, N / 320 B S 


//ex ambigua (Michx.) Torr 


Anstulochia reticulata Jacq., T 29428 B 

Arislohn hm s< rpentaria L, N 1418 Pi 


Asclepias hirtella (Pennell) Woods., 76702^ 

■n. /('/'a J'. /(J/l./ifi 

V 2627 (MCN)B 
:.,M&N2628 (LSU) 

fi/dem frondosa L, A 

// 102 D,FM,S 


Kiiichwiti uuttallii 1 . 

Boltonia asteroides ( 

1 .) ['Her., N 7 726 D, 

«o/?onw (//7Y»s<7Fll. r 

TV 937 BM,Pr 

Chrysopsis mariana 1 

_.,AV&M /593(MCN)Pi 


num (L.) DC, W263D, Pi, P 

i . ')V < ' ' «' - ' 

L.) Cronq.,* 7?i 66922 D, Pr 

_.) Cronq, /V /458 D, Pr 

L.,W / 22 ID, Pi, Pr 

Ctveopsis linilolia Nt. 

( oreopvs tun total N 

pKkopis amplexicai. 

Echinacea sanguine. 

L,/V 698 D,FM,S 

Elephantopus carolir 

iram;sRaeusch.,A/ 7 705 B 





cngeroe \['ja, u \ M 

jhl.exWilld., A/ 7773 D, Pr 

i A. Gray, N 546 D, Pr 

Av 7/i"kr- ,]',", 


Torr.& A. Gray) Greene, 

(Walt.) H.F.L Rock, N718 Pi 
Foug., N 1225 D 

3 (MCN) C 
indelia papposa N 

'5 anthemifolia Jus ' < >'D 
morum (Raf.) H.F.L. Rock, N1389D 

Rock, /V 577 Pi 
exuosum Raf.,A/ 660 B,D,S 

3/7more 3589 (LSU) P 

iray) Heiser Correll & Co/7* 

,)Cass.,W / 002 BM,D f FM,Pr, 

& Burtt, N 1035 Pi 

us (Walt.) DC, Costanza 

28897 Pr 
dbeckia gmndiflora (D. Dc 
Robichaux 4 (MCN) D, P 

< stand ., / 

Rudbeckia texana ( 

Senecio glabellus F 

927 B 

Solidago caesia L. 
Solidago canader 

Solidago odora Ai 

Solidago rugosa F 
Soiid(Hh) ■-.miner:: 

1 363 BM,FM 
Solidago a act a A 

Solidago tortifolia 


Sonchus oleraceu. 


1607 (MCN) P 

Beadle &F.E.Boynt.,W2S7 Pi 
M/tonfa cordifolia (L. f.) Willd., Hi 23840 D, Fl 
M/ton/o scandens (L.) Willd., N 37/ D, FM, S 
Oligoneuron nitidum (Torr.& A.Gray) Small, A 

Oligoneuron rigidum var. glabratum (E.L. Br, 

Parthenium hysterophorus L* N 1529 D 

' : 7' . ^ f ' In h- Nutt \ 355 F 
P/ucheo camp/iorafa (L.) DC, W 375 BM, D, F 


B, Pi, S Cciastiuni c)loniciatun: [huill 

"",, • „;„,,, | l ' , ,.,, ,., RuU'dU./VJ^'B, wW"M ,Ji i h'"/ , ", I II MI 


yens P. Mill., Cs.n. D Euonymus americana L, A 

3 Engelm , N 1233 B, D, S C ERATOPHYLLACEAE 


i upsrlla bursa [>a< 



Suaeda linearis (BL) 


<r- , IA.DC.A//29SC 

i ', , /-,,< / I \ ■ iB,S 

, , //! , in.ll , > Pi 

Lobelia puberula var. 

Triodanis perfoliata v 


.I, ' '.*- d,f; 

W M03 B, D, Pi, 


II . ' C 
W840 D,S 

Lonicerajaponica Thunl 


Phyllanthus polygonoides Nutt, 77i 66994 D 

Penthorum sedoides L, N 1 100 B, D, S 

Phyilanthus urinaria L* N915D 


Sapium sebiferurn (L.) Roxb.,* N 350 B, BM,D,FM, 

Sebastiania fruticosa (Bartr.) Fern., A/ 69/ S 
5f/7//ng/o sylvatica Garden ex L, A/ 1846 Pi 

Cucumis melo L * N 996 D 

Melothria pendula L, W 270 B, D, FM, BM, S 


frag/a sma/7/7 Shinners, Th 66995 Pi 

Cuscufa cuspidata Engelm., W 7482 P 

W ,/,/ii/ HoIia Michx., N & M 1591 (MCN) Pi 

Cuscuta glomerata Choisy, N 1795P 

■ niciai rd// (Hemsl.) Airy-Shaw,* W/220B 

Cuscuta pentagona Engelm. var. pentagona, N 


/\cooa farnesiana (L.) Willd., A/ 620 BM, FM, S 


Cyrilla racemiflora L, N 856 B, Pi 

Amorpha fruticosa L, W r?/'/ D, Pr 


/\p/05 americana Medik., A/ 750/ B 

Astragalus distortus Torr. & A. Gray, Cs.n. Pi 

Baptisia alba var.macrophylla (Larisey) Isely, A/ 608 



Diospyros virginiana L, A/ 73 75 B, Pi 

Baptisia bracteata var. laevicaulis (A. Gray ex 



Elatine triandra Schkuhr, G/ve/w 2/88 (LSU) D 

eopf/5/a sphaerocarpa Nutt. /.osse/gne 74 // (LAF) 

Centrosema virginianum (L) Benth., /V 860 D, Pi, 


Monotropa uniflora L, Brooks 343 (MCN) B 

Rhododendron canescens (Michx.) Sweet, N 659 

Cera's canadensis L,N46 1 B, S 


Acalypha gracilens A. Gray, 729898 D, Pr 
Acalypha monococca (Engelm. ex A.Gray) L 

7672 Pi 
Acalypha ostryifolia Riddell, A/ 848 D 
Acalypha rhomboidea Raf., A/ 962 D 
Acalypha virginica L, N 7629 D, Pi 
Caperonia palustns A. St.-Hil., A/ 789 D, Pr 
Chamaesyce hypericifolia (L.) Millsp.,* W/o//<er 

(MCN) D 
Chamaesyce hyssopifolia (L.) Small,* A/ 825 D 
Chamaesyce maculata (L.) Small, A/ 968 D 

Crotalaria s< 

Desmodiun cilian (Willd.) DC, ( n.D,P 
Desmodium glabellum (Michx.) DC, W 983 B 

T/v lodiumi a < W< ) » V 77 B 

Diociea multiflora (Torr. & A. Gray) Mohr, A/ 7 

Chamaesyce nutans (Lag.) Small, A/ 80/ D 


G/eo7fs/a triacanthos L,N&Komo 1635 (MCN) S 

Cnidoscolus texanus (Muell. Arg.) Small, N 72/2 
r fi jp tafus Michx.,A/ 733 D 

Glottidium vesicarium (Jacq.) Harper, A7 7027 D, 

Kumm'erowia striata (Thunb.) Schindl.,* 

Lasseigne 1434 (LAF) D, Pi 

: /. .*• )■■ .- -Jennwn G.L. Webster, A/ J/79 D 

Lespedeza cuneata (Dum-Cours.) G. Don,* 

Euphorbia corollata L, A/ 305 Pi, Pr 

Lasseigne 1597 (LAF) D 

Euphorbia dentata Michx., A/ 7046 D 

Lup/ni/s fexens/s Hook.,* W 7223 D 

Euphorbia heterophylla L.,* A/ 707 7 D 

Medicago arabica (L.) Huds.,* N & Komo 76/0 

Euphorbia meganoesos Featherman, A/ 686 D 

(MCN) D 

:..;.•.■ -.-.., .'exono Boiss., W& M 7/76 (MCN) D 

Medicago lupulina L,* A/442 D 

'.■■■■; .-, :..-.■■. ,'.l • ■;■■. ; :?,'.■ |. D 

Quercus acutissima Carruthers* N 1785 D 

Mt\, .,■ , - lymorphaL* N 1174D 

6 ' B 

, ■' ' 1 ^ 1 .) 1 1111 " I ( ://)( ' i/W 

Quercus falcata Michx., A/ 646 B, D, Pi 

(LAF) D 

Quercus hemisphaerica Bartr., 728055 D, Pi 

Quercus incana Bartr., 725652 Pi 

ill ill > 1 Mil II ' ! 'II 

B, S 

1 ut net, W / 544 D, Pi, Pr 



D, Pi 

. M Tort & A Gray, N 752 D, Pr 


5 7 B,D,Pi,Pr,S 

Quercus pagoda Raf., A / /47 B 

> ■ u- , , 1 . , " If M || , | v : l 1t 


Quercus shumardii Buckl., W& M /7/7 B 

OihcMlum s/nip/exdorr.&A.Citav) Hyritx. /V ? -77 

< , dh» ii ; Pi 

Quercus sfe//ofaWangenh., A / 083 Pi 

Parkinsonia aculeata L„* N 57/ D 

Quercus fexono Buckley, A/ & K/'tt /704 D, Pi 

Quercus virginiana R Mill., W 475 B, D, Pi, Pr 

-U: /;. /;. ',,',; luiii^hu i hm I, i i . A. 1 ii.iv, ( ' > • 
Her 16 (MCN) Pi 


« >7 r ' [)tLJce,*W/30/ D, 

' ' i.7 il ' 1 " \ '026 D, Pi 

' , ' i s F ji hi 1 i 1 , t ifll l i) 
D, Pi 

' ll 1 ■ i 1 Ml ', " ' 


' 7' ; ,i in i,Am N 1104 D,Pi,S 
Sabatiacamp ,i , \ ' .r -'. D,Pi 

D, FM,Pi,Pr 

Sabof/o dodecandra L. (B.5.P.) var. fo//oso (Fern.) 

Wi bui.fiuu /07 (LSU) Pi 






\<.'-,ihini:.i laiinicu ii ] i 1 nth " i / s / ;n / M 


(LAF) D, BM, FM 

Strophostyles helvula (L.) Ell, W 928 D, Pr 

Itea Virginia L,N 491 S 

sfM/'ficsii'A-, umbeiinui (,'vU,hl.e\Wi!klJ IMUon, 


N 1 540 D,Pr 

\ n " < ' "o'u (L) B5P N7/9Pi,Pr 

■ ■ ■ >nobrychoides Nutt.,/V /396 Pi 





i ,- , lout ilacq) Benth.,W9< 

; ' D, BM 



40 B 




■no 16 1 7 (MO 



Carya aquatica (Michx.f.) Nutt., N 1042 B 
Carya cordiformis (Wangenh ) K.H < n h \ . 


Linum medium (Planch.) Britt. var. te 

(MCN) B 

1 Mm i , t N 1085 B, Pi 


■ D,Pr 

n a; ,m Walt., N 7774 D, Pi 


flat nigra L.; D, Pi 

Hedeoma hispida Pursh, N 1326 D, Pr 

, , , 1 1T In ii r 'J T ^D,Pi,Pr 

■ . ' 1 t, '. / »79D,FM 

nperv/rens (L) J. St.-Hil,N 

Mitreola petiolata (J. F. Gmel.) Torr. & A. 
743 D 

' j i/j i-"a(l r Gmel.) G. Don, N. 

Ammannia coccinea Rottb, N 980 D 

/I'm nkircia in ulheimeii A. ( ,ray, Com 1 // & Co/r<?// y/> /2 
Monarda punctata L, W 84 7 D, Pr 

Cuphea carthagenensis (Jacq.) Macbr.,* 1 
Didiplis'diandra (DC.) Wood, N 1752 D 

Pe; 'Mmfe cem L) Britt ,* N 7268 D 
' '■! i f j , / , i , n II N 50D,Pr 

in^ino N 643 D,Pi (52) 

. ' , " ■ ' Mith ssp. praemorsa 

& A. Gray ex Rothrock, A/ 339 D, FM 
Rotala ramosior (L.) Koehne, N 924 D, FIV 

Magnolia grandifloro I ., (V 1381 B 

Salvia azurea Lam., A/ 7050 D,Pr 

Ca///rfjoe popawr (Cav.) A. Gray, W 733 7 D 

Sa/wa coccinea Buchoz ex Etlinger,* Cofo 45 

Mh ' 282 D, Pi 

(MCN) D 

'i i | L ; m. <irpus (Cav.) O.J 

•uilvialytala 1 .,( ostanza 9 (MCN) D, Pi 


; )pfty//o Engelm.& A.Gray, 723535 

W6fecusmo5c/jeiyfos Lssp.mosc/ieufosL.A/ 744^ 

Scutellaria drummondii Benth., 77i 66867 Pi 

TCosre/'etz^ya virgimca (L) Presl.ex A.Gray, A/ 74* 

Scutellaria integrifolia L, A/578 B, Pi 

S/ac/iys crenofo Raf„ W 7 776 D 

Stachys floridana Shuttlew.ex Benth., N 1217D 

(Torr. & A. Gray) Schery,*W 7472 D, Pr 

i Lion, N 1732 D, Pr 

Teucrium cubense Jacq!, N 1302 D 

i„ n >//<;L W726D 

. . ^r ' 1 Th 66905 D 



-I,!,, 7 , - mi Presl,*W 606 B, D, 




l>h m i/ f ? ,MWalt,A/ 283 Pi 





" , a' i -'* jL V654 D,S 

Coccu/us cara//nus (L.) DC, N 7504 D 



. 'i' , i- , , . . - ' iiiihess) H.Hara,TV7 

,, t Ihunix Ndkdi*W 1143B 


< 1 il hneider,*7V7036B,D 

Oenothera biennis L, TV 7022 D 

Moms alba L*N 514 D 

- ' '■ 'l ,' . lift i 111, til ,1 ' '- 

Moms ruhia L, N 1503 B,D,Pi,Pr 



Oenothera heterophylla Spach, TV 1453 D 

Myrica cerifera L, TV 35 1 B, D, Pi, Pr 

' il V A! 7598 (MCN) Pi 

i - V e<j , rufc7 Hill, A/ 560 D 
Oenothera linifolia Nutt., AT 1241 Pi 
Oenothera spachi,,' a \ n l i 



()eih)thera s/xY/osr; Null., TV T i,'T D 







■ ni >a, /An ; ( i(Aiton)Aitonf.,NM2D,BM,FM, 

Ox. As AeA/A - oi. i i»j o/ il n loin * ^ l . 

7778 (MCN) D 

Nymphaea elegans Hook., W 7006 BM (S1/S2) 

Nymphaea odorata Alton, TV 7064 D, BM, FM 

Oxalis dillenii Jacq, Cosranza T 7 (MCN) D 




i i lonealhifi ua llornem., N 1736D 

Nyssa sylvatica Marshall, TV 7250 Pi 


A. > L ; >. >mataL,N346D 



Tomoh^ nvixv/oin,; ! ., TVS, tomo 7679 (MCN) B, 


Phytolacca anwricann 1 .,/Vt>A'D 

Fraxinus caroliniana P. Mill., N 664 S 


Imxints fvnnsylvanica Marshall, TV 7076 B 

Plantago aristata Michx., TV 596 D 


Plonniaohcic!uph\!ia Null M,;' t >o V / (M( N) 1 

n (' ii Mt.K t * "V& /M T595 (MCN) 

-" a> >;,„!, ■ , , f L, TV 642 D 

Plantago virginica L t N 444 D 

L/gusfrum s/nen* L>ur.* A/ 348 D,Pr 



P/afonus occidentals L.N513 B,S 

Gouro longiflora Spach, N941 D 

Gaura parviflora Douglas ex Lehm.,/V 7406 D,BM 

, I ks 394 (MCN) B, D, Polygala 

Ludwigia decurrens AAA .. A >A •'< D, 
, i tA< Ana Walt, N 922 C 

i " . ' 'fA/.i Raf,W7395Pi 

lud'viiva iriitcxaica (NuU.i Haia, ■■ 

, I , ill 1 I I I I l.H| 


/uo/o, rt/iMl Ull *J"A'D,S 

i, A25650 P 

V A»P 


i,o . i ■ ! utiA Ion &A.Gray,A 
> i i ' J' nTjfl Ell ,/V 288 Pi 


i" w , i i pitosum Blume, TV 7T 

Po/Vt/ffiiJOO Jcnsiiloni!'! Mm n \ S 
Polygonum '":/'. r In to 

, , Cll N592D 

x obovatus Danser,* N 1362D 
x pulcher L * Hebert 4 (MCN) D, Pi 

s Hook., N 690 S 
.., N / 269 B,D,S 


:., Brooks 379 (MCN) C 

. P-mat , ",pa L, N 762 B.D.S 

ematis terniflora DC,* 

Delphinium carolinianum Walt., 


* J 29393 D 

03.;V; ; ;-;:' 

.) Scop., N 591 D,Pr,5 
Fosb., W 58/ D, Pr 

w.j ■ . » ,M f /. (J i' hmners) Terrell, A/ 44 

Mitchella repens L.N276 (MCN) B 

B ! 
I \ V! 7670 (MCN) P 
')<',» i-'inwi L,*W439D 

Sa//x humilis Marshall, T 22975 Pr (S2) 
Salix nigra Marshall, N 468 D, S 


Aronia arbutifolia (I )EII.,W7 Fp 
Crataegus marshallii EggL, W 7204 B 
Crataegus opaca Hook.& Arn., N 7683 


Sideroxylon lanuginosum Michx. 

7625 (MCN) Pi 


Sarracenia alata (Wood) Wood, W /496 Pi 


l.epuropctalon spathuiaium (Muhl.) Ell., A/ /2 

4c/c///r u „ ,(CII iKaf,N7055D,Pi,P 

Prunu5 serar/no Ehrh.,/V 462 B,D 
Prunusamericana Marshall, A/ &M 760/ (MCN) B 
Pyrus calleryana Decne.,* N 1072 D 
Pyrtis communis L* N 829 D 

.) Robins., W 299 D,FM, 

■: ,(',/;//< 'M . :v . in,\i !l I '.prrrni., U;il!h'tnnin 
(LSU) Pr 
■ ' i / ' i / ><„; R a f,/V 298 D,S 

1 1 p i rider W 7454 D,FM 

■■i/uspumilus (Ruim.DSteenis," W J6, 
^cjiiioniii ticunvnaiu V. ill null " 

KSaMicht 1 

: . Gmel., A/ 7 7 95 B,S 



1 ,j'i f fHf N 'Mfn.U'li- 


Nutt., /Vfonfz 3030 (LSU) BM 

3 ray,Wn06B,S 

rte Pursh, N1 729 D 




a L,/V 272 B, Pi, Pr,S 

ens/5 (L) Small, M;7som 78 

u/cM/a (Sweet) Tro 

lenahalei Small, W 520 D 
>ena rig/da Spreng.,* N 5< 


ICorrell&Correll 9566 (LSU) 




"J, , j, f r ,[lli!,,/V 466 B 

Sfyrax americanus Lam., N 7275 B, S 

Styrax grandifolius Aiton, Wog/e s.a (LAF) B 


Symplocos tinctona (L) L'Her., W 465 B,S 




Carex complanata Torr. & Hook., N 604 B 
- -,> 1 1u | lA N&M 1727 (MCN)B 
Carex a7g/ra//s Willd., M & N 2772 (MCN) B 
Corex flaccosperma Dewey, N 553 Pr 


c /-'.--. . , - 1( ns Ell., N 802 B,D, Pi 

Echinodorus berteroi (Spreng.) Fassett, N 1012 D, 

■ re- * , •• eni Rudge, TV 603 B 


Carex joorii Bailey, 120606 B, FM 

■, i u 'hi tngelm.vaLcalycinoJh 30114 

■r/j/7 Dewey,N&/W 7775 (MCN) D 


Carex iupuhnu Willd., 7b 14405 D, Pr 

Sagittaria lanci folia L, W 7442 BM, D, FM 

Carex m/crodonra Torr. & Hook., M&N2765 (MCN) 

J45 D, FM, Pr 

i - 7 7 D,FM 

Carex oxylepis Torr. & Hook., A/ & /W / 77 ->' (M( N ; B 

Sagittaria papillosa Buch., Tollman 19 (LSU) D, 

s (Bailey) Small, M&W2775 (MCN) 

< *. 7D/iW/aEngelm.,A/372BM,D,FM 

Carex styloflexa Buckl., W&/W 7725 (MCN) B 

Auh\.,Orzell& Bridges (NL\J)BM, 


Arisaema dracontium (L) Schott, W 493 B 

( j^\ u 7 'u/</eu Michx.,/4 70027 (LAF) D,Pr 

Cladium mariscus (L.) Pohl ssp. jamaicense 

(BucklJHuttleston, A/ 500 B 

(Crantz) Kukenth., A/ 747 BM, FM 

Colocasia esculenta (L.) Schott,* W 7037 D, FM 

Cyperus articulatus L, A/ 984 BM, D, FM, S 

V 492 D,FM,S 

Cyperus compress us L, Th 66923 D, Pr 

Cyperus croceus Va hi, A/ 745 D, Pr 


Cyperus elegans L, A/ 995 D, Pr 

Cypert, < ' ( -i,if]usBoekler,Carrer8730(VDB)D 

Saba/ pa/meffo (Walt.) Lodd.exSchult.& Schult./* 

Cyperus erythrorhizos Muhl., A/ 866 BM, FM 

W 7506 D 

Cyperus esculentus L, /V 805 D, FM 


Tillandsia recurvata (L) L, N 7846 E 

Cyperus flavescens L, W 7077 D, Pi 
Cyperus giganteus Vahl * 8 977 9 FM 

Tillandsia usneoides (L) L, N 7376 E 

1 °07D,Pr' 


Cyperus odoratus L., A/ 694 BM, FM 

Cyperus oxy/ep/s Nees ex Steud., 723505 D, FM 


Cyperus pseudovegetus Steud., W 9 70 D, Pr 
Cyperus retrorsus Chapm., N 834 D, Pr 

Commelina diffusa Burm.f, A/ 97S B, D, FM 

Commelina erecta L, A/ 809 D, Pr 

'. 7207 D,Pr 

•on) Smyth, 7b 

Cyperus rotundus L* A7 7073 D, Pr 

Cyperus strigosus L.N911 D, Pr, FM 

Tradescantia ohiensis Raf, N 7220 BM, D 

' j, j . i a r/ ; 'r < r » r I u h Orzell & Bridges 

Cyperus surinamensis Rottb., 73 7645 D, B 

Cyperus virens Michx., N 663 B, D, S 

Eleocharis acicularis (L.) Roem.& Schult, Th 87978 



Eleocharis baldwinii (Torr.) Chapm., N 927 D 

Carex abscondita Mackenzie, A/ &M 7726 (MCN) 

• ' , i 7 67 S D, Pr 


Eleocharis montcr -hut! F •» rr hull 7 

Carex a/Mrans Willd. var.australis (Bailey) J.Rettig, 

M & A/ 277 7 (MCN) B 

Eleocharis montevidensis Kunth, N 528 D,Pr 

Carex onnecfens (Bickn.) Bickn.,/W&A72766 (MCN) 

Eleocharis obtusa (Willd.) Schult., 7294 75 D, Pr 


i (Michx.) Roem.&Schul 
.) Roem.&Schult., N 964 1 

■ " > < , i 'nttb "npQ24D,Pr 

i 1 , . e 405 (LAF) Pi 

i . '2n> (LSU) D,FM 

Kyllinga pumila Michx., N 9 13 D, Pr 


Rhynchospora caduca El!., W 908 Pr 


ildlu II. i ^ < . ,. ' 

/,7/,/^'s 7404 (NLU) Pi 

(NLU) Pi, S (SI) 

■ 'v ' r " J )/ < >: il .! H Pr. ill. i ! ." , " 'V 


(MCN) D,Pi,Pr 

• v - • ,') l, " , ,' 1 iir • .[ i 

407 (LAF) D, BM 


i jfim ex M.A.Curtis, 

Eriocaulon decangulare I „ N 796 Pi 


/.<!. hnoamlon anceps (Walt.) Morong, W 72/ 9 Pi 

Rhynchospora eltiowi A. Dietr., Th 14366 D, Pr 


' r ' • < , ' , , i 1 ,ih . r-i 

fgerio densa Planch.,* W 720 BM, FM, S 

Rhvm h:';pora 'ililoihi A. uuy, CV.v// ^ Htniqes 


Ofte//o alismoides (L.) Pers.,* W 7007 BM 

i h\ .i 'i - /) )/., , oln, ,- i. hapm i m ill .V ' 

Vallisneria americana Michx., N 7467 BM 


• ' ■ ' ■ u , , ! Vahl., T 31652 D,Pi 


* ,' ' ■ , , , ' ' i i . 'Pi 


Herbertia lahue (Molina) Goldblatt s^.aiotulea 

»H ' i i il il 1 iii II i Ilium i 

(Herbert) Goldblatt, N 1273 D, Pr 

Rhynchospora rr 

Rhynchospora microcarpa Baldwin e 

, . - ,, Mil nl D,FM 

lailhciman sji. (LSU)B,S 

\,N 501 D,Pr 
•r 393 (MCN)D, 

, i i V 722 D, Pi, Pr 
)//Wiegand,7?i87587 Pr 
s L,W 543 D,FM,Pr 
//■ Chapm., Brooks 609 (MCN) C 

A/ncusnoc/aO/s Coville,C 

Spirodela polyrrhiza (L.) Schleid., W 708( 
Spirodela punctata (G. F.W. Mey.) C.H.Th 

M<>/' bi , " ns/ Weddell W7208S 
L<Vo//7w columbiana Karst., TV 7 7 '> / S 
Wolffiella lingulata (Hegelm.) Hegelm, 


Habranthus tubispathus (L 

Platanthera ciliaris 
Platanthera flava (L 

,/vj /m/ - ,1 S Watson, A/ 582 P 

Spiranthes vernalis Engelm.& A.Gray, TV 669 E 

Tipularia discolor (Pursh) Null, TV / 62-'? B 

Andropogon ternarius Mic 
Andropogon virginicus L, i 

V 565 Pi 

ii ■".<", < -i i (Reyel)Ownbey, 


^r/st/da oligantha Michx., 7" 9644 D, Pi, Pr 
jistidapa i,iiis (Chapm.) Vasey,N 760(7 Pi 

' '-., Brooks 802 (MCN)C 

Arundinaria gigantea (Walt.) Muhl., TV 264 B, Pi 
Axonopus fissifollus (Raddi) Kuhlm., A/974 D 

; (Fluegge) Hitchc, 8et a/. 8668 

Bouteloua hirsuta Lag., C 

,: D L 

Brachiaha platyphylla (IV 

A'ncht'! Nash, 


Bradi/ara fexona (Buckl. 

) ST. Blake, 7?i 6689 ID 

Briza minor L* Costanza 


' Np.-w D 

Bromusjaponicus Thunb 




Chasmanthium latifoliun 

Mb ', 


/ 4372 B,S 

C/i/or/s canterai Arechav., 


< . UiSUiD 

Chloris virgata Sw., N 697 


Cinna arundinacea L,Cs 


Coelorachis rugosa (Nutt 

Coelorachis cylindrica (i 


i ynoJon iiih !ylan (1 .) Pers,,'* W 5 J ■ 


T aegyptu 

1 '") ciciculare 

/m (L.) P. E 

!eauv.,*N 894 


Poir.) Gould & 

CACIark,fi<>f« ( ,-, , Pi.. Pr 

Dichanthelium acuminai 

Gould & C.A. 

ata{l.) Greene, N 987 BM 

' . | Link,*W 7425D 
', , I P Beauv.*/V 852 D 

i) Heller,* W 865 BM, 
* Gdsfan 30 (MCN) D 

, Gray, 77? 66898 

3 (Michx.) P. Kunth, Th 66899 

Dichanthelium leucothrix (f\ 

Pish) Freckman,A 

,1 • • , ,• 


r -D 

Dichanthelium scabriusculurr 

I (Ell.) Gould & CA 

Clark, W 586 Pi 

.s.n. (LAI )D 

Dichanthelium scoparium (Pa 

m.) Gould, N 598 D 

Dichanthelium sphaerocarpc 

>n (HI.) Gould var. 

s.n, (1 SU) D 

sphaerocarpon,N498 D 

Dichanthelium strigosum (Mu 


var.leucoblepharis (Trin.) F 

D/g/fooa c/7/or/s (Retz.) Koeler 



) Roem. & Schult, 

■ ■.< 

Pan/cum repens L, N 989 BM, D, FM 

LeLong, 6 ef a/. 867/ Pi 

t r/0!>/(//L.'r?? Bosc ex Noes van riqidulum, 

G/7more & Sm/t/j 346 1 (NLU) D, FM, Pi 

,V Pi, Pr 

(MCN) Pi 

Paspalum boscianum Fluegge, B 5370 D 
Paspalum conjugatum BetgiusJ 20530 D 

5fus 50 (MCN) D 
Paspalum distichum L, N 992 BM, D, FM 

. . - > < .■ . Mi 'i '6 738 (LAF)D, Pi, 

!1 0022 (LAF)D, Pi 

reus (Michx.) Kunth,W 

Paspalum minus Fourn, 73 7649 D, Pi 

Paspalum notatum var. saurae Parcxlip Auijustns 

79 (MCN) D 
Paspalum plicatulum Michx., Augustus 1 1 (MCN) 


.flraofa 805 (MCN) Pi, Pr 

Trlpsacum dactyloides (L.) L, A/5 79 D 
l/u/p/o ocrof/ora (Walt.) Rydb., N 552 D 
Zizan/o oqt/af/ca L, N 1459 BM, FM 
Zizaniopsis miliacea (Michx.) Doell. & / 



, [,m, i. ud * N940D 

=tum glaucum (L.) R. Br.,* N 779 D 

s angusta Nees ex Trin., Dutton & Taylor 

;EII„ Griggs 77(L5U)B 
,- (L.) Desf.,* N 1222 D 
isis (Lour.) W.Clayton,* 


., N 266 D,F, 


Potamogeton diversifolius Raf., N 799 D, FM, S 
41 FM, BM,S 
Potamogeton pusillus L, N 1393 BM, FM 

7630 (MCN) B,D 
. , N 74 73 Pi, S 
mila m 'a Walt., N 1 132 B 

' .L N534 D,Pi 
Sm/7ox smo//// Morong, N 606 B, D, S 
Smilax tamnoides L,N&M 1632 (MCN) B, S 
Sm/7ax walteri Pursh.N 73 1 B,S 

Typha angustifolia !_.,* A/ 7364 BM, FM 
Typha domingensis Pers, N 13 18 BM, D 


riataB\.,Orzell& Bridges 7 

ilia var. iridifolia (Chapm.) t- 
a Chapm.,/V 862 Pi 

■ ■(.■,u)P 

/6/4(MCN)Pi S 

D, 3( Pi 

Pi Xyris torta Sm. in Rees, N&A4 7592(MCN)Pi 



m ii port ill oll< [« mi i il j i.mi mii h M i ii ti >t Mi. I nt m 1 1088) or by Thomas 

(1993 1 

listed alphabet!, nil, I f , mi\ 



Ruellia noctiflora (Nee 


Asimina triloba (L.) Du 

Polytaenia nuttallii DC 

Arnoglossum plan tag i, 
Chloracantha spinosa 




Lobelia nuttallii JAScI 

Echinacea angustifolia DC. 

1 ' a < , • Mil Mil* 

Erigeron pulchellus Michx. 
Eupatorium altissimum L. 

luiLHuiunn incamahi,m\\l,\\l 
Eupatorium pilosum Walt. 
Eupatorium pinnatifidum Ell. 

/ nlhiinuu uianunii, ,11, i M ] Nu 

,, MMiJ; ■ ■ 



mnanthum Enge\m.& A.Gray Ceanothus ameiicanw, 1 . 

Hypericum nuditn it M 



Chamaesyce hirta (L.) Millsp. 

1 J- i /< i'.c- prostata (Aiton) Smal 

Crataegus sabineana Ashe 

■_•, ' ■ >,!^tmenanus Scheele 

Crataegus spathulata Michx. 

. tnoqynous Michx. 

' / 'i/ana Duchesne 

Pruni/s angustifolia Marshall 

Phyl Ian thus carol miensis Walt. 



Go//um p//05um Aiton 








Agalinis pinetorum Pennell 

Castilleja indivisa Engelm. 
Gratiola neglecta Torr. 
Verbascum virgatum Stokes 

Physalis heterophylla Nees 
Solarium americanum P. Mill. 



Plantago rugelii Decnt 
Plantago wrightiana C 

Poly gala curtlsii A. Gra' 
Polygala cymosa Walt. 

Polygala lutea L 
Polygonum convolvuk 


: ■.■^e.'-.^'i pnlv>:achyos R^trb.wi 
Eleocharis elongata Chapm. 
- > ' c '.mcrostachya Britt. 
- alastrisd.) Roem.l 

■w/? ( !',?r., 

' ' " > u 

& C.A.Clark 

(T p 9 e,..e 

x A. Gray 


n ;-,v <'- i 

h ,,„ „ ti 


[Ell.) Gould &C.A. 

':;;';' : ';;;; 


Clark ,raiLU,nu ™ umL 


Ptilmnium nuttallu (DC.) Britt. Physostegia pukheihi I undell 

ASTERACEAE Sa/v/a rc/fexa Hornem. 

L/aiY/s sp/cofa (L.) Willd. LAURACEAE 

.) Beadle Persea borbonia (L.) Spreng. 


i!Htlth\ hi, I !iihii:i 




We thank R. Dale Thomas (NLU), Charles Allen (NLU), Carrie Landry (LAF), and 
Bradburn (NO) for assisting us in locating specimens at their respective herbaria;L 
Smith (louisi.ina Natuial Hentage Program) for his technical assistance; Paul 
Pobei t Pumsey (McNeese State University) for helping us locate several unc omn 

McNeese State Univootv ihmugh the Shearman Research Initiative Fund. 

ilen, CM. 1992. Grasses of Louisiana 2nd Edition. Cajun Prairie Habitat Preservation Soci 
ety, Eunice LA. 
ridges, E.L.andS.LOrzell.1 989. Longleafpine communities of the West Gulf Coastal Plair 

Nat. Areas J. 9:246-263. 

Brummitt, R.K. and C.E. Powell. 1 992. Authors of plant names. Royal Botanic Gardens, Kew. 

Correll, D.S.and H.B.Correll, 1 941 .A collection of plants from Louisiana. Amer. Midi. Natu- 
ralist 26:30-64. 

Correll, D.D. and M.C. Johnston. 1 970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner,TX. 

versity of North Carolina Press, Chapel Hill. 

Duncan, W.H. 1 975. Woody vines of the southeastern United States.The University of Geor- 
gia Press, Athens. 

Fenneman, M.N. 1 938. Physiography of eastern United States. McGraw Hill Book. Co. Inc., 

Flora of North America Editorial Committee. 1 993. Flora of North America north of Mexico. 
Vol. 2. Pteriodophytes and Gymnosperms. Oxford University Press, New York & Oxford. 

Flora of North America Editorial Committee. 1 997. Flora of North America north of Mexico. 
Vol 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford University Press, New 
York & Oxford. 

Godfrey, R.K. 1 988. Trees, shrubs and woody vines of northern Florida and adjacent Geor- 
gia and Alabama. The University of Georgia Press, Athens and London. 

Godfrey, R.K.and J.W. Wootfn. 1 979. Aquatic and wetland plants c 
States. Monocotyledons. University of Georgia Press, Athen 

Godfrey, R.K.and J.W. Wooten. 1 981 . Aguatic and wetland plants of the s 
States. Dicotyledons. University of Georgia Press, Athens. 

Gould, F.W. 1 975. The grasses of Texas. Texas A & M University Press, College Station. 

Hardner, A.H. 1 960 The geology and ground-water resources of Calcasieu Parish, Louisi- 
ana. United States Government Printing Office, Washington. 

Holland, WC.,L.W. Hough and G.C.Murry. 1952. Geology of Beauregard and Allen Parishes. 
Louisiana Geol.Surv.Bull.27. 

Jones, P.H., A.N. Turcan, and H.E. Skibitzke. 1 954. Geology and ground-water resources of 
southwestern Louisiana. Department of Conservation, Louisiana Geological Survey, 
Baton Rouge. 

Kartesz, J.T. 1 999. A synonomized checklist of the vascular flora of the United States, Canada, 
and Greenland. 1 st. ed. In: Kartesz, J.T. and C.A. Meacham, eds. Synthesis of the North 
American flora, Version 1 .0. North Carolina Botanical Garden, Chapel Hill, NC. 

Mac Roberts, D.T 1 988. A documented checklist and atlas of the vascular flora of Louisiana. 
Published by the author, Shreveport, LA. 

Radford, A.E., H.E. Ahles, and C.R. Bell 1 968. Manual of the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill. 

Roy, AJ. and C.TMidkiff. 1988. Soil survey of Calcasieu Parish, Louisiana. USDA, Soil Conser- 

Small, J. K. 1933. Manual of the southeastern flora. University of North Carolina Press,Chapel 

Smith, L.M. 1996. The rare and 

Baton Rouge. 

Thomas, R.D. and CM. Allen. 1 993. Atlas of the vascular flora of Louisiana. Vol. I: Ferns & fern 
allies, conifers, & monocotyledons. Louisiana Department of Wildlife & Fisheries, Natu- 
ral Heritage Program, Baton Rouge, LA. 

Thomas, R.D. and C.M.Allen. 1996. Atlas of the vascular flora of Louisiana Volume II: Dicoty- 
ledons Acanthaceae - Euphorbiaceae. Louisiana Department of Wildlife & Fisheries, 
Natural Heritage Program, Baton Rouge, LA. 

Thomas, R.D.and CM. Allln. 1998. Atlas of the vascular flora of Louisiana Volume III: Dicoty- 
ledons Fabaceae - Zygophyllaceae. Louisiana Department of Wildlife & Fisheries, Natural 
Heritage Program, Baton Rouge, LA. 




Kevin D.Janni 

Botanical Research Institute of Texas 

509 Pecan Street 

Fort Worth, TX 76 1 02-4060, U.S.A. 

Joseph W.Bastien 

Department of Anthropology and Sociology 
University of Texas, Arlington 

> Kallawaya and should 
-ograms.The rapid loss 

3, Conservation, Ethnobotany, Kallawaya, Pharmacopoeia 

lawayade Bolivia siguen una tradiciondecu radon de mcis 

splantasa lo largo del tiemporc 

The Kallawaya herbalists of Bolivia am mnowned thioughoui Anjentina, Bolivia, Chile 
and Peru (Bastien 1987) (Fig. 1). Living at altitudes of 2700-4300m and frequently travel- 
ing to communities in varied ecological zones, the Kallawaya have not only established 
a continuity in Andean folk medicine, they have also had the opportunity to greatly 
augment their pharmacopoeia alone; die way (Bastien 1 983; Abdel-Malek et al. 1996; 
Janni & Bastien n.d.). They follow a medical tradition from the Tihuanaco (400-1145), 
Mollo(1145-1453),lnca (1438-1 532),Spanish (1532-1 825),and Bolivian Republic (1825- 
present) periods (Oblitas-Poblete 1 969; Bastien 1 982, 1 983, 1 987; Abdel-Malek et al. 1 996). 
The Kallawaya utilize nearly 900 plants (Girault 1987) of the 2000 medicinal plants re- 
ported in all of Bolivia (De Lucca & Zalles-Asin 1 992). In contemporary times a syndicate 
of herbalists known as the Society for Bolivian rraditional Medk ine (SOBOMETRA) has 
been responsible lm | i in i i I ii > nut e n i I ill > \ il il knowledge, while 

the Servicio Integrado en Salud (SIENS) clinics in La Paz utilize both physicians and 

(Abdel-Malek etal. 1996). 

Compiled from a survey of historical and ethnobotanical literature are 28 plants 
that have been present in the health and healing practices of the Kallawaya since pre- 
Columbian times. This indigenous cultural knowledge extends deep into Kallawaya 

'-it v. L-v taiqeting pLinlmn indigenous pharmee ot icmias foi c onset va Lion priority we not 
only help sustain traditional cultural knowledge and biological diversity, but also the 
ethnomedical practices of the community. Focusing ethn botai ca I research on com- 
munity level priorities helps target plants of cultural importance that frequently go un- 
noticed by global conservation programs that are, "...the action of outsiders who are 
culturally and politically detached from the threatened envininnmnts and who identify 
species for conservation through western economic models"(Etkin 1998). Discussion of 
the conservation priorities of ethnobotanical res- ,m heeiv, ten mmhably rare (Alcorn 
1 995; Benz et al. 1 996; Ftkin 1 998; Eisner & Beiring 1 994; Laird et al. 1 997; Posey & Balee 

Data on pre-Columl n < tin n i i n i iml plants were com- 

piled by a survey of the historical literature (Anonimo 1703;Calancha 1638;Cobo 1891, 
1892, 1893;Contreras & Valverde 1650; Jimeneza 1965; Monardes 1569;Oviedo 1535; 
Polo deOndegardol585;Vega1 609; Yacovleff & Herrera 1935). Information on medicinal 
plants was often fragmentary and incomplete, but fourteen plants had two or more 
references. Information from modern ethnobotanical literature was considerably more 
substantial and was compiled to see what plane, had petsemd since pre-Columbian- 
era. Data on the Kallawaya pharmacopoeia wet.- - ompiled b i , > u 982,1983,1987), 
including information et 1 therapeute eees and nor median.! 1 m economic uses. The 



|:|| : |L Charazaniii* A # P0 M3 piri * / ) 

^v^requipa . /// //V ^ # Cochabamba *— , 



:::;::::. : :;J::;::::.: : :ipj:-? : :; : :.:,;:-:::-.:::.i / y 

1 ; / /-~ 

A Kallawaya Ayllus ) y 
• Cities & Towns ( \ 





1 : ^ 





( f' -'■-"- 


/ /"" '"" 


n of University of Utah Press). 

Ilawaya reportedly have an unwritten pharmacopoeia of over 900 medicinal plar 
r the purpose of this study, the pharmacopoeia is limited to published informati 
d follows the nomenclature therein (Bastien 1 982, 1 983, 1 987). 

Relative importance values were calculated for each plant using a normalized nu 
rof pharmacoloi|K il propeith .1 < md a normalized number of body systems (i 
ated.This approach follows the one used by Bennett and Prance (2000) for measuri 

whole. For 

example, Minthostachys andina is used to treat 1 body systems,the most of any plant in 
the pre-Columbian pharmacopoeia. Therefore, it has a normalized BS value of 1 (10/1 0). 
Erythroxylum coca is used to tteal i I ui . tun muiI ih n T "nthostachys andina. 
The BS value for E u\ ii» " 10 '' nhostachys andina has seven pharmacological 
properties,again the mostof ati\ | t i jinl mi | it it it I . awt pharmacopoeia. 
Thus its PH value is 1(7/7). The combined PH and BS values of M. andina equal 2.0,which 
is then divided bvtw i \< l n ilnphr 1 1 , I » n I ul ]t. tl i> t. I i'i.e importance of pre- 
Columbian M. andina, 1 00. 1 his approat. h is useful for calculating the relative importance 
of a plant by taking into account the differences in number of pharmacological proper- 
u i I d\ systems It it d I a < an I t < has i | h nt i I n i | t >p ttios 
and treats six body systems giving u a pre Columbian relative importance of 65. Nicoti- 
ana rustica also has five pharmacological properties, but treats only five body systems, 
giving it a relative importance of 60. The relative importance scores for each plant reflect 
differences in versatility. 

The relative importance of each pre-Columbian plant is then analyzed compara- 

Columbian relative importance and contemporary relative ir 

relative importance of each plant. For example, E. coca has a pre-Columbian relative im- 
portance of 65 and a contemporary relative importance of 70, thus having an overall 
relative importance of 68. By comparative analysis, those plants used in pre-Columbian 
times are recognized lot then i ontmuity. Seven plants, Psoralea pubescens, Mutisa 
acuminata, Salvia haenkii, Verbena hispida, Peperomia anaequifolia, Gnaphalium 
quadichaudium, Ambrosia peruviana, are included in this table because they were cited 
in the literature without specific pharmacological or therapeutic details other than be- 
ing medicinal. There is no known pre-Columbian importance for these plants, but their 
contemporary relative importance is included.This comparative analysis is designed to 
show the changes in numbei of plum u > >. e il pi opt tties and body systems treated 
. Medical terminology follows that of 


ere are 28 plants in the Kallawaya phar 
This is a small portion of the Kallawaya pharmacopoeia and information on these 
nt i no itheis it Mi. mod a In tatun is considerably more comprehensive. These 
mts have a wide variety of therapeutic uses (1 9 in all) mainly as analgesics, diuretics, 
tiseptics, and expectorants (Table 2). Aside from medicinal uses there are 12 plants 
it overlap economically as aromatos, ornamentals, dyes, foods, intoxicants, etc (Table 
Minthostachys andina recorded an overall relative important e of 80, the highest of all 
pre-Columbian plants (Table 4). Three plants, Erythroxvlt /to ot a, Urtica flabellate, and 
■otiana rustica .scored in the sixties and only one plant, Polypodium angustifolium ,scored 

Table 1 .Pre-Columbian medicinal plants of the Ka 

awaya pharmacopoeia. 

Genus, species (Family) 

Vernacular Name 


Ambrosia peruviana Willd. (Asteraceae) 



Azorella biloba Schlecht. (Apiaceae) 


Baccharispentandii DC (Asteraceae) 


7- ion, -une/form/s R&R(Schrophulariaceae) 


Calceoiaria aff. engleriana (Schrophulariaceae) 

Puru Puru 


Chenopodium ambrosioides L. (Chenopodiaceae) 

Paico Lombrio 


Cinchona calvisa Wedd. (Rubiaceae) 

Quina Cascarilla 

Datura sanguinea L.(Solanaceae) 



Equisetum bogotense HBK. (Equisetaceae) 

Cola deCaballo 

/ sea Lam. (Erythroxylaceae) 



Gentiana lutea L. (Gentianaceae) 



Gnaphalium quadichaudium DC. (Asteraceae) 

Minthostachys andina Benth. (Lamiaceae) 

Myroxylon balsamum L. (Fabaceae) 



Nasturtium officinale R.Br. (Brassicaceae) 



Nicotiana rustica L (Solanaceae) 

Peperomia inaequalifolia R&R (Piperaceae) 


Plantago tomentosa Lam. (Plantaginaceae) 


Polypodium angustifolium SW. (Polypodiaceae) 



Polystichum aculeatum SW. (Polypodiaceae) 

Helecho Macho 



Psittaca tm s in liu Ht P.(l ) anth i ea 


So/wa /joenW/ Benth. (Lamiaceae) 

S:i-.! Cr,:-',ii(- 


LCalancha 1638 


7. Polo de Ondegardo 1 585 

in the fifties (Table 4). Twenty-three of the 28 pre-Columbian plants have a relative \rr\- 
portance under 50.Seven plants, Polypodium angustifolium, Plantago tomentosa,Onchona 

calvisa, Gentiana lutea, Polystichum aculeatum, Psittachanthus cuneifolius,and Solanum 
radicans, show an increase in pharmacological properties and body systems treated 

Um Alhorapeutk properties of he i olumhian nieilu inal plants in ;'■< K.illawaya pharmacop 

A analgesic. A1 oiUhepti*. A2 astringent. C . aulmlonn. .D disinflammatoiy.DI Dili 

m duphoi<Mk.D3 hemukenl.E emetu. El expectorant. F febnfuge.R - '- refngofat 

S 1 V 

A A1 A2 C D D1 D2 D3 E El F R R1 S SI S2 T V VI 

Equisetum bogotens* 

^-Columbian pharmacopoeia elucidates the importance of medicinal 
:h and healing practices of the Kallawaya. Outside the pharmacopoeia 
ulturally useful as food, 'ot hygienic purposes, ornamental, and other 

i t u if i [It ii arguably impoount | nt of the traditional cul 

A B D E 

Baccharis pentand 

! lit ,' , ' I , t , 

Diological knowledge of the Kallawaya. Along with plants hav- 
', the Kallawaya have integrated exotic plants into their 
pharmacopoeia (Janni & Bastien n.d.). Despite the integration of exotic species into 
the pharmacopoeia, the 28 plants discussed herein have retained much of their cultural 
and medicinal importance for over a thousand years. 

The diversity of therapeutic uses of these plants is remarkable. Clearly, by the time 
of Spanish invasion, the Kallawaya had actively investigated the phytomedicinal poten- 
tial of the local and regional flora. The diversity of therapeutic uses indicates the 
ethnomedical sophistication of the Kallawaya, and offers a picture of the health and 
healing concerns of pre-Columbian Kallawaya culture.This information is useful in un- 
derstanding the epidemiological fluctuations of the Kallawaya throughout time by re- 
vealing the patterns of health and sickness that enable us to ask questions as to why 
they changed. Also, by compiling such data we find the plants that are not only impor- 
tant medicinally, but also have been an integral part of Kallawaya cultural heritage. 

Many ethnobotanical investigations compile information in an effort to identify 
potential new drugs; we have compiled this information to identify plants of cultural 
importance as well as plants of priority for biological conservation and sustainability 
programs. Local efforts in conservation offer greater potential results than those of west- 
ern economists because they represent the intimate local knowledge of the native ecol- 
ogy and long experience with the species in question (Etkin 1998). Assessing the 
significance of specific taxa with cultural and ecological importance gives us a frame- 
work by which conservation of local biota is based on local values (Benz et al. 1 996). 

The list of 28 plants discussed herein describes a portion of the pharmacopoeia 
that has been analyzed based on local values. The long-standing persistence of these 
plants in the Kallawaya pharmacopoeia indicate continued cultural reliance on these 
taxa in ethnomedical and economic practices. The versatility of these species within the 

ximum value of 1). 16-1 7 th BS = number of body systems treated, sited in I ■. "16 1 7 th Rel BS = relative 

:ems treated, sited in 16 th & 17 th Century Literature, (normals i r , i up of 1) 16-1 7 th Rl = relative importance of 1( 

dicinal Piants.20 th PH = number of pharmacological properties, 20 :h Century.20 th Rel PH = relative number of pharmacological prop* 

' ■' - 20*' BS - ~, ^ 20 th Rel BS = relative number of pi, . i 

itury.20 th RI-re . ORI Rl 

16-1 7 th 16-1 7 th 16-1 7 th 16-1 7 th 

BS Rel BS Rl 

Gnaphahum quaaicnauaiu 

0.0 00 

pharmacopoeia and outside it also makes conservation priorities more compelling. Sev- 
eral taxa are used for more than one therapeutic (Table 2) or pharmacological (Table 4) 
' of economic uses (Table 3). Conservation 
ne neglect the socio-cultural importance of 

a chance to protect biological diversity and in the process also protect and sustain tradi- 
tional cultural knowledge, indigenous health caie systems, and plants of particular cul- 
tural significance. Conservation i-il j i hi bililv mi n - I locus on community 
level priorities allow us to work on several problems at onus Direction based from an 
ethnobotanical pcrs| i i. ana i iridic] i i i [ i lives us ih 

opportunity to address problems of social, cultural and biological importance.The data 
discussed here provides only a part or a broader invi stigation that should include field- 

The loss of local knowledge and biological diversity should be the primary concern 
of every ethnobotanist. Organizing at local levels with an ethnobotanical framework 

simultaneously piolec is biological and cultural diversity. I osmo tribal elders and the 
knowledge they enc i| .Cm I io niioimati m nihlil n i\ environment of the 
surrounding area ( on ervatiui md sust lin ibilii > programs that are sensitive to bio- 
< ul .ii I i in lib thi ill I i )iot i i it. io tionab ultural i ' Iqi md biod i il 

for future generations. 

C )iNc I i ill )N 


sustainability programs. The persistence of use of these plants throughout centuries of 
healing, as well as their role outside the pharmacopoeia for a variety of economic uses 
elucidates the importance of these plants to the cultural heritage of the Kallawaya. By 
focusing conservation efforts on [.bants of cultural importance we not only recognize 

III Mil I in i . ll < ' I ll i p [I II ill I ]l 

cal diversity. With overwhelming predictions of 60,000 plus higher plant species to be- 
come extinct by the middle of the next century, ethnobotany must be a leader in iden- 
tifying conservation and sustainability priorities by analysis of local needs. Just as local 
een tapped for new drug leads, we must go to it in the future to deter- 


of this manuscriptTed Barkley, Carol Janni,and 
BRIT and UTA libraries for assistance. 


Abdel-Malek, S., J.W.Bastien, W.F. Mahler, Q. Jia, M.G. Reinecke, W.E. Robinson, Y. Shu, and J. Zalles- 
Asin. 1 996. Drug leads from the Kallawaya herbalists of Bolivia. 1 . Background, rationale, 
protocol and Anti-HIV activity. J. Ethno-pharmacol. 50:1 57-1 66. 

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Anonimo, Jesuita. 1703.Technologia indigena. Revista Inca, 1923. Volume 1:Nos. 2 and 3. 

Bastien, J. W.I 982. Herbal curing byQollahuaya Andeans. J. Ethno-pharmacol. 6:1 3-28. 

1983. Pharmacopoeia of Qollahuaya Andeans.J. Ethno-pharmacol. 8:97-1 1 1. 

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University of Utah Press. Salt Lake City 
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of Northern South America. Econ. Bot. 54:90-1 02. 
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case study from the Sierra de Manantlan Biosphere Reserve. Sida 1 7:1 -1 6. 
Calancha, Fr. Antonio de la. 1 638.Cronica Moralizadora del orden de San Angustin. Barcelona 

Cobo, Fr. Bernabe. 1 891 -1 893. Historia del Nuevo Mundo. In Yacovleff and Herrera. 1 935. El 
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De Lucca, M.and J. Zalles-Asin. 1 992. Flora medicinal Boliviana. Los Amigos del Libra, La Paz, 

Dorland's Medical Dictionary. Shorter Edition. 1 980. The Saunders Press, Philadelpia, PA. 

EiSNER,T.and E. Beiring. 1 994.Biotic exploration fund: Protecting biodiversity through chemi- 
cal prospecting. BioScience 44:95-98. 

Etkin, N.L. 1 998. Indigenous patterns of conserving biodiversity: Pharmacological impli- 
cations. J. Ethno-pharmacol. 63:233-245. 

Girault, L. 1 997. Kallawaya, curanderos itinerantes de los Andes. UNICEF, ORSTOM, La Paz. 

Janni, K.D. and J.W. Bastien. n.d. Exotic botanicals in the Kallawaya pharmacopoeia. Econ. 
Bot. Submitted, {available from author} 

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tion coexist? New York Botanical Garden, Bronx. 
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quesirven de medicina.ln Yacovleff and Herrera 1935. El mundo ve 
Peruanos. Revista del Museo Nacional, Lima. 

Oblitas-Pobi eii,E. 1969. Plantas medicinalesde Bolivia. Farmacopea Callawaya.Los Amk 
del Libra, Cochabamba. 

Ovibi G n Fi r m t i 1 L 35 ( aft i dmgida al emperador Carlos V. in Collecion 
documentos ineditos relativos al descumbrimiento, conuuesUi v i olcm/a* inn de 
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Polo Db Ondegardo, Juan. 1 585. Lor errors y supersticiones de los Indios. In Yacovleff i 
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Posey, DA and W.E 

folk strategies.! 


David J.Rosen 

U.S. Army Corps of Engineers 

■ district, P.O. Box 1229 
Galveston, TX 77553, U.S.A. 

The genus Cryptocoryne Fisch. ex Wydl. comprises approximately 50 species distributed 
on islands and coastal areas of South East Asia (Muhlberg 1 982). Cryptocoryne beckettii 
Thw. ex R.Trim., a native of Sri Lanka (Muhlberg 1 982), was collected in 1 996 in the San 
Marcos River in the City of San Marcos.This taxon has not been previously reported in 
Texas according to Jones et al.(1 997), Hatch et al.(1 990),and Correll and Johnston (1 970). 
Large, naturalized colonies of C beckettii were observed growing in open shallow riffles 
as well as in shaded deep pools. Cryptocoryne beckettii is a valued aquarium plant col- 

theSan Marcos River is likely due to escape from cultivation or dumping of aquariums as 
has been proposed for the introduced aquatic fern Cemtoptens thalichtroides by Hannan 
(1 969). A description of C. beckettii modified from Nicolson (1 987) and illustration (Fig. 1 ) 

Cryptocoryne beckettii Thw. ex R.Trim. J. Bot. 23:269. 1 885. 

Perennial, rhizomatous emergent-submerged herbs. Leaves basal with elongate, sheath- 
ing petioles to 1 5 cm; blades glabrous, ovate to narrowly ovate, 3— 9(— 1 3) cm long, (1 -)1 .5- 
3.5 (-4) cm wide, upper surface green to dark green to brown and marbled to red-brown, 
lower surface red-tinged to more or less brownish or green; veins usually conspicuously 
red; apex acute to acuminate; base obtuse to cordate, margin entire, sometimes undu- 
late; submerged specimens mostly with larger, thinner leaves, often brownish marbled. 
Inflorescence (not seen) short peduncled;spathe 4-1 2(-20) cm long, limb greenish brown, 
narrowly ovate, 0.5-1 .2 cm wide, 1 .5-3 cm long, twisted, upright to somewhat recurved 
and twisted; spadix 1 .0 cm long. 

hree closely related specie 

•s, C w 

alkeri Schott, C. 

.A key to separate the four 


an be found in I 

i:TEXAS. Hays Co.: San Marco: 

5 River,. 

exposed bottom a 

g 1996, Rosen 202 (SAT, 5WT). 

) Don Bryne (Suwannee Labs, Florida) for I 

\.Cryptocorynebeckettii (Drawn by J.E.Dawson III from herbarium specin 


Iorrell, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation. Renner. 

UNNAN,H.H.1969.The introduction and establishment of Ceratopteris in Texas. Amer. Fern 
J. 59:1 22. 

Utch, S.L,K.N. Gandhi, and LE.Bf ■ ! 990. Ch< Mi - fthe vascular plants of Texas. Texas 
Agric.Exp.Sta. Bill. MP-1655. 

checklist including synonymy, bibliography, and index. University of Texas Press, 

TGDR.391 p., 

.H.1987.Araceae.ln:M.D.Sassanaykeand F.R.Fosberg,eds. Flora ofCeylc 
:hsonian Institute and the National Science Foundation, Washingtc 
i Publishing Co. Pvt. Ltd., New Delhi. Pp. 117-101 

DM. Persall 2000 Paleoethnobotany. A Handbook of Procedures, second edition. 

Academic Press, 525 B Street, Suite 1 900, San Diego, CA 921 01 -4495, U.S.A. 
Eleven years after its debut, Paleoethn Kv,< ' < \- I Procedures returns in second edition. 
This classic by University of Missouri-Columbia anthropologist Deborah Pearsall is back -700 pages 

nli i lull iii'h well as a geneial giowth in p iIihm urn. I m mi ,l lib num I it ill i i I 

makes a case that reconstructing past human diets may be the biggest and most immediate con- 
tribution by the field of paleoethnobotany. 

The book is laid out in six chapters followed by references and an index. Chapter one, "The 
Paleoethnobotanical A|)| o,.-k h"mclud< t inttoduction to the field, a historical overview and a 
discussion of the nature and status of ethnobotany. Chapters two and three are devoted to 
macroremainsand tht ti i migut omeco unq.iclontil n and interpreting them. Chapter four, 
"Pollen Analysis" covers the nature and production of pollen, field sampling, laboratory analysis, 
and a discussion of issues and directions in archaeological pollen analysis. Chapter five,"Phytolith 
Analysis," covers the nature and occurrence I phylolith nd methods of field sampling, lab test- 
ing, and interpreting the results. Chapter 6, "Integrating Biological Data," is the most welcomed 
addition to this volume. Broken into two parts/Indicators of Diet and HealtlY'and'The Interplay of 
Dietary Indicators," respectively, Pearsall gives a welcomed big-picture framework to the field. Part 
one of chapter six dm u m , indue, i dmtar\ factors, such as botanical and faunal data,and then 
discusses direct indicators like gut contents and coprolites, stable isotopes, trace elements, and 

ies from Ecuador and Peru. This chapter is a valuable additi 

literature by developing an approach for intern iting mtl . v iluaung multiple lines of e 
concerning diets. Pearsall discusses eight diets chosen to represent diets spanning the tr 
from hunting and gathering to agriculture in the New World tropics, which is ihem illusli 
two Neotropical case studies. Reconsl UK turn diets c, an impmtanl an hueulnqu ,il mid hio 
I I in issue k Iik It investigation can help us understand how pasi populations mii vr 
pi ' p ' Ikiis h i hi i is followed by a list of lefoiences and t Id b i im i I mill 
while figuies and tables appear frequently throughout to illustrate concepts discussed th 
Paleoethnobotany: A h m a > > ilu A I iu its < omprehensive outlin 


Walter C.Holmes 

Department of Biology 

i a lo! Urn > <f) 

WacoJX 76798-7388, U.S.A. 


United States National Herbarium 

Department of Botany 

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

Jason R.Singhurst 

Wildlife Diversity Program 

Texas Parks and Wildlife Department 

AustinJX 78704, U.S.A. 

jason.siiHihurst@tpwd. state, 

Thymelaea passerina (L.) Cosson & J.Germain was first reported in the United States by 
Pohl (1955). In reporting T. passerina as new to Ohio, Vincent and Thieret (1987) pre- 
sented an expanded account of the occurrence of the species in the United States,men- 
tioning its presence in Illinois, Iowa, Kansas, and Nebraska. Additionally, Thymelaea 
passerina has also been reported from Alabama (Webb et al. 1 997), Mississippi (Wofford 
&DeSelm 1988), Washington (USDA, NRCS 1999), and Wisconsin (Harriman 1979).The 
species is not cited in any of the recent references or checklists treating the flora ofTexas 
(Correll & Johnston 1 970; Hatch et al. 1 990; Johnston 1 990; Jones et al. 1 997), nor is it 
included in Diggs et al. (1 999) in their flora of the north central part of the state. A de- 
scription of the species and pertinent synonymy follow. 
Thymelaea passerina (L) Cosson & J. Germain, Syn. Fl. Env. Paris, ed. 2, 360. 1 859. (Fig. 1 ). 

Stellera passerina L., Sp. PI. 559. 1 753. Complete synonymy given by Tan (1980). 
Taprooted annual to 55-60 cm tall; stems erect, slender, simple or more commonly with 
few to several ascending branches in distal half, glabrous or weakly pilose distatly, yel- 
lowish green; leaves alternate, simple, exstipulate, sessile or nearly so, linear-lanceolate, 
7-1 5 mm long, 1-2 mm wide,stifflychartaceous or subcoriaceous,glabrous or less com- 
monly abaxially weakly puberulent, apex acute, margins entire. Inflorescence axillary, 
flowers often 3 but ranging from 1-7 in distal leaf axils, cluster commonly subtended by 
2 green bracts, bracts lanceolate to broadly so, to ca. 1.5 mm long, basally ciliate; flowers 
bisexual, actinomorphic,perigynous,sessile,2-3 mm long,corollasabsent,sepals4, weakly 

i, manifoM.'d i 

rcapsule,apically c 

it i ji . 'n I II •■ 

he urceolate, persistent hypanthium, hypanthium 
and calyx lobes substrigose, stamens episepalous, 8, bkydu , included, filament much 

hoil ih ir i ilu iiiihu i i i puiui ipi< ill hit i ul i I I ,] 

hypogynousdisk,bicarpellatebutuniloculai by abortion, stvlo one, terminal, short, ovule 
one, pendulous. Fruit a dry, indehiscent, 1 -seeded, 1-locul; 

' Ii".c0 within tin- 1 insistent hypanthium; seed ovoid, 2- ] i 

I ins I urasian weed ot 'I hviricLiea section lyqia (Ian I 
the United States from Alabama (Webb et al. 1997), lllinoi 
Iowa (Pohl 1955), Kansas (McGregor et al. 1986), Mississippi (Wofford & DeSelm 1 
Nebraska (Pohl 1955), Ohio (Vincent & Thieret 1987), Washington (USDA, NRCS 1 
Wisconsin (Harriman 1979), and is here newl\ document I t n 'he flora of Texas. I 
alsobeen inttodu ii r i ti Hi i I i ijtim tn> nh American popula' 

ranges from (April) June to September. 


Specimens cited: TEXAS. Denton Co.: approximately 5 mi E of Sanger on FM 455, 24 Jun 1 999, 
Singhurst8156 (BAYLUJEX, US).Fannin Co.: 0.5 mi E of Haiie Community on FR 1 550,N 33.51 1 73,W 
96.05437, 5 Jun 1999, Holmes 10173 &Singhurst (BAYLUJEX, US). 

We believe that the species was likely introduced to Texas through the use of agricul- 
tural machinery imported from further north and used to harvest wheat, the dominant 
crop in both areas.The Denton County specimen is from the Cross Timbers and Prairies 
vegetational region of the state, an area characterized by slightly acidic to acidic sandy 
loam soils (Correll & Johnston 1970). Associated species included Dalea purpurea, 
Indigofera miniota (Leguminosae), Froelichio floridana (Amaranthaceae), Helianthemum 
georgianum, Lechea mucronata, L. tenuifolia (Cistaceae), Hypericum drummondii 
(Hypericaceae), and Krameria lanceolota (Krameriaceae).The Fannin County specimen 
occurred in the margins of roads and wheat fields in clay over limestone "chalk"on the 
Gober Limestone Formation of the Blackland Prairie vegetation region of the state.Com- 
mon associates included Asclepias asperula (Asclepiadaceae), Forestiera pubescens 
(Oleaceae), Hypericum perforatum (Hypericaceae), Rhus aromatica (Anacardiaceae), Se- 
dum pulchellum (Crassulaceae),and Sophora affmis (Leguminosae). 

The documentation of Thymelaea passerina as new to Texas is not only a report of a 
new genus and species to the known non-cultivated flora of the state, but another fam- 
ily, the Thymelaeaceae. Jones et al. (1 997), in their checklist of the vascular flora for the 
state, included two species of Thymelaeaceae, Daphne cannabina Wall. (= D.papyracea 
Wall, ex Steud. according to Huxley 1992) and D.cneorum L, but both of these are culti- 

We thank James Solomon (MO) and Michael Nee (NY) for searching for specimens of 
North American Thymelaea passerina in MO and NY, respectively; and Aaron Goldberg 
(US) and Monique Reed for helpful comments on the manuscript. We are grateful to 
Tom Wendt (TEX) for providing other assistance and to Michael Vincent (MU) and John 


Correll, D.S. and M.C.Johnston. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner. 

Diggs, G.M, Jr., B.L. Lipscomb, and R.J. O'kennon. 1 999. Shinners & Mahler's illustrated flora of 
north central Texas. Botanical Research Institute of Texas, Fort Worth. 

Harriman, N.A.I 979. Four additions to the Wisconsin flora. Michigan Bot. 18:143-145. 

Hatch, S.L, K.N.Gandhi and LE Bi i l990.Chi Mm 'I the vascular plants of Texas. The 
Texas Agricultural Experiment Station, Texas A&M University, College Station. 

Huxley, A. (Ed.-in-Chief ). 1 992. The new Royal Horticultural Society Dictionary of garden- 
ing, vol. 2, D to K, Macmillan Press, London & Stockton Press, New York. 

Johnston, M.C.1990.The vascular plants of Texas. A list, up-dating the manual of the vascu- 
lar plants of Texas. Published by the author, Austin. 

1 1 klist in. fueling synonymy, bibliography, and indo 

McGregor, R.L.JM B n RLE. i ,md t.K.5cHofiELD(eds.).' 

University Press of Kansas, Lawrence. 
MmiLLNBRoi k, R.H. and D.M. Ladd. 1978. Distribution of lllinc 

Illinois University Press, Carbondale. 
Pohl, R.W. 1 955. Thymeloea passerina, a new weed in the Ur 

Sci. 62:152-154. 
Tan, K. 1 980. Studies in the Thymelaeaceae II: a revision of 

Royal Bot.Gard. Edinburgh 38:189-246. 
USDA, NRCS. 1999. The PLANTS database (http://plants.usa 

Data Center, Baton Rouge, LA 70874-4490 USA. 
Vincent, MA and J.W. Thieret. 1 987. Thymelaea passerina (Tr 

Webb, D.H., H.R.DlSi i w, and VV.M. [)i r,\r, 1 997. Studies of prai 

Alabama.Castanea 62:173-184. 
Wofford, B.E. and H.R.DeSelm. 1 988. Distribution of and first re 
ted States. Castanea 53:305-3' 

of the Great Plains. 

r plants. Southern 


Timothy W.Chumley 1 and Ronald LHartman 

While conducting a floristic survey of central Colorado (Chumley 1 998), two collections 
of a gooseberry from the Canon City area, Fremont County, proved to be Ribes niveum 
Lindl. These collections represent the rediscovery of a taxon known in Colorado from a 
single,neglected gathering byT.S.Brandegeein 1873 [Bmndegee 697, S\ur\o\X 1985).These 
new collections come from the drainage of Cottonwood Creek, 1 3 to 1 5 miles north- 
west of Canon City. Wilson Creek, the Brandegee locality, is 3-4 miles to the east or ca. 8 

The Brandegee specimen was cited by Porter and Coulter (1 874) as R.irriguum Dou- 
glas, a species of the Pacific Northwest. Rydberg (1906) excluded this taxon from his 
treatment of the Colorado flora. Presumably, he did not examine the material and ig- 
nored the taxon due to its great disjunction from its normal range (being one of several 
species"accredited to Colorado but not the intervening states/page xii).The occurrence 
of "R.irriguum" has been similarly ignored or overlooked in subsequent treatments of the 
flora (Harrington 1 954; Weber 1 953, 1 967, 1 990; Weber & Wittmann 1 992, 1 996). Sinnott 
(1985) examined the Brandegee collection in preparing his treatment of Ribes section 
Grossularia and determined it to be R. niveum, another species of the Pacific Northwest, 
rather than R. irriguum (R. oxyacanthoides L ssp. irriguum (Douglas) Q.P. Sinnott). This is 
consistent with Porter and Coulter's description of the specimen,whichm tch R.n t 
and not R. irriguum. Sinnott, however, failed to relocate the Colorado population. Exami- 
nation of the specimen {Brandegee 697) at the Missouri Botanical Garden confirmed his 
determination, and the new collections document its persistence in Colorado for over 

The normal geographical range of R. niveum is in three separate areas:the northern 
group of populations are in southeastern Washington,eastern and northeastern Oregon, 
and adjacent western and central ldaho;the middle group,southeastern Oregon,south- 
ern Idaho, and adjacent northeastern Nevada; and the southern group, west-central 
Nevada (Churchill, Lander, Pershing, and possibly Nye counties; Holmgren 1 997; Sinnott 

its occurrence in Colorado prior to settlement was found in an account of the Long 
expedition to the Rocky Mountains in 1820 (Goodman & Lawson 1995). Edwin James, 
the botanist of the expedition, collected a specimen of Ribes in the area that was deter- 
mined byTorrey and Gray as R.irriguum.A description of the fragment provided byTorrey 

Goodman and Lawson were unable to locate the specimen and thus could not confirm 
its identity.The pi. <t - nth tegion therefore is not documented before 

Brandegee's collection in 1 873. Prior to that time, Canon City was a major gateway and 
supplier to the gold and silver mines of South Park and Leadville,and it is possible that R. 
niveum was introduced into Colorado from the Pacific Northwest during the 1860s by 
gold or silver miners, it is also possible that Native Americans may have been respon- 
sible for dispersal since the localities are close to old Ute trails into South Park.Brandegee's 
collection may thus represent the parental population at or near its initial point of es- 

The plants were in flower by middune, in fruit by late July. Fruits persisted through 
at ieast early October. They were found growing along cieeks or dry washes in pinyon- 
juniper on sandy soils derived from gneiss. Intel. ,[uniK • , re in Colorado always 
grows in clumps with other woody taxa including Ptelea trifoliata L, Ribes cereum Dou- 
glas, Ericameria (Chrysothmanus) nauseousus (Pall, ex Pursh) G.L.Nesom & G.LBaird, Rhus 
'///C/'M'/i/Nutt.,and hmoer, pem/m/h him Phis could ■. hpemhot seeds by buds. 

In floral features, the Colorado material of R. niveum is a close match with material 
from the main geographical range.The most striking characters of R.niveum are those of 
the stamens. With the petals extended or erect,the stamens surpass them by 2 to 3 mm; 
filaments range from / re °> mm in lemiih.i uilheimoie, boih the anther, and filaments 
are at least moderately pubescent with prominent silky hairs. All other Colorado species 
have glabrous filaments and anthers.Vegetatively,/?. niveum has a distinctive rich reddish 
brown bark on new growth and is unarmed except for stout spines at the nodes. Ribes 
lacustre (Pers.) Poir. may have a similar hue, but the internodes are usually guite spiny. 
Another species h • In imih i ml, it n| hi il oung twigs are pale 

yellow. With age the bark of most Colorado gooseberries becomes gray and exfoliates, 

II ! I ! I I I I I III It 

Leaf blades of specimens of R.i e, zolleeted it the three known sites in Colo- 
rado are moderately to densely covered with minute, stalked glands mixed with erect to 
curved eglandulartrichomes. Those m the mam populations to the west and northwest 
vary from glabrous and ciliolate to densely clothed in minute.erect to curved eglandular 
trichomesonlyalthough the p. tih mmihi .■ till, i i em I hj. - to thefickle nature 
of glandular trichomes in many groups of plants and the variability in pubescence in 
populations over th main i. iiai n il mge >fth eiln iifference may not be 


and flowers glabrous externally, but differs in anthers purple,greenish with age vs.white. 
Ignoring anther color, it would key to R. inerme ( 1 1a), although not easily.Foran excellent 
treatment of Ribes that is largely relevant to Colorado, the reader is referred to the Inter- 
mountain Flora (Holmgren 1 997). All but two Colorado taxa are treated: R. americanum 

troduction (W. Jennings, Louisville, CO, pers. comm,), R. divaricatum Douglas. Ribes 
coloradoense Coville is placed in synonymy with the major disjunct (to the northwest),/?. 
laxiflorum Pursh. 

It is always possible that additional populations of Ribes niveum remain to be dis- 
covered east of the Continental Divide in Colorado. Two recent floristic projects have 
contributed substantially to our understanding of the distribution of species of Ribes, 
and to vascular plants in general, in this area (see The first, 
the Central Colorado project, which led to the rediscovery of R. niveum (Chumley 1998), 
included the Mosquito and Rampart ranges and Pikes Peak. It extended from Morrison 
(near Denver) west to Webster and Hoosier passes, south on the west side along the 
crest of the Mosquito Range to near Salida and south on the east side to Pueblo. The 
second,the Sangre project, included the SangredeCristo and Wet mountains, Mesa de 
Maya, and the Spanish Peaks. It was to the south from the first area, with the Arkansas 
River, in part, forming the boundary in common. Canon City is on the Arkansas River, 
north side, midway along this line of contiguity.The Sangre project extended from Pueblo 
west to Monarch Pass, south on the west side to North Pass, Saguache, San Luis, and the 
New Mexico line and south on the east side to Trinidad and Branson (collections mostly 
by B. Elliott and R. Hartman). Together, these two study areas cover nearly 1 0,000 mi 2 for 
which ca. 27,500 numbered collections were obtained. In the process, 322 populations 
of nine of the 1 3 species of Ribes reported for Colorado (Weber & Wittmann 1 992 and 
this report) were sampled during four field seasons (1 995-96, 1 998-99). Despite the high 
level of intensity in collecting, no new sites for R. niveum were discovered south of the 

i: COLORADO. Fremont Co.: Sand Gulch south along a drainage intersecting 
. 1 5 air mi NW of Canon City, 1 6 Jun 1 995, Chumley 895 (RM); Cottonwood Creek, 
' of Canon City, 25 Jul 1995, Chumley 2454 (RM); along Wilson Creek and sur- 
: east, ca. 7.8 air mi NW of Canon City, 28 Jun 1 997, Chumley 6887 (RM). 


e agreement betv 

Chumiiy, T.W. 1998. A flonstic inventory of the East Slope, central Colorado. M.S. thesis, 

I..H 111 o ii , of Wyoming, I .iinm 

Goodman, GJ. and C.A.Lawson. 1995. Retracing Majoi Stephen I I.I ong's 1820 expedition: 

ll» "n- i it it . I I i .i n it 1 1 - i of Oklahoma Press, Norman. 
Harringion,H.D. 1954. Manual of the plants of Colorado. Sage Press, Denver. 
Hoi mgren,P,K. 1997. In: A.Cronquist,N.H.Holmgren,and P.K. Holmgren. Intermountain flora: 

den, Bronx. Pp. 12-26. 
Porii r,T.C. and J.M. Coi . i o 1 874. Synopsis of the flora of Colorado. Department of the 

Interior, U.S.Geological and Geographical Survey oftheTerritories,Miscellaneous Publ. 

no. 4. Washington, D.C. 
Pa i .0 ii: ,, P.A. 1 906. Flora of Colorado. Agric. Exp. Sta. Agric. Coll. Colorado Bull. 1 00. 
Sinnoi i, Q. P. 1985. A revision of Ribes Lsubg.Grossularia (Mill.) Pers. sect. Grossularia (Mill.) 

Nutt.(Grossulariaceae) in North Amci io. Rhodora 87:189-286. 
ToRREYj.and A.Gray. 1838-1 843. A flora of North America. Wiley & Putnam, New York. 
Weber, W. A. 1953. Handbook of plants of the Colorado front range. University of Colorado 

Press, Boulder. 
Weber, W.A. 1 967. Rocky Mountain flora. University of Colorado Press, Boulder. 
\ i \A * P>oo Colorado flora: Eastern slope. University Press of Colorado, Niwot. 
Weber, W.A.and R.C.Wittmann. 1992. Catalog of the Colorado flora: A biodiversity baseline. 

University Press of Colorado, Niwot. 
WrnrR, W.A. and R.C. Wn imann. 1 996. Colorado flora: Eastern slope. Revised edition. Univer- 

otv 'loss ol Colorado, Niwol. 



, mwlcv M.' . >4 ' , i 

Charles M.Allen 

Biology Department 

ei'sii) ■ uisioi •''••« 
Monroe, LA 71 209, USA. 

William D.Reese 

Biology Department 

70504-2451, USA 

2 Parish in northeast Louisiana are apparently the 
first records for the state. Specimens of Alopecurus myosuroides Huds. and Sclerochloa 
dura (L.) Beauv. were collected from the same field on the same day. Neither species is in 
Allen (1993) or Thomas and Allen (1 993). Sclerochloa dura (L) Beauv was reported for 
Louisiana by MacRoberts (1 977) and then excluded by Allen (1 980). Brandenburg et al. 
(1991) did not find any records for this species from Louisiana but did report it for all 
three adjacent states: Arkansas, Mississippi, and Texas. Alopecurus myosuroides Huds. is 
listed by the USDA-NRCS (1 999) database from adjoining Mississippi andTexas as well as 
Alabama, Kansas, New Mexico, and most east and west coast states. 

s Huds): LOUISIANA.Morehouse Parish: F 

section of Sunshine and Tucker roads, E side of Tucker road about 0.5 i 

mi S of Sunshine Road, fielc 

fallow through the winter and not flooded, scattered throughout the fi 

eld and along the edges,27 

Apr 2000, Saichuks.n. (LAF.NLU). 

Voucher specimen (Sclerochloa dura (L.) Beauv.): LOUISIANA. Morehc 

>use Parish: Rice field area 

on the Harold Tucker farm, about 7.7 mi due S of the Arkansas/Louisia 

section of Sunshine and Tucker roads, E side ofTucker Road about 0.5 i 

mi S of Sunshine Road, fielc 

fallow through the winter and not flooded, growing along the field ro 

ad, 27 Apr 27 2000, Saichul 

R i in DM JR[ Hid IW In i 1 991. Hard grass {Sclerochload 

the United States. Sida 14:369-376. 
MacRoberis, D.T. 1 977. Additions to the Louisiana flora. Sida 7:220-222. 
Thomas, R.D. and CM. An i n. 1 993. Atlas of the vascular flora of Louisiana/ 

ferns allies, conifers,and monocotyledons. Louisiana Department of W 

eries, Baton Rouge, LA. 
USDA,NRCS. 1999.The PLANTS database (; 

I lata Center, Baton Rouge, LA 70874-4490 USA. 


Guy LNesom 

North Carolina Botanical Garden 

Coker Hall CB 3280 

University of North Carolina 

( hope! Hill. NC 27599-3280, U.S.A. 
Two collections extend the known range of Gamochaeta simplicicaulis (Willd.ex Spreng.) 
Cabrera into Georgia. This native South American species was previously first reported 
from North America in North Carolina,South Carolina, Florida, and Alabama (Nesom 1 999). 
An additional collection from Florida confirms its presence in that state; the previously 
cited Florida specimen also was collected in Walton County by H.A. Davis. These new 
records are uneguivocal in identity. The hospitality of the staff at BRIT/VDB is greatly 

Voucher specimens: FLORIDA. Walton Co.: 4.5 mi S of Freeport, by overflow pond, 12 Jul 1972, 
Davis 16204 (VDB). GEORGIA. Bulloch Co.: wet pine woods 1 mi S of Statesboro, 8 Aug 1 975, Krai 

56228 (VDB). Wayne Co.: longleaf pine-turkey oak sandridge by US 301 at 5 side of Jesup, 8 Aug 
} '- ■ . r *20 (VDB). 

Nesom, G.L. 1999 CjiJ' ' > ^t r i i i i if 1 -n-i in four southeast- 


Bobby J Ward 1 999. A contemplation Upon Flowers: Garden Plants in Myth and Lit- 
erature. (ISBN 0-88192-469^5, hbk.).Timber Press. Inc., The Haseltine Building, 1 33 
S.W. Second Avenue. Suite 450. Portland,OR 97204-3527, U.S.A.(503-227-2878,503- 
227-3070 fax; $24.95 hbk. 447pp. Line drawings. 

This work represents t t to i i ii i i u n II n i into the subject. Not only does the 

author describe the i i i n i t i i | I ml I th r return n im h 

also relates them v\ her I t nit I I i I i it th mological accounts of 

scientific names and i uninion names., he narrates mylhs and leuencts and cites poetry and prose 
referring to the species from the works of authors throughout the range of Western literature. Well- 
indexed by authors nam. mi |ini ||i i hu tin i II i >mmon), this work be- 

1 1 I nVV I if 1 999 Douglas Chandor: An English Artist and His Texas Garden. 

(No ISBN, hbk.) Antler Press, 31 7 Cleveland Ave., WeatherfordJX 76086, U.S.A. (No 

price given), 1 73 pp., b/w photographs. 
Douglas Chandor pami | n i ilt i hi i r t I i m tycoons and society 

matrons.and in the rocky ground of Weatherford.Texas, he created an extensive garden that even- 
tually rea< hed national mcoqn it ion. "Whim Shadows," later ( hamioi can lens, opened to the public 
in the 1940s. Eclectic in style, the gardens employed principles of English landscape gardening 
with many Oriental fe^tm In In | in I, m in , ,atd Cod gave me the talent to paint 

!) Chandor and Kuteman Family Matters;3) 
md the Garden; 5) Public Response to the 

■: omenta. 




n, in the LIS. 

1 )<iuijlas eino In 

a (. hamlou, 


IThe Studic 

rials; 7) i handod 

s Garden 

Vaughn's Brocln 





)l Ii u i» ) ,i) mi tirnl.erf n m ,t 

hbk. 128 pp,271 color photos, 14 b/w illustrations, 1 map. 
From the close-up m 1 |l t | ,| i i tMllinUip. the 1 

color photographs by Ron and Margaret Mackay are enouoh to make vou want to grow orchids or 

hum thcdiisijackci. "A thoroughly revised and updated version of Millar's earlier book on 
the same subject Mill it i th l t it \ \ t' it ul t it tee the German occupation of New 
Guinea ended more th in ■ , ,t , i mhiI t S n t It i e m ii on hid thishool- 

is a truly personal account of the author's experience with the orchids. Decades of study— obses- 
sion? love?— of the orchids native to this geographic d it t n i I I, i produced an incredibly 

account L t II t I , I - t j I r I hi r < 




Cheryl A.Lawson 

The University of Oklahoma and the State of Oklahoma lost their most distinguished 
and renowned plant taxonomist on May 23, 1999, with the death of Dr. George J. 
Goodman, Regents Professor Emeritus and Curator Emeritus of the Bebb Herbarium. 

Dr. Goodman was born November 5, 1 904, in Evanston, Wyoming, to Arthur and 
Elizabeth Jones Goodman. As a young girl his mother had come in 1 886 to this country 
from Wales, and Dr. Goodman would proudly point out that half of him was a first-gen- 
eration American. A ranch, located some twenty-five miles south of Evanston and home- 
steaded by his Grandfather Goodman in 1 883, is where Dr. Goodman spent the first five 
years of his life. His love of the outdoors and the West thus began. 

After graduating from Evanston High School in 1922, Dr. Goodman, who had no 
plans to attend college, worked as a ranch hand,as a clerk in a drug store,as a sacker in a 
grain elevator,as a salesman of pianos and phonographs and of contract printing, and as 
a compass man for the U.S. Forest Service in the Kaibab of Arizona. About 1925 and 
during the time he was delivering groceries in Ogden, Utah, Dr.Goodman's father bought 
another ranch which was located due south of Evanston a mile over the Utah line. Dr. 

occurred that would begin his botanical career which lasted nearly three-quarters of a 

Dr. Edwin Payson,a botany professor from the University of Wyoming who had done 
his graduate work on the Cruciferae at the Missouri Botanical Garden under Dr.Jesse M. 
Greenman,and his wife Lois came to Evanston to find someone to take them up into the 
Uintah Mountains to collect plants. It was there that they learned of George Goodman, 
who by that time had already climbed several peaks in the Uintah Mountains.They made 
their way to the Goodman Ranch and camped there for a week or two. Dr. Goodman 
took them up to Stillwater Fork and Hayden's Peak where Sereno Watson had collected 

Impressed by the young man who became their guide, the Paysons suggested to 
Dr. Goodman that he should go to college and offered to give him a room for the year 
and to try to find him a job on the Laramie campus. After a discussion with his parents, 
a year of college couldn't hurt, Dr. Good man agreed to go.While still camped 
? Goodman Ranch, the Paysons wrote to Dr. Aven Nelson, botany professor and 
irium curator, and told him they had someone they thought would make a good 

i was. He didn't ask, but learn he did! 

The Medicine Bow Mountains, located just 

ite collecting site during the three years and a 

his bachelor of arts degree with honors in botany. It was to these mountains that he 

would return many, many summers throughout hi ifi Vhether h< was there to teach 

In 1929 Dr. Goodman received a Rufus J. Lackland Fellowship from Washington 
University in St. Louis to do graduate work at the Missouri Botanical Garden under Dr. 
Jesse M.Greenman. His new roommate and soon-to-be closest friend was C. L. Hitchcock 
(Hitchy) who would become professor of botany and curator of the herbarium at the 
University of Washington, Seattle,and a major contributor to the Flora of the Pacific North- 
west. Lured by Eriogonum and Lycium, Goodman and Hitchcock headed out to collect in 
the West in the summer of 1930. 

This trip which began in June followed Route he ihrouaL southern Missouri, Okla- 
homa, and the Texas panhandle.This was the first time Dr. Goodman had been in Okla- 
homa, and while no plants were collected until they reached Portales, New Mexico, on 
iu '" M h« i muni rod ring in Oklahom icuriou licotyh I >nous plant with flow- 
ers that looked like an orchid I Ian > i h latei tftei coming to the University of Okla- 
homa, he learned that he was looking at Krameria. 

The Goodman/H 
were collected and later divided into sets which were then sold mostly to large herbaria 
throughout the country in order to fin, hh e the trip. Sites such as the Chiricahua Moun- 
tains of Arizona and the Abajo Mountains of Utah had been infreguently collected. From 
thru i ejections a few now taxa lesullrd. 

throughout their lives I litchy continued to lease George Goodman about events 
which transpired duiii 'r, i i nm null J t f 1. A'erde Dr.Goodman expounded 
lengthily to two gentlemen on wheie to tuiv the c heapest gas in MonumentValley.One 
of the men proved to be the vice-presideni of Bethlehem Steel! I itchy never did agree 
with Dr.Goodman on whethei the\ had sent a mountain lion o< a .o J| in the Kaibab forests 
in northern Arizen i [ i 1 tin I h Int i I il it ( s there were wild. 

The two repairs d in i i i v i nim - iouo :> nl m Or tin nl their 

panel truck, lived on strawberry jam, and had a lifetime's worth of fun. Even as late as 
1 992 Dr. Goodman clearly recounted for me the events of that trip. 

Dr. Goodman completed his doctoral degree at Wasningion University and the 
Missouri Botanical Garden n i 933 and came once again to Oklahoma, but this time as 
an assistant professor of botany and curator of the herbarium at the University of Okla- 
homa, Norman. However, a job offer in 1 936 from Iowa State College took Dr. Goodman 

During three of the summers in Iowa Dr. Goodman ha< 

a. He and Hitchy had seen these mountains in the distance as they drove north to- 
d Mexican Hat, Utah. Very few botanists had by that time collected in the Lukachukai 
s. Among those who accompanied him was Lois Payson,who had been instru- 
mental in beginning Dr. Goodman's botanical career and who had been widowed in the 
spring of 1927. The group camped near Canyon de Chelly and collected on the Navajo 
Reservation. Lasting friendships developed with the wonderful Navajo.On one occasion 
at least they were invited to an Indian ceremony, which lasted the entire night.This asso- 
ciation with the Navajo people made a lasting impression upon Dr. Goodman. 

Fortunately for those of us who came later as students to the University of Okla- 
homa, Dr. Goodman accepted in 1 945 the offer from Dr. George L. Cross, a former col- 
league and close friend in the OU Botany Department and by then president of the 
University of Oklahoma, to return to OU and the Bebb Herbarium. During his tenure as 
curator,Dr. Goodman builtthe collection from a few thousand plant specimens into one 
that before his retirement in 1 975 would amass nearly a quarter-million specimens.To- 
day the Bebb Herbarium houses the world's finest collection of the flora of Oklahoma, 
along with strong holdings of the flora from surrounding states,the southwestern United 
States, the Great Plains, and Mexico. 

The University of Oklahoma recognized Dr.Goodman's exceptional contribution to 
the University and to the Bebb Herbarium by appointing him Regents Professor in 1 967. 
He was also awarded the Distinguish Service Citation,the University's highest honorjn 1 978. 

Dr. Goodman possessed not only a lively intellectual curiosity and an exhaustive 
knowledge of plants, but also a warmth and cordiality that would make the herbarium a 
place where colleagues and students gathered over the years for meetings, parties, and 
informal coffee and conversation. 

During his career as a botanist Dr.Goodman came to be known as a leading expert 
in the field of plant taxonomy for Oklahoma and the western United States. He authored 
seventy-three publications, described thirty-six new plant taxa, made eleven new com- 
binations, and had four plants named for him. 

Dr.Goodman was a charter member of the American Society of Plant Taxonomists, 
the International Association of PlantTaxonomists,the Society for the Study of Evolution, 
the Southwestern Association of Naturalists, and the Colorado-Wyoming Academy of 
Science.lnaddition,hewasamemberofPhi Beta Kappa,Phi Kappa Phi.Phi Sigma,Sigma 
Xi,and the Oklahoma Academy of Science. He received the Phi Sigma Ortenburger Award 
and the Oklahoma Academy of Science Award of Merit in addition to the Distinguished 
Service Citation from the University of Oklahoma. 

I was Dr.Goodman's last graduate student. We continued to work side by side both 
in the field and in the herbarium for nearly a quarter century after his so-called "retire- 
ment." During the field work for our book, Retracing Major Stephen H. Long's 1820 Expedi- 
tion: The Itinerary and Botany (OU Press, 1995), we traveled over 10,000 miles through 

Nebraska, Colorado,New Mexico, Texas, and Oklahoma. Our field trips in ihcWesi follow 
Working as I did with Dr.Goodman was an incredible experience filled with indelible memo- 

Dr. Goodman was as enthusiastic about life as he was about plants. Once when 
asked how he would describe his life, Dr.Goodman replied/It's been a blast!" It, too, was 
a"blast"for those of us lucky enough to have had him in a part of our lives! 

In August, 1 999, Dr.Goodman's ashes were taken by Marcia, his wife of fifty years, his 
daughter Sula Grace Henrichsen,and other relatives and friends to the vicinity of West 
Glacier Lake near Centennial, Wyoming, in the Medicine Bow Mountains.The circle closed, 
and he was back for perpetuity in the mountains he loved. 


NOVEMBER 8, 1908-SEPTEMBER 13, 2000 


Anne S. Bradburn 

h,hi>\ u ■' i- ,,M 

Dept. of Ecology and Evolutionary Biology 

The History of Natural History has lost another champion. Although she was most often 
in the background, those of us who were fortunate enough to know the Ewans were 

meticulous researcher spending days tracking down the most minute details. Hers was 

Born near Saskatoon, in Saskatchewan, Canada, Nesta with her family moved to 
California where she attended the University of California at Los Angeles, earned a de- 
gree in botany in 1933, and met another young botanist, Joseph Ewan,whom she mar- 
ried in Reno, Nevada in 1935. They produced three daughters, Kathleen Harris of 

enjoyable, Dorothy Nemecekof Chattanooga, Tennessee, and Marjorie Ewan who now 
lives in Albuguerque, New Mexico. All share their parents love of nature, if not book col- 
lecting, and have happy memories of family camping trips.There are five grandchildren. 

During their next 64 years together the Ewans also produced nearly 500 titles. Be- 
ginning in 1 963 with "John Lyon, Nurseryman and Plant Hunter, and his Journal, 1 799- 
I814"published in theTransactions of the American Philosophical Society (n.s. 53: pt.2.), 
Nesta was frequently listed as co-author. Her participation was more widely recognized 
after"Ewania:the writings of Joe and Nesta Ewan"was published in 1989 (The American 
Botanist, Booksellers, Chillicothe, Illinois). 

During their 40 years at Tulane, Nesta was famous for providing fabulous Sunday 
brunches to generations of graduate students. David White, now Professor of Biology at 
Loyola University, fondly remembers devouring stacks of buckwheat cakes loaded with 
jams and jellies, a tradition which continued after the Ewans moved to the Missouri Bo- 
tanical Garden in Saint Louis. 

Once established in the old Museum building where they had palatial quarters and 
for the first time ample space for their burgeoning library, Nesta told me "When I die I 
don't have to go to heaven, I am already there."Joe and Nesta were both great lovers of 
plants,animals,and the great outdoors, and they reveled in the morning walks through 
the splendor of the Gardens. So while we at Tulane missed them sorely, we were glad 
indeed that they had found a wonderful place to continue their studies. 


Richard V.Francavigl a 2000 The Cast Iron Forest: A Natural and Cultural History of the 
North American Cross Timbers. (ISBN 0-292-72515-9,hbk.;0-292-725l6-7,pbk.). 
Univ.ofTexas Press,RO.Box 781 9,Austin,TX 7871 3-781 9 r U.S.A (51 2-471 -4032). $45.00 
hbk., 24.95 pbk. 269 pp., 1 5 color and 52 b&w photos, 8 maps, 1 2 line drawings, 
6" x 9". having a lift ut it n i in i ! h n i tu > i m ,\ it mo tho than nt. t 
including the author. While iho book is an obj< liveanah illusion il map i, el Hum i 
notni< statistics, and census record many pa i, i il mi n n nal fascination with 

the Cross Timbers Mi li ii I i ilii ihm t ,| I ,r rl nrersity of Texas at Ar- 

lington. Like many pro* mnl in I nth i n Cmr it I i h- hi h< en nourished by Texas 
pride and now finds that his roots have grown deep into the Cross Timbers sandy clay. 

Francaviglia begins th I kbycicv I >| it iih con pt o u In I i from i e. pei 
spectives:"The Cross Timbers... are a forested archipelago largely surrounded by a sea of prairie. 
Centered roughly lit t n I nl > n nlio tl r in I i t qetation comprises 

generally north-south trending holts of scrubby oak trees." Next, "...the Cross Timbers typically 
appears as dense stands of post oak and blackjack oak trees that rarely exceed about thirty feet in 

1 nil 'i i" < im hiu i n nth ofWaco along 

'• I'-ll O'i ].| 


tinOS |: 

oi ihor Ll j| ( ,ninihi Ih, M hi > urn is i , ,n, in h I i. in I 
explorers, and Anglo settlers and farmers, and Imally the <ontemporary urbanites.This is the rea 
strength of the book and a significant contribution to the non-technical literature. In so doing 
Francaviglia is able to trace the perceptions through 5- and 1 0-year intervals. Use of these histonca 

I it.i ii. i e, hi ih ii I nt. | i t it ion- n h\ , - ii hi i ti hiu, eie oi weie not add. d to map 1 

ami diaries, and leader, will want to keep lliat in mind. However his nueipretations appear to be 

i ni it le in 1 i I it i nil nl i I ii ,1 . i I ; I i in "ie argument that pre 

As a reader not inteiosted so muc h in the histonca aspects, I bogged down in some of the 
fact, the author appeal • i , hum m hit n the details and intro- 

The a 

and increasing con 

servation awareness 

, Even 

regional literary work' 

reviewed. One of thi 

>k is, in the last chapter, 

? of the topics covere 

:d and a six-page sun 


of the author's interpre 

Dennis W. Woodland. 2000. Contemporary Plant Systematics,Third edition. (1 -883925- 
25-8,hbk.). Andrews University Press,21 3 Information Services Bldg., Berrien Springs, 
Ml 491 04-1 700, U.S.A (61 6-471-61 34, $64.99 hbk. 
560 pp., numerous b&w photos and line drawings, CD of over 4,700 color images 
Contemporary Plant Systematics flows from Woodland's teaching experience to fill the needs of his 
own students. His goal in writing a text is to provide"a well-illustrated, broad-view, beginning text 
that would give the students, wherever they may live h I h , i nt botanical understand- 

ing of vascular plants that would utilize the changing world of global information." In some ways 
he has succeeded and in others he has not. 

The book is divided roughly into three portions. Printed page tabs, which mark the chapters 
and other divisions, are a helpful innovation. The first four chapters cover an introduction of sys- 

second section (5 chapters) is by far the largest. It encompasses an extensive survey of pterido- 

special characteristics and terms.The last 

six chapters cover more advanced topics: history of the 

vascular plant groups;a survey of morphological, ch> n I I - n m conservation issues; 

and the role of botanical gardens. 

The field and identification portion 

(early chapters) is thorough, but the other chapters on 

iple,speciation is covered in one page and one diagram. 

Likewise,in the discussion of phenetic and 

dendrograms and cladograms are illustrat 

An appendix with simple four-taxon prot 

)lems, mathematical algorithms, character and distance 

ae very helpful. 

Woodland is to be congratulated fo 

loptc emtio hi, ill - i i nun din.i.i, mu , n tin. i 

ing of organisms. Conservation is rarely addressed by texts in plant systematics but is an endeavor 

that systematists ate m mi il l~ In r i i r it 

i during their careers. His survey of botanical gardens is a 

nice addition, but many instructors woul 

d rather exchange this information for more detail on 

Lersten on plant anatomy, Rolf Sattler on morphology, 

Loren Rieseberg on molecular systematic 

s, Cliff Crompton on palynology, Peter Holland on ecol- 

iouiteon families of (the, 1 / le^ oy ni-\M i ) lentospoianqiate homosnoious fcins am neater 
thediawinqs nm I he dim r if n ions are espec lallv helplul to the in>vi< e in dilietenii.jting ; 
ni/ing any panic ular segregate family. In a ceitain extent, the included CD of the Unix 
iseonsm ' ' Atla ' the Vascuiai 11 1 4 ptorth ieti n le hul I f un :t 
face to be less tl 

There is a numbei of aim < inn mm 1 pi I 1 iln impl 1 t< d hi ie < it In 1 lei 
collected from the second edition or were introdiu ed with the ihrd). I lie haidbac k hindi 
in ilil , iml 1 ' . il ■ il m hold cm well . ith m| in, I use. [he 1 ,| ' 'tin 1 c 1 , nil 
lor example, on page 410 one finds "from a wide spectra mc] of sources"and on page 4 

nomic hierarchy, the Maqnolimeae is listed as the suboider under the Asteralns and al 
Asteraceae! A num e 1 > in 11 1 1 11 1 1 | mint m u m 1 

In an apparent 
11 Ibid 1 II - , < , 1 ' I 1 1 1 1 ett plate of Zom/f 

was published il iq. 2b), is not 1 ited as the source oi the original lorn whu h ihe illust 

i' mi u i nt t 1 fqionu i ml ne 1 lb 1 n e nmpil b in ,t In 

Contemporary Pliii'l ml I b nu 1 1 m m u . n il 1 u il ui 1 1 in 1 ill 

qlossuiv of m r h h 1 il tn wee mmlaoed with 1 shorn il 1 m 1 f 1 f 1 

famines and it the book weie designed to use a protossmmafy bustmtod companion arlat 
Wendy Zomlek 1 q , t 

ager, Botanical Research Institute of lexw,. 

Man and J.M. Saravanan. 1 999 Pollination Ecology and Evolution in Compositae 

(Asteraceae). (ISBN 1-57808-058-4,pbk.).SciencesPublishers,lnc, P.O.Box 699,May 

Street, Enfield, NH 03748, U.S.A. $49.50 + shipping. 166 pp. Line drawings, tables,and 

y thorough account of our c 

ost complex family of flowering plants. I he book begins 

research on pollination in general and on Compositae in 

that are commonly ton id u tlm n,v ,e , n the Composite family 

considerable dama t« 1 t t , r j I h 1 m > h ( ter I 
specific parts of the composite capitulum, anc 

out the 'importance of the structural differences of these organs, and how pollinators have influ- 
ences the evolution of those differences. Chapters on nectaries and pappus complete the discus- 
sion o' I he specializations found in cuin[ < • puuLi i'.'h ed in with the chapters on each of the 

floral organs is a section on sexual j> I, ' \\ i tl, in me Compositae. In this author's opinion, 
ih s chapter is a bit misplaced, an the overall structure 

chapter is well written and well referenced. The final three chapters bring all of this information 
together with a well-presented discussion of floral biology, pollination and evolutionary trends 
within the family.This book is very well written and thoroughly researched. It includes a glossary of 

learns which is guite useful, espei all-, t r il t familiar with terminology specific to the 

Compositae. The entire volume is well references, with nearly 450 citations contributing signifi- 
cantly to the importance of this worl uld i nimend thi < >lun in to all students of the 
Compositae, and to anyone who i nt u i n| II i n biology in general.— DebraTrock. 

Alan Hopkins (Editor). 1 999. Grass: Its Production and Utilization. Third Edition. (ISBN 

0-632-0501 7-9, pbk.). Blackwell Science Ltd., 350 Main Street, Maiden, MA 02148- 

501 8, U.S.A. Distributed for Blackwell Science by Iowa State University Press, 21 21 

South State Street, Ames, IA 50014-8300, U.S.A.(Orders:800-862-6657, 5 15-292-01 55; $49.95 pbk. xiii + 440 pp., numerous figures. 

It has been a treat to review this book, and here is why: for many years I was a biology professor at 

Kansas State Univers ) notably grassy state.There I was 

familiar with the native grasslands, plus the managemrni and utilization of grasslands for many 

purposes. The book under review it i| n Briti lands and it offers a different dimen- 

M( tilt nllH II h ! I I I II Nil I I 1 I Ul I ) 

pages, each with a several page list of references. Most of the cited papers are from the past de- 
cade. There are chapters on pastures, herbage production, weeds & pests in grasslands, feeding 
values of grass (50 pages), conservation, Iunh" .mo in, ing behavior, landscape and wildlife, etc. 
I here is no coverage of grassland systematics. Two chapters drew my attention; one on sward 
estaolishmentand renovation, and th- rh i i i in ,, land. The climate of Britain comfort- 

acy supports lawns of great expau i ' n I i m iihiuilln nhl< u< ot decorative purposes 
(amenity grass), and the approach to lawn and turf management is rather different from what we 
are at c ustomed to. The introductory chapter is an absolute gem for summarizing the thrusts of 
nook and for supplying a grass-oriented land-use map of Great Britain. 

How useful would this bool- ' the 

applied ecology and physiology of me whose studies 

focus on intensely managed gra c Lin i lit il i n hi h i ,i i ularlv valuable, for they 

are lamely from the Europi mlit-niur- -- it i nl i- - I Americans— Theodore 

ite of Texas. 

Cm II iiii.hkjBm I i * 1 i999.The New Central Texas Gardener. (ISBN 0-89096- 

871-3, pbk.).Texas A&M University Press, College Station, TX 77843-4354, U.S.A. (409- 

845-1436,409-847-8752 fax). $24.95 hbk., $14.95 pbk. 204 pp., 4 pp. color plates, 

b/w illustrations. 

This "basic gardeninq t t r m r it i il m i Innil updated version of the 1980 publication 

I >v Ml i I I t t 1 1 i 1 1 i | i 

advice about landsi j[ in i I- i ' itin.ti n - iM* lf , ,| , | Tint selection; and gar- 
den purines that work." 

V vini nditi n Mhi it n i itut i i u A ntr is a chapter on 

1 I ih W M i h in In I i t 1 i ti ii I (i I ,iii, fin nt and information 

and ponds are briefly described and discussed. I lh i tut < immi includes maps, ink drawings of 
plants by Kate Bergquist, mini i 8 |-u, se< turn of color photographs. A month-by-month calen- 

dm Ms inintinq i. ntn ludes the book. 

\hh hi ih id - in I ten nn it< in n I . i in d i ndi in i tli hit i /ill find it somewhat 
» Ii m mmM hik tl nihil Ml tl hi lit m th i nit ntral Texas gardening 

RoblrtW. Kiger,Cmm in AT i iM.m.ln MM. t 1 "99 Index to Scientific Names 
of Organisms Cited in the Linnaean Dissertations Together With a Synoptic 
Bibliography of the Dissertations and a Concordance for Selected Editions. 

IM5N0 M iMb-r, t,hl ) Hunt In MM. lot Bol muul I wnim M ition ( nn |i« 
i II ti IMi i i! Fttt Ininih PA 1 I m [ , Mi n 300 pp 6 b/w 

figures, 8" x 11". 

I imvieus himself. I he c 

Ihem is Mill) 

loqmphii i nl.-iloq M thenriqinal 


ons, plus a succinct overv 





an expLinalion n! tin' Men numbeis, vvhn 11 , 

The dis: 

the texts of all represent 

1 n "i, t if ioli t - on / In 

Piun.i in inn ,l 

1 ol i I I'lf - ' l h i^t- Ik i 

)M n In In' ,| i ■ 1 ii .in iin-n F it 


i - -ti- Ml 


omfortably accessible to 

nksand good wishes. Mnv thev i. 

their work. — Theodore A/1. 


tute of Texas. 


e CailletJ.Farron Campbell, Kevin C.Vaughn, and Dennis Vercher (editors). 2000. The Loui- 
siana lris;the taming of a native American wildflower. Second Edition (ISBN 
881 92-477-6, hbk.). Timber Press. Inc., The Haseltine Building, 1 33 S.W. Second Av- 
enue, Suite 450, Portland, OR 97204-3527, U.S.A. (503-227-2878, 503-227-3070 fax; $34.95 hbk. 254 pp., 1 1 1 color photos, 5 watercolors, 1 1 
b/w photos, 14 line drawings, 1 1 tables. 

n Apogon) belong to series Hexagonae of the genus Ins in 

ises.The editors 

— 1) History of Louisiana Irises; 2' ( la: sifii ation and Species;3) Description of the 
•) Collecting of the Species and Natural Hybrids;5) Propagation of Louisiana lrises;6) 
ridizing to 1988; 7) History of Hybridizing since 1 988; 8} Tetraploid Hybridizing; 9) 
lybridizing; 10) Culture; 11) Landscaping with Louisiana Irises; 12) Flower Arranging 
Irises; Appendix A, Society for Louisiana Irises; Appendix B, Popular Cultivars of Loui- 
ipendix C, Mary Swords DeBaillon Medal; Glossary; f 

Stanley L Bentley. 2000 Native Orchids of the Southern Appalachian Mountains. (ISBN 
0-8078-2563-8,hbk.;0-8078-4872-7,pbk.).Univ.of North Carolina Press,P.O.Box 2288, 
Chapel Hill, NC 2751 5-2288, U.S.A. (800-848-6225, 91 9-966-3829 fax;www.uncpress. $39.95 hbk; $24.95 pbk.xviii + 256 pp., 1 1 9 color photographs. 
Another orchid book. Well, maybe not! This is quite a guide filled with great photography of 52 
species of orchids found in the mountains of southern Appalachians. That is to say, a region en- 
compassing western Virginia and North Carolina and eastern West Virginia, Kentucky, and Tennes- 
see. This book is quite a treat for you amateur naturalists or professional botanists who happen to 
live, vacation or botanize in the southern Appalachians. Each species is provided with a scientific 
name and common name(s), a description of the flower (color, shape, and size), information on 
flowering time, distribution and typical habitat. The author offers much advice when hunting for 
orchids, but one piece of advice I thought was most interesting was to carry binoculars. Have you 
ever used binoculars to look for orchids? Well, you just might want to consider it according to the 
author/Time after time, when I have spotted a plant high on a bank, well past the 'telling for sure' 
point, binoculars have saved me from having to scramble up the bank to try and determine its 
species. As the years go by, I appreciate more than ever the steps saved by my trusty binoculars." 
Contents— Preface, Acknowledgments, Please do not Dig Native Orchids, Introduction, What 
is an orchid?,Rarity among Orchids, Looking for Wild Orchids,Special Orchid Places in the Southern 
Appalachians, Preserving Wild Orchids and Their Habitat, Using This Book, List of Native Orchid 
Genera of the Southern Appalachians, Glossary, Bibliography, and Index. —Barney Lipscomb. 

.Pi in and John P. P,au 2000. The color Encyclopedia of Daylilies. (ISBN 0-881 92- 
488-1, hbk.).Timber Press. IncTheHaseltine Building, 1 33 S.W.Second Avenue, Suite 
450, Portland, OR 97204-3527, U.S.A. (503-227-2878, 503-227-3070 fax; 

I t In h 1 i hh t hi In 1 Ml - i ii | In 1 i, In iii I it imp. tionol tl 

m in i iipt T« II n 11 i I ii ii | 1 n i h- i n I i t botanyand hortia 

till 1 i ih I I I ' I I LI Ii | h I i l| Ml II Ml I II M il I 

the world combine too ii ih in i ornj m< ■hun nl Midaylili. t ) i il. h 

other book so completely catalogs ihe woild of daylilies." 

Contents.— Foreword; Preface and Acknowledgments; 1 ) The Daylily Plant; 2) History of tr 
Daylily;3) History ot I II H il i 4) i 5) i i II ii m i nellies; 6) Doubles;' 

I lybi idi/ing; 1 1 ) Cultivation; 1 2) Daylilies o 

Pc-ilr Goldblatt and J i mm M Wildflowers of the Fairest Cape. (ISBN 620 

24787-8, pbk.). Red Roof Design cc,Cape Town, SOUTH AFRICA. Available fromTim- 

ber Press. Inc.Jhe Haseltine Building Hi VV H m r u nie 450,Portland, 


hbk.,8 1/4" x 11",315 pp., 663 color photos, 1 color map. 

"OH Lord, how manifold aie ihy woiks! in wisdom hast thou made ih.Mii all: the earth is full of thy 

riches."— David, Psalm 104 '4 \n<t thefa rest < ape (Cape of Good Hope) has its share of botanical 

riches! No wondei it i t u < i i m> 4 I i m I i h i n t the earth. I don't think 

anyone will argue | n h< i in ni i i I n in f'otancial Garden, St. 

■.:i. in no! iViaiininu i National h.Mauk a nsi [iile,Hnilh Ala. a) Uolaii :> : <2 of tla most oomm.m .n 

(onspiiuou wildflow i ml loin lilt i in ih ler I lime are presently in 

(.nit ition ()n. iiii ith Ii i i n I I I I I i I i ill gasp at the beauty. 

Well done thou good and faithful servants! 

i o 4 it Pn t i e the i jp. 'i n I i I In I i - I I .it I II t I ii 1 

Cmiu i ■ . h V, ildtlowers Routes:TheCapePeninsuL the We* m a. Bokkeveld and Hantam, 

TheOlifants Rivet Vail, m in-lih. lainheils! n mim h< M uninin IH v>\ Ihe South Coast;How 
to use this Book; Wildflower Sch( dul 1» il ' i n I ' tl . I nnilie, lllu ti i 

Aa an. I Nam. ■, Ma, m. 2000 Mosses and Other Bryophytes. An Illustrated 

Glossary.(ISBN0 4 \ )6 50-7,hbk.) no Optics Press,Box320,Nelson,NEWZEALAND. 

Available from Timber Press In I . I ml Jmg, 133 S.W. Second Avenue, 

Suite 450, Portland, OR 97204-3527, U.S.A. (503-227-2878, 503-227-3070 fax; ^ m s> I, I I ^ 8" x 8 1/4", 220 pp., 968 color photos, 22 

ine drawings. 
:olor glossary covers mosses, liverworts, and hot 1 1 t t iin 1 ' > the authors/The three 

as were chosen b' 

milarenough that a term which apfiln .too,,! hi n i\ \ -ties to the other two as well. "Nearly 
photos— representing nearly 400 species— v,m> t ?l n it an >u I- els of magnification, 
ly with a microscope. A scale bat i |i i- i nl li t h t , n Tin authors have in- 

-referenced terms used elsewhere in the glossary. Terms in the glossary are printed 



i poini 

: somewhat longer than a 


and cuspidate, ending i 

er).More than one illustrati 

vided for terms that desci 

variable structures sue 


finitely be useful to students anc 


that often 

are st 

udied in the laboratory. 

i rW i hi 2000. Tropical Ornamentals: A Guide. (ISBN 0-881 92-475-X, pbk.;0- 
881 92-448-2, hbk.). Timber Press. Inc., The Haseltine Building, 1 33 S.W. Second Av- 
enue. Suite 450. Portland, OR 97204-3527, U.S.A. (503-227-2878, 503-227-3070 fax; $34.95 pbk„ $59.95 hbk. 542 pp. 458 color photos, 50 line 
drawings, 6" x 9". 

: ;•_-, no doubt this honk is about mostly tropica! unc-mn-nidls and a few others. iV'anv of tlv: 

Sotidago nemoralis, etc.). The 400 plus plants treated in this book are arranc 
by scientific name and not common name. You can find the common name in the index. This of 
course allows closely related plants to be placed together. About two-thirds of the plants are in 20 
plant families and are accounted for in an appendix, Twenty Common Plant Families. Each plant is 
organized as follows:Genus and family;Scientific name and authority; Synonyms; Distinguish- 
ing characteristics; and Description. It really is a guide to the plants most frequently encoun- 
tered in gardens of the tropics. The photos are splendid. This is an excellent -eference on tropical 

Contents— Preface, Introduction, Organization of the Information in Tropical Ornamentalsjhe 
Tropical Ornamental Plants, Twenty Common Plant Families, Identification Key, Glossary, and In- 
de> Hume) tipscomb. 

ioyd. 2000. Christopher Lloyd's Garden Flowers: Perennials, Bulbs, Grasses, 

..(ISBN 0-881 92-492-X,hbk.; 0-881 92-448-2,hbk.).Timber Press. Inc./The Haseltine 
ing, 1 33 S.W. Second Avenue. Suite 450. Portland, OR 97204-3527, U.S.A. (503- 
'878,503-227-3070 fax; $39.95 pbk.CAN 54.95 hbk.448 
9 color photos, 50 line drawings, 71/2" x 10". 

man band covering eunihi i i n m ' , n ; Im i< nngl\ i nough tin 

All the plants included h i thin pi nence during a long life of garden- 

1 1 have grown myself at some time i Igui Ion Find vour favorite plant in 

e, then I suppose it is one not grown by 'lie auihoi, suiely that will include a lot of 

he author has learnt, thought, seen tii lib i i il >ut the plant, and that 


).The Illustrated Flora of lllinois:Sedges:Carex. (ISBN 80 
tern Illinois University Press, P.O. Box 3697, CarbondaleJL 629 
5-6633,618-453-1221 fax;$59.95 hbk.328 
1 59 line drawings, 1 59 maps. 

Wow 1 Hums volume M of the llhi.inif. j i loi, ■ > hli<\ w senes, iIh snip in. i List volume devotee 
monocots.The 1 59 detailed illustrations by Paul Nelson are excellent and will aid any person in 
identification of these species. A key to all the species is inc. luded. I he usual stuff is included v 
each species: Genu | w pla I publi t nonymy,d< rip ommon name, habi 

range.and Illinois distribution (.with map of Illinois); this is ioliowed >>v a discussion of the nom 
clature and habitat of the species. 

Irani (he dustjacket.— "Since more than three fourths ol the speues of Latex in Illinois 
inhabitants of wetlands,an rm t th . nn i tit ,1 i , working in wetlar 

Amateur and professional botaniM .veil find th information! tiemeh ikiable, as will envin 
menial ,ind ion gtoups jar den lub iaim bureaus, hoi in extension out oi 
fit H mi Hid li II t i I i i il i i ii i nn I n i t i i in incl in ran and end 

|Mt I | it j r > i it i I tli f ii i ii it nn Mil ii ir u i i essments and wile 

management prnjei ts will also hub the infoi ination per tiruMH." Barney I ipaomb. 

ST. Runkel and DM. Roosa. 1 999. Wildflowers and Other Plants of Iowa Wetlands. (ISBN 
0-81 38-2 1 74-6, pbk.). Iowa State University Press, 2121 South State Avenue, Ames, 

From the Preface - With th. inweasi j awamn. of th. inpuitano »f wetlands, a book was 

newlwl |, . s It, ini u im spei kill i understand ' .-. wetlands aw ■ in- , w a ! i f ■ 
1 1 m n . t wet lam I pi ml I dm aie wondetiul nuniiik books i I i I h i if w • 

so technical that all hut \ i , ilii- i mil ■ u I im [In e manuals r. h." 

trate many users, particularly beginners. We think wetlands are grand place 
them as we do. Hence, we offer this book as a starting place for those v 
about Iowa's wetlands and wetland plants. Plant descriptions are present! 
or aquatic), then refined by habit (e.g. floating or submerged) or by taxo 
and allies ortrees and shrubs). Common names vary throughout the coun 
those in frequent use,although others may be used regionally or locally. \M 
plant's Latin name,along with the authority." 

aylor. 1998. Desert Wildflowers of North America. (ISBN 0-87842-3 -I pbl- 

untain Press Publishing Company, P.O. Box 2399,Missoula,MT 59806, U.S.A. (406- 
i-1 900, 800-234-5308). $24.00 pbk.349 pp.500 + color photos, line drawings, maps. 
ock cover.— "Each ,piin , ,h m i nt II w pm nurns u ,1 n< n |u i i n I i 
of wildflowers transform the drserts of North America into a sea of vibrant color. With 
is and photographs of more than 500 species of flowering plants, this full-color filed 
arough the flora of the blooming desert. 
he book includes an overview of desert ecology, a simplified botanical key, and an illustrated 

Contents. — Preface; Inrro In nil i III ■ I i j Mi ntilvmg Nmii, mm i ml <r 

/ildflowers to Family; Plant Anatorm H in J i |illu it n I] lm ted References; and 

idex. Desert Wildflowers of North America covers the Mojave, Sonoran, Painted, Chihuahuan, and 
m n i i 1 1 I ( i i i t i i i 

ppreciates wildflown pmru ul il in I - n n nmmm The plants are arranged alphabeti- 
ally by family, that is the common name of thp fdmil i mtlm F imilv, Allthorn Family, Ama- 

inth Family, Barberry Tarn um < IM I i ' i i list plants on a desert-by-desert basis, 

Sonoran and Chihuahuan 

Jean Andrews. 1 999The Pepper Trail: History and Recipes from Around the World. (ISBN 
1 -57441-070-9, hbk.). University of North Texas Press, P.O. Box 31 1 336, Denton, TX 
76203-1 336, U.S.A.(Orders: 800-826-891 1,940-565-4590 fax). $50.00 hbk.264 pp.57 
color illustrations, 7 maps, 7" x 1 2". 

1 t - i i I I i i i nth 'prppei ti.nC IF n| > tin tun 

ning addition to Jean A/ di- . I i |- Mrews, who has been called 'the first 

lady of chili peppers, "the godmothf i I ml il II , t i n u yistered trademark 

'The Pepper Lady/follows the spice trade and early movements of capsicums along the spice roads, 
through much of turkey and the Middle East, Africa and Monsoon Asia (India, Nepal, Bhutan, Sri 
lanka,Thailand,and Indonesia) plus » i luanai I u nan pro m China and the Silk Route. 

This latest offering of Andri- u In M | i i in ' Ik! such as the etymol- 

ogy of the word 'cayenne.'" 

"The first spice to I ). u ,< d b\ m,n | < ppu «n nnnil hum o Guatemala, much of 

Part I I 

Notes; Bibliography; Subject Index; and Recipe Index. 

M 'I , ■■ i t .iliiihi ■, m eveu . mi.'d in I now about Peppers 

cipes! 1 didn't count the 


: many are listed for tr 

H' 1 Hi '.'.in 1 il In 




?.s; Breads; Desserts; ai 

jine Chilli Pepper Jelly,; 

y when 1 was raised c 

•n Wild Plum Jelly 

, but who knows? 

Mowering Plants of the Galapagos. (ISBN 0-801 4-8621-1 , pbk .; 

I)-"! I i/IO - hbk ) ( >rn< li I Inivetuh I rcss.Sacjc I louse: 51 I oih .in et iho a 
NY 14850, U.S.A. (Orders: 800-826-891 1,940-565-4590 fax). $59.95 hbk., $29.95 pbk. 
370 pp. 266 color photos, 1 map, 6" x 9". 
mm the back cove i I In J \\ \ -.1 n . I i n. iiuou i ik n! ol unusual plants 

nd uiunals The i lano luinn I iin m ,1 ih. iu otihil. I ' it in n eatch leading to his 
heory of evolution Is no it J el. ion ind thnit nanniticenl flnia md fauna continue to draw 
isitors from around ih .III I nth inh. i". . it 't i .oik and featuring his 

xceptional photoqr ipr \ . ■ • - ■ ; "■ ■ .,,",..! i ih li t ihle and in-depth, yet 

onlenty I on ■wend; Piel,n e; At knowiedgineiits; Oalapau. is Islands M.i[i; I low lo Use I his 
lUide, Illustrated PlantTerms; A Bind Itu- i h ti n i 1 i,a! i| u Lin. i i I nil Descriptions,Plant 
e\ i I i itv Lit. t .in i I | | n i I I I J ! i i I 111 i i t I nit "Tie iU I iii the 

csxt; Appendix .A Selened Visitot sites and Flevveiona Plants I ikelv to he hs. oenteied; and Index. 
Other than |ii I th ( pin 1 1 i |. t t u in ih in . | tin t I mil. v m p| (i nt in "hand, 
oe inn in him mini, id i ih f>| t pn i< no II I. tin II ' ' | n in it I it i n I n 

ae bookin one of fiu ,i i| i ti I nib I tl n. n i I i I ttln live groups is then 

ivided into groups based on how a plant's leaves are typically arranged on its stems. Options 

ided basedonflowo i di Ih hi n hi II i t im piJ i 1 m purple,blue,green, 
nd brown. Once you have made vent selection, them is a poie number that takes you to the 
I action 

arough the photographs until he or she finds one tha 

William A.Weber (Editor). 2000The American Cockerell: A Naturalist's Life, 1 866-1 948. 

(ISBN 0-8708 1-544-X, hbk.). University Press of Colorado, 5589 Arapahoe Avenue, 
Suite 206C, Boulder, CO 30303, U.S.A. Distributed by: University of Oklahoma Press, 
Book Distribution Center, 41 00 28 th Ave., N.W. Norman, OK 73069-821 8, U.S.A. (Or- 
ders:800-627-7377,800-735-0476 fax). $29.95 hbk.352 pp. 7 b/w photos, line drawings. 
From the dustjacket.—"\r) The American Cockerell: A Naturalist's Life, 1866-1948, botanist William A. 
Weber pulls together pieces of the life ofT.D.A.Theo' Cockerell, a man who was an internationally 
known scientist, a prolific writer, and a highly regarded teacher at the University of Colorado in 
Boulder. The elder brother of the noted scholar Sir Sydney Cockerell, Theo labored in relative ob- 
scurity in America while his brothers and their fan n i>- , >■ ili Miih limelight of smart Brit- 

"Despite his alienation from his elite background, he nevertheless became a great teacher; a 
mentor, a kindly artist and writer of rhymes for children, and the greatest specialist on bees in the 
?s is monumental— he catalogued over 

merits.— Foreword by Norma LeVeque; Joseph A. Ewan on Cockerell; Acknowledgm 
I Chronology;Autobiographical Papers; Philosophical Papers; Academic Matters;MiS( 
ife and Habits of Bumblebees; Postcript; Biographies, Obituaries, and; Fndn 

W.H.F 2000. Rain Forest Exchanges: Industry and Community on a 

Frontier.(ISBN 1-56098-983-1, pbk.). Smithsonian Institution Press, 470 L'Enfant Plaza, 
Suite 71 00, Washington, D.C 20560-0950, U.S.A. (Orders: PO Box 960, Herndon, Va 
201 72-0960; 1 -800-782-461 2). $ 1 9.95 pbk. 222 pp. 1 1 figures, 4 maps. 

Fisher argues that decisions to cooperate with frontier industries are best understood by taking 
into account the power of native social systems to shape the acquisition of trade goods. Charting 
the history, politics, economics, and ecology of the regions, he telis how subsistence practices such 
as hunting and gardening have been altered be sedentarization, how villagers interact with In- 
dian-agency and extractive-firm personneLand how notions of barter or sale only loosely describe 
the transfer of goods that take place in the village. In Rain Forest Exchanges, Fisher contends that 
efforts to encourage conservation and sustainable practices among indigenous Amazonian groups 
remain problematic unless, in addition to the influence of the frontier, the dynamics of each groups 
social and economic organization are recognized. — Review forthcoming in Sida 19(3), 2001. 

W.K. Chapman, V.A. Chapman, A.E., Bi ssi in, A.R. Blssi i n , and D.R. Plns. 

New York in Color. (ISBN 0-81 56-0470-X,pbk). Syracuse University Press, Syracuse, 
NY 1 3244-5 1 50, U.S.A. $24.95 pbk. 1 64 pp. 350 + color photos. 

'i i l i > u WiidllowcrsofNtwYork in Color i i field guide iln ill give inline enlhusi- 

and concise descriptions, written in easy-to-follow, nontechnical language. The color illustrations 
have been selected for their scientific accuracy as well as their aesthetic quality.This field guide is 
keyed in a manner that easily leads the reader to major groups based on flower color and other 
physical characteristics. Wildflowers of New York in Color contains descriptions of both commonly 
encountered and rarer, protected species. Included are color illustrations and descriptions of spe- 
cies not seen in other field guides. Nomenclature has been updated to reflect current usage.This 
companion for nature lovers and anyone interested in the wildflow- 

D J Raynal and D.J Leopold. 1 999. Landowner's Guide to State-Protected Plants of For- 
ests in New York. (ISBN 0-9670681 -0-X, pbk). State University of New York, College 
of Environmental Science and Forestry, Syracuse, NY, U.S.A. $19.95 pbk.92 pp.Color 
From the Introduction.— The purpose of this book is to present the key diagnostic characteristics of 
the protected plants that are found in forests of New York State using photographs and brief non- 

hbk. 1214 pp.995 line drawings,2433 range 
From the dustjacket— "Designed as both a reference work an. 
ume contains more than 2,400 range maps and 995 line drawings. Fc 
students interested in an up-to-date treatment of the vascular flora of 

The geographic scope of the work e> 
Island and west to the Hudson River.The"General 
such groups as aquatic plants, vines, and 
herbaceous families, the keys cover flowering 

The "Descriptive Flora" section includes I- 

onous or hallucinogenic properties. 

The distributions of more than 2,400 sp< 
les line drawings of 995 species, showing d 
i in the keys. Near the end of the volume tr 
for dicots and one for woody plants in wintf 
ecimen displaying limited information, or t< 
acteristics for use in teaching." 

The line drawings, although not for ever> 


Rick Darke. 2000. The Color Encyclopedia of Ornamental Grasses on CD-ROM. (ISBN 0- 

881 92-479-2, PC format only.Windows 95 or higher).Timber Press. Inc.Jhe Haseltine 

Building, 1 33 S.W. Second Avenue. Suite 450. Portland, OR 97204-3527, U.S.A. (503- 

227-2878,503-227-3070fax; color photos. 

This is the CD-ROM version of Darke's hard copy reference, Co/or Encyclopedia of Ornamental Grasses. 

h i 1 in n ir - ) i h l | ' i i i i J i i 

cat-tails;selected bamboos are included as well. The CD version includes nearly 24 new plants and 
200+ color photos not found in the hardcover.The photos from the CD can be printed in full color 
for use in design work, plant identification, and nursery signage.There is an Interactive USDA Zone 
Map, a useful "Search" function that allows you to track down plants and plant names, as well as 

d A. Zahler and Edward C. Jensen. 1 999. Conifers of the Pacific Northwest [on CD- 
ROM.]. Oregon State University College of Forestry, Forestry Media Center,248Peavy 
Dept/fmc; Price unknown. CD-ROM. 

in which conifers flourish. Nearly 30 species of conifers grow naturally here, organized into 4 
lies and 1 3 genera. Many other species of conifers have been introduced from other parts of 

A Compatible PCs; 486 DX, SX or greater; Windows 95, Windows NT 3,51 
! RAM; 8- or 16-bit sound card. MACINTOSH; All Macintosh with 8 MB of a 

n Technology category 


Daniel B.Ward and Robert T. Inc. 1 997.BigTrees.The Florida Register. (ISBN 1 -885258-06- 

.\pbkj. Florida Native Plant Society. $20.00 pbk.viii + 223 pp. 
From the Preface.— "Thr f lorn Li Koqotoi document eh) native and .'-1 Minn native trees, for a total 
of 853 listed trees. For cadi individual tuv, inioimaiion is uiven as to its common and scientific 

tion, owner, and nomina: ■ nd t tank i i mp n t the- tat iitmrK] the elite trees of 

tional champions, while 106 o- da champions." 

Contents.— Preface; Foreword; Intiodcctiondnvvntoiv of Big teen , units of Yesteryear; How 
They Died; Rates of Growth; I uiqost and Smallest; Spei ios Banked by I li-i(]hi;The "Average" Cham- 
pion; Large Vines;County hiv f toid I ist;Nominalon. ami Owners; Acknovvli^iqments; Selected Ref- 
erences; Appendices; A. What K a free' 1 B. Me<. hanit s of the Survey. C. Memorandum of Under- 
standing. D. How to N ■ E. . '.' ■ nTree.F. Evaluation 
of the Formula. G. A Case of Volume Measurement; H. William BattmmS Biq Trees; and Index to 

)E. Brown, Frank Rj mi mo and s i[ I i 1 998. A Classification of North 

American Biotic Communities. (ISBN-87480-562-7, pbk.). University of Utah Press, 
179S [ south Campus Drive, Suite 101, Salt Lake City, UT 841 12-9402, U.S.A. (801- 
585-9786, 801-581-3365 fax). $ 1 9.95 text; $20.00 map; book and map set, $34.95. 
1 52 pp. 1 1 8 b/w figures (photos, maps), 8" x 1 2". 

tin ha< kiovei. ACIn \nn ' i in < In i n hi 

ities delineated in the \ 


t of Figures; List of Table 

s; Acknowledgments; Int 


"he Biogeographi- 

'Classification System;- 

l)The Biotic Communiti 

Ml 'I'll Ml 

lerica Map; Plates; 

Teresa Carri ra Caci iOn. 1 999. Orquideas de Chiapas. (ISBN 968-5025-44-4, hbk.). Consejo 
Estatal para la Cultura y las Artes de Chiapas, Polyfotum MrsiMmeticano, < a ede 
Andres Serrra Rojass/n.,TuxtlaGutierez,Chiapas,CP2Q0 to ( s in *ppio hi' 
hbk. 1 96 pp. Color photos, 9" x 1 2". 

delaflor;Partesdelaiolu ntn h if nennTn i, I, ft m i ,, ,, i, de formas;Mapa de 
Regiom f i ioqr iti i II nl I ,, hU umi nil \ i i h familia Orchidaceae 

Daphne Gail Fautin, Douglas J. Futuyma, and Frances C.James (editors). 1999. Annual Review 
of Ecology and Systematics. Volume 30. (ISBN 0-8243-1 430-1 , hbk.; ISSN 0066- 
41 67). Annual Reviews lnc.,41 39 El Camino Way, P.O. Box 101 39, Palo Alto.CA 94303- 
01 39, U.S.A. (650-493-4400,800-523-8635,650-424-091 fax; 
$1 20.00 hbk. 641 pp., b/w illustrations and one color plate. 
Annual Review of Ecology and Systematics for 1 999 is packed once again with a mix of articles on 
ecology and systematics. Volume 30 has a total of 21 articles followed by a Subject Index, Cumula- 
tive Index of Contributing Authors.and a Cumulative Index of Chapter Titles, Volumes 26-30. 

The Origin and Early Evolution of Turtles 
Uses of Evolutionary Theory in the Human Genome Project 
Streams in Mediterranean Climate Regions: Abiotic Influences and Biotic Responses to Predictable 

Choosing the Appropriate Scale of Reserves for Conservation 

Con .pec ific Sperm and Pollen Precedence and Speciation 

Global Amphibian Declines:A Problem in Applied Ecology 

Using Phylogenetic Approaches for the Analysis of Plant Breeding System Evolution 

Evolution of Diversity in Warning Color and Mimicry: Polymorphisms, Shifting Balance, and 

Consequences of Evolving with Bacterial Symbionts: Insights From the Squid-Vibrio Associations 

The Relationship Between Productivity and Species Richness 

Analysis of Selection on Enzyme Polymorphisms 

Polymorphisms in Systematics and Comparative Biology 

Physical Biological Coupling in Streams:The Pervasive Effects of Flow on Benthi Oi jani n 

/ obiolog E plot n ] Origin l olution,and DistribuTk i n i J i tl Univet 

Evolution of the Eastern Asian and Eastern North American Disjunct Distributions in Flowering 

Full of Sound and Fury:The Recent History of Ancient DNA 
Do Plant Populations Purge Their Genetic Load? Effects of Population Size and Mating History on 


Gene Flow and Introgression From Domesticated Plants into Their Wild Relatives 

Barbara LBowriNG.2000.The Berry Grower's Companion. (ISBN 0-881 92-489-X ; hbk.).Timber 
Press. Inc.Jhe Haseltine Building, 1 33 S.W. Second Avenue. Suite 450. Portland, OR 
97204 3527 U.S.A.(503 1 27-2878,503-227 3070 fax; $29.95 
hbk. 308 pp. 40 color photos, 25 b/w illustrations 1 7 tables, 6" x 9". 

-i;1) General Principles; 2) Berries in the 
jmbles;5) Blueberries;6) Grapes; 7) Minor Crops; Appendix: North 
fry Plants; Glossary; References and Other Resources; Plant Name 
w forthcoming in Sida 19(3), 2001. 

G.Terry Sharrer.2000. A Kind of Fate: Agricultural Change in Virginia, 1 861 -1 920. (ISBN 
0-81 38-2569-5, hbk.). Iowa State University Press, 21 21 South State Street, Ames, IA 
500 14-8300, U.S.A. (Orders: 800-862-6657, 5 15-292-0 15 5, w m i ii,v, ,edu) 1" ' 
pbk.256 pp.,b/w photos, 6" x 9". 

From the du5tjacket—"A Kind of fiitt \, , , > j addresses how mod- 

intervention in agricultui m m .nin n f >\ il if n i, i , mi imodity prices, land 

from the period, poignant photographs capture the essence of these farmers' daily trials and tri- 

Contents.— Ackno I i m ,, In i ni.tnnl) M t\ i I t.-m ,e f pidemics and Epizootics, 
A New Order of Thing i u n u I i n .») < u , t , i, , ,,,,,,, ,| lu 

Crops and Crises; 3) loil and Tumble: 1 ile in the Country, Debt, Axes, and Despair; 4) Professing 
Change: Growing Knowledge: I he Vn<]inia Agricultural I xgcnment Station, Reaching the Farmers; 
5) The New Farming: Drive for Production, Dairying: Progressive Exemplar, Capital and Credit; 6) 
Reforming Fate: Farmers and Tenant f A I ur ,l[ A r 1 unuHhi i mi A nd ofan Era;Conclusion: 

K. Morton and Joan M. VhNN 2000 The Flora of Manitoulin Island and the Adjacent 
Islands of Lake Huron, Georgian Bay and the North Cannel.Third Edition. (ISSN 
031 7-3348, pbk.). University of Waterloo Biolng , n nimli r 40 Biology Series, 
Department of Biology, University of Waterloo, Waterloo,ON,CANADAN2L3G1 (519- 
888-4567, ext. 3751; 519-746-0614 fax; jvenn@sciboiq.uwnloiki()(d) $ A -A pit il 
bound (Canada);$37.50 spiral bound (outside Canada)/A( A ; hbl U minim s ,000 
hbk. (outside Canada). 374 pp., 1 24 color photos, 997 range maps, 7" x 1 0". 

1 mi' ulin I I nd in 1 th hi ii il t i I lid in th it tin it i r inn [ r 1 ake 

uron and the Nortlu f in An in tit - n i i i i .i nt-a of unique biological 

terestwitha remaiAA, n u il . A i ,t the total i ulai t m 1 Canada)." 

The Flora accounts for 1 350 kinds of vascular plants.The 997 distribution maps are computer 
"awn from a database t m " i id ti i , ih i nt iu t n\ chapters deal with 

lion in tin 1 iciiuMi lii I 1 -la A In u -n n i ,n i li i nt the more interesting 
id beautiful plants of the region. 

Text and the maps have been completely revised in the > edition; the color figures are um 
lanqed from the A" 1 edition I hem is no family key not do families have keys to genera and gen- 

i t pe, ie, an >>, > , ,|, , M ,| | , i pi . hk d io s| me, /' ant , , p Sl om h 

Ronald M. Lanner. 2000. Conifers of California. (ISBN 09628505-3-5, pbk.; 09628505-4-3, 
hbk.). Cachuma Press, P.O. Box 560, Los Olivos, CA 93441 , U.S.A. (805-688-041 3; $24.95 pbk., $36.95 hbk. 288 pp., 54 original full-page wa- 
tercolor illustrations, 7" x 9". 

From the cover.— "Conifers of California is the first book devoted to all of the state's conifers. Its au- 
thor, Ronald M. Lanner (professor emeritu U1 mi ni i itv) has explored, studied, and taught 
about forest trees in the West for 40 years. In Conifers of California he shares his expertise and intro- 
duces each of California's cone-bearers in an engaging text that serves as both natural history and 
field guide. Lanner's narratives are accompanied by detailed identification information, watercolor 
botanic illustrations by the late Eugene O. Murman, color photographs of each species by well- 
known landscape photographers, and distribution maps." 

Contents.— Preface; Acknowledgments; Eugene O. Murman, the Artist; The Names of Trees; 
Cones; Pinaceae, The Pine Family; Cupressaceae, The Cypress Family;Taxodiaceae,The Baldcypress 
Family;Taxaceae,The Yew Family; Appendices: A: California's Soft Pines and Hard Pines; B: Conifer 
Hybrids in California; C: A Key to the Genera Based on Characters of Mature; D: A Key to the Genera 
Based on Leaf Characters From Lower Crown Branches; E: Alphabetical List of Conifers Growing 
Naturally Within California; Annotated Bibliography; and Index. 

the photography is stunning and shows off the las ihn i .ml -in jm ruiesque California land- 
scape hie nook is beautifully designed and vvsujlc r i a in in il I i i n i anv 

Den Bown. 2000 Aroids: Plants of the Arum Family. Second Edition. (ISBN 0-881 5 

485-7, hbkj.Timber Press. Inc.Jhe Haseltine Building, 1 33 S.W.Second Avenue.Su 

450. Portland, OR 97204-3527, U.S.A. (503-227-2878, 503-227-3070 fax; wv\ $34.95 hbk. 468 pp. 108 color photos, 50 line drawings, 6" x ' 

Table of Contents— Foreword to Second Edition hot, id iln I n ,t Edition, Preface; Ackno 

-iim n tin i Mian 1 ) > anations on a Theme; 2) Of Tails and traps and the Underworld 

Woodlanders;4) Aquatics and Amphibians;5) A Place in the Sun;6) In the Shadows; 7) Towards ■ 

Light; 8) The Titans; 9) An Acquired Taste; 1 0) Acids and Crystals; Aroids in Cultivation; Checklis' 

Aroid Genera; Glossary; Referent - mihi « 

i litot ) 7000. Breeding Ornamental Plants. 

if i i H, Hasi Itine Building, 133 S.W.Sec- 

d,OR 97204-3527, U.S.A. (503-227-2878,503-227-3070 
S4.95 hbk.359 pp.88 color photos, 10 b/w photos,26 

MJmilMi iii s)i ^ing Siberian lris;6) Breeding Hostas; 

g Ornamental Aroids; 8) Breeding AfricanViolets; 9) Breeding Gesneriads; 1 0) Breeding 

II) Breeding Penstemon; 12) Breeding Rhododendrons and Azaleas;1 3) Bn i' ; 

Laurel and its relatives; 1 4) Breeding Camellia 1 r ) in H i Inn ' 

ow; 16) Breeding Magnolias; 17) Breeding c ] >\ f t mi i \ endix Sources of 

Plant Nar 

Dorothy J. Calla\ 
(ISBN 0-85 

fax; www. 

vAYand M. Brett C a 
31 92-482-2, hbk.).T 

Preface; Acknowledgments; 1 ) Genetic 
3) Breeding Daylilies (Hemerocallis);4) Bi 

\ n Field Guideto Indiana Wildlfo.-.^, :. ■ 3,pbk.).lndiana 

University Piess, 601 N. Morton Street, Bloomington, IN 47404, U.S.A. (Oi. Im mm 
842-6796, 81 2-855-8507 fax). $i P bM || Ji.l i In it mi 

5" x 8". 
Contents.— Prefa numbers, Photo- 

graphs, Drawings Sut nti'i. urn. \uthoi ot i niith nun. i , i nl it names Measurements 
of plant parts; Main pat mi .pi .1 p< i. m,, , i | I nr m I fl , i Distinguishing charac- 

teristics, Habitats an l I liu tl in : I. niii inn ill in ii tutus Glossary;What is 

a flower?; Aster famil\ t iinin^ i mil i I ' n hi I i i I i ■] inferences; and Index. 

This is a nicelv dor I ill i i i i M i ill | i i , , > * » 

Sida 19(3), 2001. 

Ai an I Bi 1 1 u, Wn i - ( . K . .i ■•-, mid Ai ■ i n I' Pi i ■< . 7000. North American Boletes: A 
Color Guide to the Fleshy Pored Mushrooms. (ISBN 81 56 0588-9, hbk .). Syra- 
lUseUnivotsity Press, Syiucuse, NY 13244-5 160, U.S.A. $95.00 hbk. 396 pp.450 color 
photos, 7" x 10". 

Contents.— Preface; Ac kno I. jgmmt k\mn ,n n l-mlet kniy in hiief; Introduction to the 

Boletes.Typical Bolete Fruiting Body: Macroscopi. I itu Mow lu U I hi Book How to Identify 
Boletes; Field Key to the Bokm I), ription ,11 let Ucneia ml pm i< Undescribed Boletes, 
' S 1 '"! ' Hid P « nl I lii ' >p I , i in mi I I mi nmmon Names; and 

Gil Nelson. 2000.The Ferns of Florida. A Reference and Field Guide. (ISBN-1 -561 64-1 97- 

9,pbk.,1-56164-193-6,hbk.).Pin< i[ | I li In P. >| i »• it <;t i I L 34 <0 

U.S.A. ( $27.95 hbk. 208 pp. 200 + color photos, 6" x 9". 

From the back cover -"At least 1' An) t in h I i v i I Honda From northern 

species whose south, it, t ,n h 'mm mi ih i li i urn Mm mi northern tier to tropi 
cal treasures that spill ivm in i h nh n tip, II i i th n [ imllel in fern diversity and 

number. No othet like i i n t th m n .in n i mi it ■> mst such an expanded 

P field gui ill fust us to It thi unazirm i t fei I of lowing in the 

tradition of John K small II >' , iiii t < rmong and Olga Pakela's 1976 edition of 

the same name,thi n mi m i t I II j n ,t II i t i md naturalized fern 

species. Color plak t. itut i i ilur i in m h t hi I t In I tare species never be- 

:ie a complete ok. mm , i ,. ik 

vlatihi. ominijin si'aA ,'A MPfMcP 

E. Rogers. 2000.The Magnificent Mesquite. (ISBN 0-292-771 05-3, hbk.). University 
Texas Press, P.O. Box 7819, Austin,TX 787 1 3-7819, U.S.A. (5 1 2-47 1 -4032). $24.95 h 
1 67 pp., 1 8 color photos, 2 line drawings, 5 maps, 1 2 tables, 5" x 9". 

i the dustjacket.— The mesquite tree has deep roots in the American Southwest, liteial v 
widely popular for barbecuing, woodworking, fl 

'ell-managed rangeland. 

vritten especially for a general readership, one of th 
>nts a wealth of information about its natural histi 
>rking uses. Ken Rogers describes the life cycle, sf 
mesquite, which is native or naturalized not only ir 

! i rig i mesquite wooc 
or and making mesquite bean jelly. He also looks into the ways that people are using mesquitt 
nature, from rangeland management in the Southwest to desertification prevention in arid coun 
bs. Color photographs and maps complement the very readable text. 

Contents.— Introdn. ii mM mi v'nar It Predators of Mesquite; Mesquite m Texas Am 
2 Southwestern United States;The Uses of Mesquite; The Mesquite in Verse; Worldwide Aspect 

i / i. Inr i i juite ,Ful | 

isopis (Family Leguminosae/Fabaceae) and Its Species; Appendix 2. Sources of Information, Ref 
mces and Further Reading; and Ii 

). Phylogenetic Analysis of Morphological Data. (ISBN 1 -56098-81 6- 

" " " :, hbk.). Smithsonian Institution Press, PO Box 960, Herndon, 


ulesVersesMorphologylnSystomiti ilm n,i , i v Hi ,\ n i 

2) Character Selection and the Methodology of Morphological Phylogenetics;3) Discovery of Phy- 
logenetic Characters In Morphometric Data;4)The Usefulness of Ontongeny in Interpreting Mor- 
phological Characters; 5) Coding Morphological Variation Within Species and HigherTaxa for Phy- 
logenetic Analysis;6) Hybridization and Phylogenetics;Special Insights From Morphology; 7) Using 
Stratigraphic Information in Phylogenetics;8) Logical Problems Associated With Including and Ex- 
cluding Characters During Tree Reconstruction and Their Implications for the Study of Morpho- 
logical Character Evolution;and Index.— Review forthcoming in Sida 19(3), 2001. 

'You don't have to be a botanist to use it! All the 

to teach plant taxonomy, family characters,' 
Excellent introductory materials on the history and geology of 
North Central Texas. Great for identifying indigenous plants in 
an ecosystem! Essential for student wildflower collection 

projects! Use this one book instead of fivi it has it ill 

E-it! Worth C 'unit I ' v School 


i i'ik n t 

and naturalized plants.This book should be in the 

every gardener, landscape architect, horticulturist, 

lanner, farmer, ranchei.and wildflower enthusiast." 

—Howard Garrett Th, Dm Do lot 

Shinners & Mahler's 


Flora of 

North Central 



INNOVATIVE. With every species illustrated - it's a new 

\e\ i irluii it lexa tl >• i 1 I i- ith f tn i i 
- ] li' '■,/" ',-'", i mi , uilh't 

"The Illustrated Flora on North Central I e> i i: a m i |nifi< ;nl ■ ( il 
This abundantly iliu i i I in I i 'it only be of use to 

those peopie wantim: to idetifiv ioxan i)fiitis, it will bv a 
model of how a local flora should be prepared." 

i Pram 

r, Royal Botanic Garc 


. ■ ', ■■!",'.. V 

■SiH " 

: >j J Of:.r:cm 



; v ->w 

1 1962. Inherited 

Mahler, Director 

Felix Llamas, Contributing Spanish Editor 

Ppi '.'i' Botanh i ra< ult id d< I- )l< qia 

1993 it has be* 
published by B 

E-2471 Leon, SPAIN 

d i C ' i mfi gionsto Botany 
Voi ume 1 9, Number 3, pages 445-766. 
23 August 2001 

G)EVEK,nl 200 I 

Botanical Research Institute of Texas 


ISSN 0036-1488 

Table of Contents 

Pa. i! M. Pe'm^on a 

Guy L. Nesom— 507 

T '] Keysseria and Pytinicarpa < ■'■ 

Guy L Nesom— 513 

Saccogi ( <■ ' M ' 

r PT H h— 519 

M ■■ • -'I ■ ■. ,i in Pennisetum ( ■', , , (>[]'' | 

I Rodriguez Tuerina— 539 

jra (0 ban 

■■.•■■ ,u n I A ,VVP :/■ 639 

Flows m , 
Gnaphalium exilifolium 

The VASCULAR flora of Aw 

.in County, Mississippi 

MacH.Alford— 645 



George Yatskievych and J 


Glochidion PUBERUM (El 

jphorbiaceae) naturali. 

Miriam L. Flarn and Low 

ei i E. Urbatsch— 71 1 

RT.Haiw, 715 


DavipJ.Homn 727 

V •■-! I A I I 

FlvAKpii F ipvsa C 

\.GR".i !'■■ TO'-'R! • ! B.l . I ',- ■ ." 


Pennisetum pennisetiforme (Hochs. & S 

Gray) Nesom, comb, nov.— 516 
(Raf.) Terrell, stat. nov.— 592 

l S4S 




j'PLF i in VTi iell.lOMB. NOV.— 610 

\T|RR|-!I,VAR. NOV. — 611 





W.Takeuchi M.Golman 

itute of Texas (BRIT) Divisional Manager 

■> t urn and Papua New Guinea Forest Authoi 

iM Res< an h Institute PNG Nation - 

1 1 ae, Mi >iohe Province 41 i P.O. Box 5055, A >i< A ) 

PUA NEW GUINEA National Capitol Distrk I PAPUA NEW GUINEA pg 

In imn u ! qui i i m i i\< \ pUU i tied < i i ij ui i ii n h n k i i 

pop liar fb di velopm i uisii tun tti ol n \ du \l hou Ii im< >l ihei il in i 

ii , in nated there haw 1 i mpl U overall s\ n! 1 j i'i . i i in : 

it Ink save na wok painim am ibin kainap plain i me 


mUftftttS:%Z£ltWbfr^&Z>\Z.bfrfrt>t>?. %<Dltli><D&-t}\t&bftT% 
fc. *HXT(i, a«E©WSE^rstT**a*d«^»"CU*fi!t***a(c:«»(fe>nT 
U*fr«jfc<D*fiElcfca*£Tfc#&, iiiio«Pia©tt«iHa<i:5Ba»S:fcwr 

In spite of its status j i lt>l> . nhiln bn,n di\< i iln u u >n I A »n.i M< w Cum. ,i 
(PNG)has thedubiou li i n lion of being on o vlak i most inadequately 
surveyed nations. Stevens (1989) had defined the relatively well-collected areas 
in PNG as having a collections density of 50-] 00 specimens per 100 sqkm. But 

even by this easy measure, the well collected localities identitied h\ l"i is criterion 
werepiimanh conlim d to t lew principal dtainag< and high mountains. Re- 
cent analysis shows that the background level of collection density averages 
less than 25 per 100 sq km over New Guinea as a whole (Conn 1994), a rate 
substantially below the standard regarded bv Stevens as a minimal baseline. 
Within the Malesian region, only the Celebes and Sumatra have comparably 
low collection indices (Stevens 1989). Although Steenis (1950) had estimated 
thai >0 \ u ol coordinated ( ploration would b requi red to document the 
New Guinea flora, the rate of documentation has not improved since that as- 
sessment was made, and has in fact dramatical!) declined within the last 30 

i l.i n f) I i In* ii ii land | I 

are to U dew id ind i in pit nu and G indigenon igcncii the pn ent trend 
in documentation must be reversed. The urgency for corrective action is espe- 
cially acute when the target country represents a biodwersity hotspot within 
which social and economic changes are expected to intensify. 

Papua New Guineas population gtowth rate of 2. Go is now one oi the high- 
est in the Pacific region. From a present base oi 4.7 a million, the number of 
people isexpect Itodoubl by 20 Kliiiinou G, I i m 1 l ><> o The demographic 
projections are especially consequential because an estimated 84% of the popu- 
Inioiii rural, and all ueli househohl are dependent to nine degree on slash 
and burn agriculture (ibid). Currently, an estimated 200,000 hectares of land 
are cleared annually for subsistence, from which 20,000-30,000 hectares rep- 
resent natural growth lorcst permanently removed by various clear eui opera 
nous including industrial logging (Filer 1995). 

With the anticipated increases; in human population, subsistence activi- 
ties are expected to progressively mimic the el Gets ol forest clear-felling as fal- 
low cycles are accelerated ([.ouman eGr Nicholls 1995). When PNG enters the 
leep pai i oi flu pi iiei I popul now rowth curve serious environmental 
im pacts are likely to occur I'he re are already mounting 1 1 id ications of impend- 
ing failure in the subsistence ,\ tern G everal pun metal areas due to intensi- 
fication of cropping rotations (Levett & Bala 1995). 

At present, PNG sttfl retains 70G ol its primary forest cover (Me Alpine & 
Quigley 1998) and is one of only four tropical countries with extensive tracts of 
original vegetation tSuzuki 199 3 J. This remarkable state of preservation has 
unfortunately lostei d i complacency unong ciei i pi olessionals by encour- 
aging an expectation that current inadequacies l n docunu utatuon ran alvavs 
be reversed by future action. The underlying assumptions are not likely to pcr- 

The poor slate ol I lot istic documentation lias maim and significant mani- 
I est ar ions m Papuastan botany \ disproporl innate number oi taxa are known 
from single col lections, and t economic knowledge even at family level is often 
highly superficial (Johns 199 G Virtually nothing can be said oi' the basic biology 

and populational variation for many of the most important Papuasian plant 

groups (Stevens 19H9 While ii genu II no led that certain mountainous 
areasarehotspotskii i Km in ( nn -nil in ii i nnpo ibK o ul< u \ lou i K \ non 
centers with the scant v data at hand. Some consequences ol the past emphasis 
on high elevation exploration can b< ,een in recent m nil trom I he >cnin 
Conservancy's lowland m\i\ol Joseph , mi u Ii n 9 pi i ics were newly de- 
scribed from accessibK i n u rh prin i| tin tional highway in Madang 
Province (Huynh 2000; Takeuchi 2000a, 2001). Lowland environments are pre- 
ferred venues for economic development, so there is considerable potential for 
floristic losses in tin oik < ont inn, d n In ol l>, , nu 1 1 documentation in 
low I aid howi io i • in ha\< si viae < on i qui m i m ui\ im me intensification of 
commercial logging, considering that over S0° o ol PNCs loggable forests (as 
defined by industr) tandatch arc in i In o land I i i i n i n & Nicholls 1995). 
Of the forest blocks i ui<ntl\upt< uiinp nu n h mt ibl i ml h7" ilsomou 
mmedium-crowncd lowlsnd lull low st (om n l'ai|in in 1" m i. i , . i oi\ 
Hm on Hamtnermasto- d Saunders 1995) i v. fetation type which is probably 
Papuasias richest Id on in formation (Louman i dudioll 1 995 J 

several competent >l crvci h veenumei l d lo< ho of particular value 
and urgency for exploratory survey within PNG (Johns 1993; Steems 1950; 
Stevens 1989). A common thread extending through all these recommendations 
is that the exploration status of highlighted areas has hardly improved since 
thetimeof theSteeni commentary v'ery little hn changed with respect to the 
quantity and quality of botanical data over I he past several decades. The lack 
of substantive prog n ■ adversely affect taxonoirnc andecological a , .merits 
and ultimately impedes sustainable management end development within the 

In PNG, any activity requiring access to natural resources m ust include consid- 

eration ol tin trai mil I; ncl n nun \ tern \n < tun u d 97' ml diet oiutl i ; 

is under customary o\\ net hip ubject to complex s tern of usage rights and 
social relationship win n n l hemselves snpcrim posed over a multitude of 
cultural-linguistic traditions (Crocombe 1974; Holzknecht 1995). Alienated land 
and properties otherwisi rnulrr governmental contro u irtn Id ion i 

ent. Irrespectne ohndu n In.m i i n il , > u i d I ml ibn i in 

land-use issues are the viLlage clans and landowners ol' specific forest blocks 
(i.e., the 'papa graun'). 

Because so many prerogatives reside with local villagers, direct negotiations 
with the customary lenanls tire mandatory for aim/ scientific program. This is 
not oeasibdoui hi n m mo tigation s pi incipjl u o\ et seas-based. Due 
to the proliferation ol advocacy groups with environmentalist agendas, many 
landowners have also become conditioned to regard extract i i ct iti nils i 

land (particularly by loreigneis; with considerable suspicion. This situation 
applies especial ly to bioprospecting. Survey opcratorsdo well to avoid prospec- 
tive collecting altogether, because o\ object ions i hai have been raised in relation 
to such activities. Any contemplated project would be required to explain its 
aci i vii ies loan oil eniii in keptical and uninformed audience. Having a func- 
tional ! now ledge of rh lin nib rn ( M hn i m rol pi in) is i i mi il. 

A significant consequence of l he primacy of custom a rv rights in Papuasia 
is that certain Western mechanisins lor pei mit issuance and resource access are 
culturally utele\ m.t m PNl i I'hci i loi exampk in rich n [iiirementa i "plant 
collecting permit.' Hven if such permits were established by government agen- 
cies, they could never be enforced at i he local level where botanical collecting 
actually occurs. Onlv the customary landowners can grant approval for removal 
of materials from their territory. Paradoxically, while this eliminates much of 
the bureaucracy characteristic of Eurocentric management systems, survey 
operations are oft en rendered more complicated and unpredictable, because the 
activities are entirely subject to the whims o\ individual landowners. 

Modern commentatoi have been unanui u m< pi ing a need for de- 
veloping local capacity as a prerequisite for longterm assessment and manage 
ment of PNG's biodiversity (Beehler 1993: Conn 1994; Hamas 1998;Johns 1993; 
Sekhran & Miller 1995). Public sector agencies in PNG are subject to unpredictable 
changes in government support due tosfults in political direction. Based on pre- 
vious trends, it is vci v unlikely that vital commitments to science capacity 
building will occur through in-country funding, t ollaborativc studies are a 
potentially effective means lor improving internal capabilities when appropriate 
agencies are engaged as partners in research. Programs which provide for partici- 
pation of qualified counterpart scan make longterm con tri but ions to bioinventory, 
but the partnerships must be careful I \ selected and not merely convenient. 

The PNC. infrastructure in science hasexpei lenccd prolound changes over 
I he last thirty w;u I Mil me tin < olom il adnurii [ration md ioi t bi icl turn iln i 
wards, many of the functionsassociated witfi florist ic survey were centrally invested 
in highly capable units such as 1 .ae Herbarium (J .A l : .) and the former Department 
of Forests. Government facilities in the 1960s and early 70s virtually monopo- 
lized plant exploration within PNG. In contrast, nongovernment organizations 
(NGOs) were conspicuous by their overall absence Irotn activities involving 
botanical documental ion. I lowever ai lei PNC 1 became a sovereign state, the capa- 
bilitn ol national agencii lor i Ion i work progn aveb. declined as budgets 
were subjected to a political reordering oi priorities (cl . Conn 1994). The earlier 
priorities are unlikely to be restored in the future because institutional and social 
realities have been so completely transformed. Por example, there are now no 
Ph.D.-level professionals in the PNG National Forest Service, and currently only 
one M.Sc recipient is seiwing with the national herbarium. 

Tosomcextcntjln e trend magen y capabilities haw been mitii iterl w thin 

the last 10 years b\ countervailing dew lopmentsm the nongovernmental sector. 
A seminal event was the Conservation Needs Assessment (.CNA, Beehler 1993), 
the first country wicl plan to define c< npreh nsiv< priorities for conservation 
and research action. The CNA stimulated establishment of a multitude of Wildlife 
Management A ica si w M Wandcl ,\ -01 iai d pwj. . i b ■ .1 d on the Integrated 
Conservation and Development (ICAD) model (cf. Saulei & Ellis 1998). Not co- 
incidentally, many ol tl post CNA initiaiiv< ncomp at least in part, the 
areas prioritized by the CNA. Together with these developments, a number of 
NGO entities have assumed effective jurisdiction over the VvM As and their land- 
owner groups Mam of PNCTs prime wildi iu environments are presently 
included under NGt Main em< in \t i| iin i hi • I In total biodiversity 
represented by such partnership i \ r\ ul iantiale\ n chough the arrange- 
ments collectively comprise on I 'i 1 ! rf the P G land area. In addition, a 
large backlog ol sites is under consideration lor lul are conservation action. 

The combined effect of these events is that the hierarchy of PNG science- 
related administration has been transformed by the creation ol a new infra- 
si i uciure Hue to the r< lative n « < n< \ ol the in w ai rang* merits, there has been 
hlil eh in m h w i\ biof ic il in \ u < mdi i ceclii I i ( . Howew i n ii\ 
allthe WMA-related NGOs maintain a fulltiine presence within their respective 
wards, so the opportunity lor a< hie wing el lei ivi coi irnumtx integration with 
surveys are now very promising. The WMA typically incl d i id nt -coord i 
nators and protocol s lor rn a i i 1 1 a i 1 1 i ng con tin uous 1 i a i son with 1 andowner groups, 
critical functions the i government interests can no longei provide in the remote 
areas. From an operational perspective, each of the NGO/WMA combinations 
is the equivalent of a research lacilit) I h< elements loi multilateral surveys 
combining professional and village participants are thus in place, requiring only 
that the individual components be drawn together under a common plan. A 
future schedule for comprehensive Inoinwmioi nes could be eo istructed using 
the government planning instruments on one hand and community level 
implementors composed of WMA/NGOs on the other. Linkages of this sort are 
already the basis for several contemplated operations. 

The sheer numbers ol jk u involved in docum m i ion and inventory are 

overwhelming when a ppi.M In d in > n ti ulri L ] »'t | < 1 1 \ es m collection and 

curation. Time honored traditions in tropical exploration are no longer adequate 
to the tasks of acquiring and analyzing large collection sets. A revised approach 
to floristic documentation is clearly required, and is perhaps best adapted from 
the experiences of entomologists faced with problems similar to those in bo- 
th rough refinement of comparable methodologies first developed by INBio 
in Costa Rica (Janzen et al. 1993), PNG based researchers at the Parataxonomy 

Training Center tPTC) have devised practical solutions to the time demands 
presented by intensive sampling in rich tropical habitats (Basset et al. 2000; 
Novotip el al ITT i.l hcirapproaeh has been terrain local vil lagers (as parataxono- 
i ni^lsi m i he liuulainentalsol col lectin;;, specimen sorting, idem if ical ion. and 
compLttei l.i .d.l n i nanaia neni I he lew international professionals are pri- 
marily engaged as instructors, quality control agents, and ultimately as data 
interpreters. By ionising the intervention oi individuals to the points where 
their expertise is most eltective, the research process is thus streamlined and 
accelerated. The immediate product of these arrangements is [hat extensive 
specimen sets haw been acquired and processed within time frames that would 
ordinarily require prohibitive inputs. The demonstrated success oi such orga- 
nization inentomolog\ b\ itsell shows that the protocols can work for botany. 
Insects after all, exceed the floristic diversity bv several orders of magnitude. 

Asan example of the enhancements offered by the new procedures, within 
a period of 5 years the PTC studies in insect herbivon haw collected, sorted, 
mid mounted over 100.000 specimens oi leal chewingand sap sucking insects 
representing ca. 1,300 species (cf. Basset et al. 2000: Novotny et al. 1997). Such 
outputs considerably exceed those obtained bv convem lonal efforts without 
parataxonomist assistance. Over 40 scholarly papers have been published by 
the research team, in stark contrast to the normal downtime between study 
inception and publication t usually To years) lor the kind ol eco-entomological 
inquiry being undertaken by PTC researchers (krwtn 0)9.0. These improve- 
ments are a direct result ol the use oi parataxonomists in time-intensive ac- 
tions such as collecting and sorting, allowing other participants to optimize 
their own activities on cost-effective schedules. With the marked increases in 
sampling outputs, new insights haw emerged which could only have arisen 
from statistically large datascts, such as arc now being generated by the new 
protocols I he PI '. ampling piogram ha led to a reevalual ion ol insect plain 
relationships, with wide ranging implications lor understanding t he i ompo 
nentsol invertebrate eliwrsilx in tropical systems (Basset et al. 2000; Novotny 
ei al. 100/ i. In an analogous manner, quant Hal ivcl\ boosted floristic surveys 
haw the promise oi spawning comparable adwnces in our knowledge of the 
taxonomy and ecology o\' the Papttasian Malesian flora. This isespecially likely 
when the existing sampling co\'erage for plants is so erratic and sparse. 

A significant factor in the success of the PTC operation is the fact that instruc- 
tors and students share ful hi me residence m a combined laboratory dormitory 
complex while pursuing con in ion research object iws. Continuous interactions 
between mentor-trainers ami parataxonomists instill a sense of fraternity and 
purpose which is not easily replicated by conventional piopcm. even though 
the latter may otherwise superficially mimic the PTC program structure and 
objectives. Preservation o\ the social relationships will be crucial to effective 
translerance of the parataxonomist concept to floristic survey. Similar patterns 

for success were previously pioneered .it the Chnstensen Research Institute 
(Orsak 1993) and more recentlx h\ tin \ illage Devi lopment Trust. Botanical 
planners should note the methodological paradigms, particularly the conditions 
contributing to their effectiveness a rh< nipln uion lor floristic survey are 
both timely and considerable. 

A point worth repeating is that the parataxonomist concept has been thus 
fit apph.-( pi nn u il\ bN entomologist I noiiini i I \ t In botanical profession 
has been slow to recogni c that nany pi >blem ol hi lo u al sampling in the 
tropics are universal, and applicability ol ucc< fultechniqu' i likely to cm 
icro disciplinary line; I pecially with traditional cultures such as PNG, folk 
knowledge of plants is olten more extensive than the corresponding base for 
insects (Basset et al [000) so plants are actualh vei \ appropriate subjects lor 
parataxonomist -assisted investigation (Novotny pers. comm.j. 

II comprehensive paralaxonomo program n it templed within PNG, 
complementary improvements will be required in the laci lities associated with 
biological documentation. Development of local capacitii o lloristii in \ 
is unlikely to achieve lasting results if the physical security of collections (and ot her 
survey products such as databases) cannot be assured inside the host country. 
This can be a problem in developing societies where funding priorities for sci- 
ence are generally low. A permanent instil ulional base will also be needed, and 
is best achieved through the development oi organ ions specifically devoted 
to parataxonom v, rather than by placing parataxonom ists in preexisting herbaria 
or government institutions. With scientific facilities in t he public domain, there 
is likely to be an adimui i u I u Ja\ in | ok mnal taft over individuals 
without formal credentials. The continuity ol parataxonomisis in such etna 
ronments would be less secure than in a mission-specilic unit such as PTC 

Steemss estimate of a hO year cycle oi coordinated exploration is opera- 
tionally impossible. In its reliance on outdated concepts of how nich inventories 
should be achieved, u is also incompatible with the social realities of contem- 
porary Papuasia I m i Muud) ' i )f Miii. in nhl i K lo upj mi mil i nd.d 
programs of dclem d n hi iiun imuu \ h,iw i I mining factors devolve 
exclusively upon a mall number of highly trained pi oh uonals, whether in- 
digenous or foreign, it is cloubtlul that real progress will be made toward the 
goals implicit in torn pwln n in urw\ <> n |\ | u i -^ ' i i i i i < nt expansion of the 
workforce, like that afforded by parataxonom y. will the inventory process be 
able to encompass I he diversit\ withm Papuasian ion ;i o osystcms. Method- 
ologies which improve cm ony rates ol I loi lstu do< umeutation are especially 
urgent in view of the habilai destruction winch has occurred in Malesia since 
the time of van Steems (cf. inter alios Kiew 1990). 

Future surveys should include teams composed oi purpose-trained 
parataxonomists. Participation by landowners will also permit outputs to be 
amplified across the board, result mg m survey yields substantially higher than 

convent ions. 1 expeditions \ collateral ad van tag< ol com in unit)' involvement 
is the associated opport unity lor integral me, t rada ional k nowledge systems into 
the collections documentation. Hthnobotanicul inquire can be easily assimi- 
lated when local mini mi mi an; en; igcd m surve) 

Although there is clearly a general failure ol lloristic documentation in 
Papuasia, little attention has been explicit I y devoted to the way field operations 
are actually conducted. Yet that should be the logical starting point in any analy- 
sis, because how collections are acquired and the limitations associated with 
their acquisition, cannot help but affect everything else which lollows. In any 
such examination, probably the most obvious limiting factor which would 
emerge is the mel feet iveness ol existing collecting methods. 

In die early days ol the PNd borcst Service, nl le lire was ol ten used to bring 
down Icrtile branches I rom the eanop\'. Nowada)'s, in a country where high- 
powered firearms require special permits, this is no longei a viable option for a 
number of reasons., not Iciest ol which being that possession ol such weapons 
would attract undesirable attention to the collecting, teams. In order to obtain 
specimens from high canopies, local climbers are i hus employed i^n nearly all 
surve\ s. Selected l reesarcalso I requent l\ cut down. Both mefhodsare very time 
consuming however, and it is not unusual for a single collection obtained by 
such means to take a hall hour or even longer. Other procedures using sling 
shots, extensible poles, wire saw s, etc. are use! ul only in certain situations, and 
also require a substantial amount o\' practice before the field assistants can 
achieve neisonablc prol iciencv. 

The search for suitable gatherings is often length)' just in itself. Especially 
in mature grow th. where the forest biomass and collection targets are located 
far above the ground, few taxa will be within easy reach. I rider prevailing con- 
ditions, a collector with several assistants can expect to obtain an average of 
only 30 taxa per day. Oailv rallies tend to be higher in regrowth and montane 
vegetation because of their lower statures, but general 1\ a botanist will not take 
men di n )0nurnbei evi i un.ik lavo idle cu cumstance^ Vld to these con- 
siderations the fact that colleetois in logistical! v difficult cn\ uonments are 
olten buulened In institutional quotas lor multiple duplicates and the time, 
cost demands are increased even further. 

Id forts diverted to the preparation ol duplicate samples detract from the 
documentation process, ! 'he international herbaria with significant traditions 
in Papuasian botany are lew in number, so a point oi dim mid nng returns is quickly 
reached when distributing specimens. (. ol lections con si si i rig of numerous du- 
plicates are very inefficient in terms of the costs in obtaining those duplicates. 
Flonstic inventor)' is better seined, by securing small sets t ] duplicates) of dif- 
ferent conspecifie numbers rathei than by obtainingsingle numbers with many 
duplicates. On the former procedure population variation can be effectively 
assessed, while the latter procedure contributes little. 

The reasons for the inadequate documentation of the Papuasian flora can 
be entirely understood at the most immediate and basic operational level: that 
of the individual collector laboring in the bush. Nomattei how much individual 
effort is expended, the per capita outputs are not going to increase to an extent 
necessary to reverse the current trends in botanical inventory. Since personal 
yields are not amenable lo ii ii |;hi\< in m rh < omm n in ill inin i t, < 
pand the workforce. Themost pi ■< in aland o i lb reali • means of achieving 
this in traditional societies is through the application of parataxonomists. 

An unfortunate fact okonrempoiai \ un ;. isthatparataxonomy-assisted 
outputs in themschew mm r u hi, \ ill r In d, m d nb| an , -, Even when survey 
collections attain respec ml >f solum, mam ,>. .1 -1 101 1 Jim e suffered from the 
myopic attitude that the botaim al with, umw aie an end in themselves. 
Oftentimes there has been no attempt todi rninai finding 01 even to assemble 
the results into any kind > l ibh lorn II, onn on d con 1 1 ti n I 
ization of resource manage merit within du sin veyedarea ;, which in practical 
terms is probably the most important downstream product from biosurveys. 

Perhaps die best example of rh, preceding eii m m < \ » 1 I 
current developments in the April-Salumei region of Ismi Scpik Province. No 
other classical locality in Papuasia is of such critical historical-biotic value, 
owing to the fact that ncarh ill o Ledermamn interio m h 1 ih Is If 
Kaiserin-Augusta (Sepik) Expedition fall within this n etui Veldkani] era I 
1988). Although key localities in the Hunstem Range were revisited by CSIRO 
botanists Hoogland and ( raven in 1 966, and by a National Geographic sponsored 
contingent in 198Q them is si ill no compilation oi urvivirm pec i me 11 horn tlu- 
now mostly-destroyed 1 d, im nm fi nor any published run pendi urn from 
the subsequent llunstein expeditions. 1 In the ineantiuK eon oi ignificaui 
discoveries have been recognised from the newer survc\ -• including species 
of Freycinetia from 1 he h\",0 expedition alone (Huynh 1999). The recovery of 
the endemic genus Scpikva, formerly known only from an illustration in 
Schlechter (1923), has also occurred in 1 e< ( m \< a J! I Pm 1 \v 1 <omin) 
I h clevt lopm m ■ 1 hhik n d In 1 nun < pi nm ' u 1 bu di o\, tn 
are reported in technical journals which are in an essible to government agencies 
or are d iscussed on I y wit h 1 n a small circle of botanical collaborators. The April- 
Salumei tract is currently a focal point of contention between conservation 
landowner, logging md miningirit icstsCcf. Bukl k H Piler&Sekhranl998), 
and a variety of future land uses is now under planning consideration. The re- 
sults of uninformed action in a locality with such unique biotic and scientific- 
historical values are potentially devastating. 

It is thus an imperative dice iiai: ingmec h misms Ix erected lor ensuring 
that surveys connect directly with the agencies responsible for priority-setting 

in wilderness territories. It is net suilieient tocstabhsh links between universities, 
herbaria, or NGOs with community-level jurisdictions over survey sites. These 
entities do not possess! lie statutory powers lor determining policies and priorities 
in the resource operations Uve,.. forestry and mining) winch have the greatest 
potential impacts on the environment. Without critical mputstothe planning 
facilities in government, surveys may end up as mere information-gathering 
exercises from habitats which subsequently disappear. At a minimum, floristic 
inventories should be consciously directed to the Forest Planning 
Division of the PNG Porcst Authority, and iodic Nature Conservation Division 
o( the Office of P.nvironment and t onsei \ation. 


in the last decade, a substantial amount of data has been acquired from many 
o.l PNOsloiesi environments. 'I he greater part of this work has been conducted 
i on k i ili. in ,pk i 01 I ' ,i i o|'k ralint u iilnn tin n u pecti\ < on s i \ it ion ai as 
A discouraging aspect of many such studies is that they are either never for- 
mally published 01 m In i \\ i t pp n hi id i i! n < i hnutid teadershipand 
di itibutton In tin lollowingdiscu sion. several di tinct but interrelated issues 
are considered, m some eases drawing upon data which arc available locally, 
but not readily accessible to 1 lie wider sc lent if ie community. The commentary 
addresses selected topics in L) morphospecies enumeration. 2) floristic rich- 
ness on environmental gradients, and 3) the relat lonslup between collections 
in Herbarium to timber concessional activity. 

1) Morphospecies enumeration. — Although the sice of lis t lora is ol con- 
siderable general interest, there is lit! leconsensuson the number of plant species 
within Papuasia (cf. Collins et al, 1991; Prodi n 1 984; Good I960; Hoft 1992; Johns 
1993; Womersley 1978). The only sure way of gauging the total floristic inven- 
tory for PNG is through systematic revision, but it will be many generations 
before a Flora Mu/e.simm-style compilation can be concluded (Geesink 1990). 

In spite of such concerns, the locality specific inventories are the most 
Minor! ni Din lor planning purposi While flora wide summaries may be of 
broad conceptual interest, they do not provide l lie sort of information which is 
relevant at operational lewis. Management act ions arc t vpieally evaluated and 
implemented lot pectfn localitu md th ; n em detailed information on 
the floristic profiles of individual tracts, liven if main plant families will re- 
main unreviscd ihrough the foreseeable future. 1 he enumerai ion ol Papuasian 
mot phospeciesand thedetei initial ion ol l heir spa! ial disiri but ions is an achiev- 
able objective. Knowledge of the local components ol i I mi istie diversity is a ba- 
sis tor rationalic d priori p citing, and decision in akin u i In le\el of ag,enc\ 
implementation. And ultimately it is the conservation ol local diversity which 
is the Inundation lor future i I or i si ie inquiry even t hough tin: larger regional 
patterns provide the basis for placing the local knowledge in context. 

Species checklists are thus of greater practical significance lor planning 
and conservation than is generally conceded, if they are used in conjunction 
with other information sources. 1 lowevcr. unless the compilations accurately 
reflect current taxonomy and are periodically reviewed and adjusted, they can be 
an actual disservice to govern mcni plan tiers and managers. In this connection, 
attempts to automate the process ol plant identification through interactive 
keys have promise as a heuristic and pi actical tool, although the methodology 
is likely to be const rai i ied I >v I i m i ted access to corn purer resources within coun- 
tries such as PNG. 

2. Floristic richness on environmental gradients. — The total numbei o\' 
Papuasian plant morphospcci enclosed within individual. 1 rritorie on 
elevational or horizontal environm* it I gradient i il > till unknown. Recent 
intensive surveys at ter Nit. have | I hhehigl ngh locality census 

thus far achieved in PNG (i.< 1 100 morphospeci< lakeuelu 1 999, 2000b) but the 
counts are not comprehensive. Inventories intending to c\ aluate rural i lorruic 
content within environmental !y \ iabl tract must < cut uallv qua.ntif\ species 
richness between habitats. A problem lor any manager working with limited 
resources is the question oi how to apportion survey effort through tune and 
space to a targeted flora. 

In the Neotropics the am mr to thi is u vould K i latively straightfor- 
ward fhe number of tree peck isi erselyrelal dtoelt tion, with the high- 
est counts in the lowland ; and with rich ne lading progu lively with altitude 
(Gentry 1988). When the emphasis is on evaluating th< tim floi Ka isu ,nll\ tin 
case with forestry opt ration ,) urvc\ would pro In- c maximized returns by 
concentrating at low elevations A salient qualification however, is that 
neotropical richnes i m t< U h h< ri h montane zone (cf. Henderson et 
al. 1991) if all plants i nn huh i nontrei peel* are con idered, so the optimal 
sampling plan lot urve) wouldl d \ 1 nt on the objectives. 

In Papuasian habitats floristic richness patterns arc more equivocal and 
complex than those reported, by (. .entry (1.988) and are generally consistent with 
Henderson etal. (1991 ) if the nontree component is considered. On New Ireland, 
tree counts have been reported i declinin monoiotucal h with elevation in 
the manner of the Ni otropi I ken hi i ] I9 ( However this result 

has not been replicated and may be an artefact ol widely separated sampling 
static n b( i b)U i \ | I i i in, u i pi mg methodology, 

concluded that the species richness curve lor New Ireland has a mid-elevation 
bulge, being highesi ai : >0 m and dnnuu bins ibow nd below rlia.r k:\cl 
Kulangetal.(1997)u n< .in d ni mminm i\ l n»ml >t I nOO m elevation along 
analtitudinal scquiiH , h vladang I'rovm < In a inula tudy, species diver- 
sity bom Ion I i ibil it m New Britain it un d maxi mini * aim hclw n 
600-800 m(Balun etal. 1996). Previously, the 1995 Bismarck-Ramu survey had 
determined the 600 m level as being floi'istically richest among examined sites 

(Hedemark et al. 1997). Although [he lowland montane ecotone has not been 
critically evaluated in Papuasi a, there are converging! i id ical ions that the point 
of highest florist ic development lies somewheie in or near that transition. An 
obvious implication for local consen tioi iniiiarivi i I hail his interval should 
din t il I) i ii il >re lor pro! d I itun urvcy can increase 

their efficiency by allocating more time lo i he low inoniane ecotone instead of 
altemptingequal coverage ol all habitaison an aliiludinal gradient. The lowland 
ramioresi and I fie high montane loresls are apparently less diverse, so survey 
o-l fort should h ,llo< an d an ordin; h 

The substantial variation (600-1,000 m ) in the elevations of beta diversity 
maxima suggests mi hi. no ol lo il fa< no l ; 'rom thedilien nces between sites it 
is also apparent thai the richest communities can be idem Tied only by actual 
survey, and not by extrapolation from results obtained m oi her areas. Site-spe- 
cific vegetation a 1 histoi ies.elmi._uie considerations including Massernerhebung 
(cf.Grubb& Stevens PXSn) subsi nil e distinctions, sampling methodology, etc., 
are factors probably responsible for l lie contrasts between locations. 

r.wn while the argument I rom raw n umbei ccm s clear enough Ik itu 
ation is still obscured bv considerations ol quality. Plant species do not have 
I he same value at hoist from conservation perspectives. The local endemic is 
understandably valued more than widely distribuied lava, so richness is only 
one aspect of site assessment. Due to dillicul ties in generating the required data 
there are no published accounts comparing eiulemism between different 
Papuasian environments. This is an obvious lac una which should be considered 
in future inventoric Uthough the unreviscd latu ol many plant families is a 
serious consti'aint, pivl i m i nar> esiunales could be obtained b\ using the lava 
covered in modern treatments. 

It is also appropriate to note that while previous el loi is haw alt erupted to 
quantify relationships between beta diversity and elevation, the critical variable 
is not elevation but I In Ion .tela ili< itionunii luwha er manner the forest/ 
community type is defined, whether by species composition, physiognomy, or 
a combination ol both it is ivm I be the lores! rvpe w fuel i underpins [he i ic hues' 
variation. Elevation is an obviousenvironmenial control but itseffectsare mani 
fested through the vegeiaiion lormation or 'life rone; and it is certainly not the 
only controlling factor. Knowing which forest formations have the highest number 
of species, in both absolute and proportional terms, is thus a more meaningful 
focus for inquiry than anv site specific relationship lot ween richness and el- 
evation perse This i t .pccialh 1 1 ut because tin .aim .peocsand or vegetation 
types often have dilierent clcvat lonal ranges al dillereui locations, which is at 
least partly responsible for the uieonsisieiii results obiauied by current inves- 
ligal ions on elevation dependent i ichness. (. m aneolotal grounds, it is apparent 
lorexample,that main' plant taxa havcanomalous low elevational occurrences 

on ultrabasics. There are also the Massernerhebung induced permutations, to 

ciii anotln i ob\ iom mlluun N< I iln m lui-'i din in< inhli.^ Mm n M >' 

! ,i i hi i km o in i« ' moil b< iv en ion i unit itIHuii in Jim i ii\ i . .mi 
plicated by the cvistMK < » an ihoiin.i-iinv^ i in f >i foi est classification 
in Papuasia (i.e., Hammermast i & Saunders 1995; Johns 1977; Paijmans 1975, 
19/p Sjuncii t- I^Gj [ In In i -,< lirnu how -m i i , now mpportcd by a GIS (Geo- 
graphic Information System) augmented b\ mm imum ovci lays foi the pic- 
existing 1:500,000 scale topographic maps (Australian Survey Corps) and also 
with separately issued 1:100,000 scale maps showing the vai tons loivst unit::-, 
foi ill of PNG. It rhu provide a very useful foundation loi planning and execut- 
ii ; iln in n \ needed to i ob < nut r i i u< 

3. The relationship between collections in Lae Herbarium to timber 
concessional activity.— Recent summaries (Balgooy et al. 1996; Welzen 1997) 
fi mi the I Umi Mah s'utmt ugg< ;r that Ni v « ininc a h is tin high. I < m ol I I 
ristic endemism in Malesia. Based on current revisions, most of the endemic 
species appear to be concentrated in montane habitats. I Ning selected genera, 
Heads (in press) arrives at some of the same conclusions regarding relation- 
ship, between montane environment; mdi ncleiui ,m 

( oIl( noc .u ii a i in Papua aa are veto eh rh keved in i i\or o\ mon- 
tane areas (Conn 1994), so to a certain extent phytogeographic summaries will 
be affected by the sampling inequalities. The greater pan o( the LAE holdings 
(and thus the overseas duplicates resulting I rom r hem ) originate from the 
Mamose 2 region and the Highlands U.:i lag 1) ,o it is inevitable that a certain 
bias has been introduced into distributional summaries derived from such a 
foundation. The magnitude ol such biases cam be mi erred i rom the geographi- 
cal unevenness ol < oll< < i i in the Lai Herbarium i i m mar ized in Table 1. 
I he i oie ol tin PNG national < )l lection in botan\ con i i oi the NGF 
and LAE series specimens (1-49,999 on the New Guinea Force numeration and 
thereafter with higher numbers on the LAI: sequence,). I he institutional sets 
currently end at ca. 85,000. Both the NGF and LAE sequences have many blank 
inn rval i on inn ii mini 1 h r blocks u hiNi urn i i d i us oil, t loi 
but never actually u <l \ ubstantial amount of niaaei lal was also rejected or 
destroyed after then )l lection tad been recorded Back numbering of contem- 
porary gatherings has been ein ployed in an attempt to fill in these gaps but 
there are still much I ever col lections than indicated by the institutional num- 
ber. Table! is thus based on a manual count of archived labels physically repre- 
sent! d bo con [-on hn |i i inn n nil hi i b. mini 

The NGF and I \l .erics constitute approximate^ >5% of the estimated 
275,000-300,000 specimens in the national herbarium, though it should be 

Fig. 1 . Provinces of Papua New Guinea. The Western Highlands are composed of Enga in the w 
Eastern Highlands are divided respectively into Chimbu (Simbu) and Eastern Highlands Provinc 
to the Island Arc terranes (Pigram & Davies 1987). 

r.The Southern Highlands have geological affinities to the Australian cr 

North Solomons (Bougainville 

West New Bi 
Subtotal Is 

I ,isi Srpik 



Subtotal Papuan Austrocraton 



noted that the size oi the total holdings is not accurately known and tends to be 
overstated because of discontinuities in the recording svstcm. A major data re- 
covery effort would he required to collate locality information on most oi the 
remaining collections. The notable exception an tie arh ( S1RO sets obtained 
by L.Craven, T.Hart lex P. I leyliger; R Hoogland K Paijmans. and R Scliodd. 
The LAE duplicates by these codec tors are entered in I iei barium logbooks and 
have been easily incorporated into Table 1. .Approximately Ql.OOO specimen 
numbers are included in the combined tallies (Table 1). 

I he obvious preponderance of specimens Irom Morobc and adjacent prov 
inces, and the opposing paucity ol material I coin Western and Gulf, are apparent 
even from casual inspection of the national collections. With 28% of the col lec- 
tions, Morobe Province is ver\ disproportionately represented. The disparities 
clearly reflect the concentration of Forest Service facilities and infrastructure 
at Lae and Bulolo. A similar situation is shown by the relatively high numbers 
oi' specimens from Central Province, especially from the Brown River and the 
Sogeri areas, owing to then proximity to Port Moresby. 

If timber concessions are tabulated bvprovi nee and region (Table 2), a general 
i n verse relationship is ev idem between the disi ribnt ion oi existing concessions 
and the collections coverage o( the corresponding areas (lag. 2). The provinces 
most susceptible to logging im pacts t Islands legion and I lie Austro-geoprovince) 
have the fewest herbarium specimensas reflected in the LAP sample, while the 
best documented regions (Mamose and 1 lighlandsj have relatively little logging 
activity. This inverse relationship shows that future lorest sector development 
is likely to be most intense in areas which are florist icalK the least known, and 

t n r i n I in is in Papua New Gui 

Total Concessional A 

New Ireland 

North Solomons (Bougaiir 
West New Britain 
Subtotal Islands Regio 

Subtotal Mamose Region 


Subtotal Papuan Peninsula 


rmihcr H'qhrrds 


Subtotal Papuan Austrocraton 

SI Ah[)S R! dlflN 

may thus involve considerable biodiversit y r isk. 

Document at i i I i ni n .lit whei 1 

cessional activity is curt mho > irrin or imminent -(.ch Fie \, reproduced from 
PapuaNewGumea I cne .1 Aailioi n \ 1.998). l ; lonsti< latai required not only to 
rationalize the loggim pi n lot tfiectedan o.,butal i urn sessthepostharvest 
consequences oi foi c i felling Die opportunity lord' o\ rmg localized species 
and of record i ng popular i onal variation eoi ild be oilier wise irretrievably lost. 

Judging from historical pattern of collecting and future lorestr\ iced th< 
highest-priority documentation targets should be the lowlands of Papua and the 






Austrocratonsubrcgion, particular!) (hull Province. This does not necessarily 
mean that such environments are more speciose or significant, only that the 

flora I hen is comparatively less explored and at greater risk of alteration before 
i hi been dcx i n Us d. P> .1 on I • \< Britain should also he a priority 
i ; i Inn nun h 01 lie island ha been previou y log; d and taxonomic losses 
have presumably aheady occurred as a result oi forest removal (oil palm plan- 

ronmentsare primarily in natural growth. 

Papuasian bioinventories involving exped 
trained specialists are relicts from a bygone e 
within a rapidly evolving PN(d indicate the appropriateness for change, even 
though the manifestations oi such need are obscured by the misconceptions 
imposed by international and cultural distance, h.xisting failures in documen- 
tation are certainly not attributable to a lack o\' collective scientific interest or 
dedication of past workers. Thcdeticienciesare pnneipath. methodological. The 
evidence of the last 100 years shows thai conventional it i nerariesare not going 
to achieve adequate collect ions sat una ion oi critical environments within accept- 
able time frames. As long as the burdens oi inventory are borne primarily by 
an elite professional corp thedocum mat ion of PMO bona richness will con- 
tinue to bean elusive objective. I he human assets lor survey must be applied in 
more effective ways than previous programs or the del leieneies will peisisi 
I mloi innately lime is running out. 

Past attempts at floristic inventor)' have been imsiiiut lonallv eetii rah. sal, 
expeditionary bnel, logistically intensive, and with participation by a select 
membership, butun opei it ion will need to beconx Icceritralized, continuous, 
participator) and effected primarily by personnel that are preferably actually' 
living at the sites being subjected to bioe\ a 1 nation. Parataxonotmsts and the in- 
frastructure to sup] xn I them, are necessary elements m ae h icving such outcomes. 
The new schedules will also have to be acutely responsive to emerging grass 
roots assertion ol community ownei hip ri In uullhen iiltmgdem no lot 
stakeholder participation m all activities mvoK ing customary resources. Un- 

loss lin u i i in i Jen n i 1 m 1 i md ow h ml o i 1 1 it k iht okijt i 

tives outlined by van Steenis half a century ago are unlikely to be realized. 

The senior author i hanks Yojtech Novoiny (Institute ol Entomology, Czech 
Academy of Sciences) lor providing access to grant proposals and supporting 
documents. Many suggestions made h\ Pi: Novotnv were incorporated into the 
final draft. Michael Heads (University o\' Ooroka) also shared unpublished in- 

formation. Assistant Director Barney Lipscomb (Botanical Research Institute 

of Texas) extended his usual snppoir ,11 nl ,1 , istance ( ulli ague Hitofumi Abe 
(Ecosystem Research Group, University of Western Australia) provided the 
Japanese translation. The PNG National Forest Authority gave permission for 
reproducing data from Service files. 

We acknowledge referees RE Stevens (Missouri Botanical Garden) and J. 
Pi poly III (Fairchilcl Tropical Garden) lor their characteristically thorough cor- 
rection of the manuscript. 


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Pp 363-387. 

he floia of Papua New S.uin 


Paul M.Peterson Yolanda Herrera-Arrieta 

Department of Botany CIIDIR Unidad Dun. mgc 

COFAA-lnstituto Politecnico Nactonal 
Smithsonian Institution Apartado Postal 57 

Washington, DC 20560-01 66, US. A. Durango, Dgo., C. P. 34000, MEXICO 

\BS1 RA< r 
Vfu/iitnhergici includes 1^1 pecic .Inmil-V World origin; I 33 speci ire indigenous to North 

\menca|althou lin.un ol tlu t i m t( m i i\ \ m . ih i ) ml outli \ , k k u 14 i| ] i 

occur in Central Ameriea i a in< k P < L u , , ndmiu J > . s pil i s ( ,uui in south America (10 are 
endemic) andonh mm nduiik ,h,k u 1 it . ,i i mm »ui Im n \ 1 1 No modern subgeneric 

1 'I' i i m I m tl nu il| i i ' n n hi] u n I I i \n in i mi 1 i i 

ol the leaf blade as viewed m transversa euionl provided a unique el ol 1 charactei i>u i 
previou Inpotlu t 1 , I i , n I p 11 , ,, t |„ t |, UIU u| n 

ibleintothut mijoi imiomi il,i ip , , , dn 1 r \ ul»,, nu u \f subg \hthhnh> ni 
and Trichochloa) and two sections t;\ I ,,im, , imllWn.viiiiiiiUiuM ubg. InYhoAdns 1 en 

arias principalm 

i u i< u pai t probai las re! i.inn. Im >n u, i 
1. u In, u ou d( lod in ,< | i u, I , 
nit/ ' ( n n i ( divi A ilu ii K 

- I \! -ail.f \/l(lli, ll/'l LM,1 \ ll (< hth r',/0,1 \ d<' 

ubg. In. /iuc/iW Aim cuando la prcsencia de 
In u.lm ion d< I i^ , pout d, Midi/, „!., i in 

AMeAvumiel subtipodefotosin 

1 he Mihi ri be M uhlcn boi y, i mac t Poaceae: C h loridoideae: Eragrostideae) was first 
circumscribed by Pilger (1 950) to include only species of MuMcnbergiaSchreb. 
with narrow single-flowered spikelets, firm glumes often shorter than the 
awned lemmas, and cylindrical caryopses. In this same treatment Pilger recog- 
nized Epicampes}. Presl \-Muldcnhergia subg. Tvichochloa A. Gray, M. sect. 
Epicampes (J. Presl) Soderstr.] in subtribe Sporobolinae Ohwi. Pilger further di- 
vided Muhknbergia into eight sections: Auvms (Trin.) Bush, Beaha (Scribn.) 
Pilg., Cinnastrum (E. Fourn.) Pilg., Clomena (P. Beauv.) Pilg., Muhknbergia, 
Podosemum (Dcsv.) Pilg.. Pscud.'NpomJ'o/u.s Parodi, and Slenodadiurn (Trin.) 
Bush,. Subsequent authors have agreed that Pilgers ml rageneric treatment o( 
Muhlenbergia was not particularly phylogenetically inlormative (Soclerstrom 
1967; Pohl 1969; Morden 1985: Peterson and Annable 1991). More recently the 
following six genera haw been shown tosharc common ancestry and have been 
placed in the Muhlenbergiinae: Bealia Scribn.. Blcpharoneuron Nash, 
Chaboissaea E. Fourn., Lycurus Kunrh, Muhlenbergia. and PereilemaJ. Presl 
(Duvall et al. 1994; Peterson 2000; Peterson et al. 1995, 1997). 

Many agrostologists have elected segiegate genera to emphasize critical 
li tun, m iln lai aid divei i gem \i i/i/t n|i< rgm I >■ van •, i 181.0) n < oj' 
nized the genus Podosem n hi. based on the caespitose. open panicled, and long- 
awned M. capilkiris. Palisot de Beauvois (1812) described the genus Clomena 
based on the annual M. peruviana, and Presl ( 18 >0) described Epicampes based 
on M. robusta. Two relatives of the type species of the genus (M. schreberi), M. 
glomerata and M. andina, were given generic status by Link (1833) as 
Dactylogramma and by Thurber (1863) as Vasexa, respectively. Nuttall (1848) 
described the genus Calveodon based on the widespread and often important 
range grass, M. mom an a. The only other generic name given to a species pres- 
ently placed in Muhknbergia is Crypsinna, described by Pournier (1886) and 
based on M. macroura. Hitchcock's (1935) transfer of many of these segregate 
genera to Muhlenbergia has been Followed by most American and European 
botanists. I he morphological characters ( hat delimit the genus are spikelets 
with single perfect florets and hyaline or membranous lemmas with three usu- 
ally prominent veins. I hese characters are not at all unique within the Eragro- 
stideae and seem to be possessed by about hall ol the genera in the tribe. 

The morphological di\ersiiy within Mulilenbergia is t rcmendous. Annuals 
less than 2 cm tall [M.depauperata.M. mi nut issima.M. peru \>iana,M-ramulosa) 
are not uncommon and there arc numerous si rough, caespitose perennials over 

2 m tall {M.gigantea.M mutu .a, M. robusta). Rh monies and ,/or stolons are found 
in 1/4 of the species and there is a single species (AT dumosa) that has a growth 
form similar to bamboos. 1 eal blades can be flat, involute, or folded with a variety 
of pubescence types located on iheabaxial and/or adaxial surface. All species ol 
Muhknbergia have open or contracted (spike-like) panicles with the branches 

generally re-branched. At maturity or anthesis. the angle of the branches 
spreading from the culm axis and t he total width of the inflorescence are diag- 
nosiii < h rat t< ri ti< used to < p i in ih p C ( i< P< di el on nr an n< m \ tin 
from appressed or spreading to nodding and reflexed bom the branches and 
the pedicels can be either lound or flail ned in ros eel ion. Most species of 
Muhlenbergia have m i uK ilmuand ph M i aUhou Inbu rre two species that 
are occasionally 2 or 3-flowered >M. aspei i folia and M. uniflora). The lemma is 
perhaps the most u us il iniiiiin imln ! mm u< h i , length, presence or 
absence of an awn or mucro, pubescence type and location, shape, and color 
can all b< i . d tmdiik u nnin nmi i tin p. u ' In im k I, mm m > \ t in.J 
(f -veined in M.palm i rc/isis J with a si our men rial vein and two lateral veins, although 
the lateral veins are sometimes very 1 la.rd i m d seem wit 1 i a good (20X) dissecting 
microscope. The caryopsi has a fused pericarp and is u ually free from both 
the lemma and palea in mm tspe< iei ml Muhlenbergia however toe length, shape 
and to h .(it ti m - ul )i u i i i ibl 

At 1 i i talb Muhlcn cd of lil sp Peterson 2000 I h 

distribution of Muhlcnba mi ilrnmsr entirely \ev World where 133 species 
are indigenous to North America [all hough many of these range to Central 
America (33) and South America (14)1; 38 species occur in Central America (a 
single species is endemic J: bm species occur in South America (10 are endemic); 
and only six endemic species ace known ro occur in southern Asia. One obvious 
hypothesis is that the genus arose where it is most diverse today i.e., northern 
Mexico/southwestern U aim hi nc idiatcd hoi i dispersal event, the 
longer t lie distance from i he place of origin, would in theory, lessen the chance 
mi isuecessful introduction fnereioi bun ire rnan\ speciesof Muhlenbergia 
m Nmith Mimm a kvm in' . nil il \im mm . \ t n u v i in mifli Mm n, i ,uiA 
Imalh' very lew in Asia o la ill ten peei n thi ubi nbe Muhlenbergiinae 
that have been investigated genetically (Pctei on md 1 1 rrera A 1995; Peterson 
and Morrone 1998; Peterson and Ortiz-Diaz 1998; Peterson et al. 1993; Sykes et 
al. 1997) exhibit a norl h to south migration pattern, including Muhlenbergia 
torreyi (Peterson and Ortiz-Diaz 1998). 

All species of Muhlenbergia previously examined exhibit kraim Um ) leaf 
anatomy, particularly the parenchyma sheath subtype which is common in 
species occupying the most arid regions (Brov nl9/ Mates hey 1984; Hattersley 
and Watson 1992). Two main subtypes, NAD-ME (nicotinamide adenine dinucle- 
otide co-factor malic enzyme) and PCK (phosphoenol pyruvate carboxykinase) 
have been found, verified by biochemical assay, to oi cut within Muhlenbergia 
(Gutierrez el al. 197 A. 1 lattersley and Browning 1981; Brown 1977; Hattersley 
and Watson 1976, 1992). These two biochemical subtypes differ in their predomi- 
nant C4 acid that is transported from primary carbon assrm 1 lation tissue (usually 
the mesophyll) to the photosynthetic carbon reduction u'CIA tissue Usranz 

sheath - parenchyma bundle sheai h ) bee I larrcrslcy and Watson 1992]. There 
usually is an associated anatomical structure in PCK-likc species that is diag- 
,„, iu mill Km c ji i una m. i ioi > hlou iu fn in i n in i ontinuou between 
adjacent vascular bundles. In typical NAD-MH species the chlorenchyma is 
tightly radiate and usually separated from one vascular bundle to the next by a 
column of colorless eel Is 1 hesc diffcicnco havt hi lorn ill\ been used to separate 
these two subtypes: however, it has been shown in ioiueopoyon Desv.ex P. Beauv, 
Eragrostis Wolf , hAmc/ine R. Br, Panieum I.., PrinkdorMou S. T. Blake, Triodia R. 
Br, and Triraphis R. Br. that these anatomically PCK-likc genera are actually 
biochemically NAD-ME (Ohsugi et al. 1982: Prcndergast et al. 1986, 1987). 

The first majorauatomical investigation of MuMchIvivm wasdone b\ 1 lohn 
(1901) who looked at 10 species and was able discern three groups: woodland types 
(=M.subg.Muhlenlvrgm):diy, rocky mountain slopes A M.siibg. Trtchoch/ofl, sect. 
Epicampes and other species]; and I ipes (= M. subg. Trichoch loa, sect. Podosemum). 
Holm (1901) pointed out that from an anatomical view point these characteris- 
tics might prove useful in dividing Muhlenbergia into sections or subgenera. 

Schwabe 1.1948J later investigated 22 species ol Muhlenbergia that occur 
in South America and found that they correspond to four major groups: hygro- 
phvies or mesic species u hi. subg. Muhleuberyju). \erophytic annuals one, 
xerophytic annuals two ( M. subg. Muhieubergia)^ xcrophylic perennials, and 
psammophytic perennials (= Msubg. lin Inn Mom sect. Podosemum). Schwabe 
(1948) also suggested that Muhleubergui should be separated from the genera 
ol Agrosrideae and incorporated into the I ragrosteae. 

On the basis of leaf blade transectional anatomy and morphology 
Soderstrom(1967klistuigLushed two subgenera in hhihlenbergiaMuhknbergiti 
and Podosemum (= Trichochloa, an older name), and divided M. subg. 
Trichlochloa into two sect ions, sect. Podosemum and seel, /m t cum pes. Soderstrom 
placed lOspcciesol Muhleubergui, which luivc parlially sclerosed phloem and 
caps of sclerenchyma associated with the primary vascular bundles, into M. 
subg. Trichochloa. Muhlenbergia sect. Epuampes was characterized as having 
a compound keel Unidvem) composed ol primary and secondary vascular 
bundles sunken in a confluent mass ol thick-walled parenchyma, whereas M. 
sect. Podosemum had a simple mid vein composed oi a single primary vascular 
bundle with additional iertiai v vascular bundles present iSoderstom 1967). 

Two years later Pohl (1969) completed a revision of 12 closely related species 
that he believed to represent the entire M. subg. Mu h len hergui in North America. 
Principal differences among the species in this study were size of the bufhform 
cells, size and degree o\ radial orientation of the chlorenchyma, and the extent 
low hi hth chlorenchyma w or; iu d ntodisi tc units surrounding each 
vascular bundle. Using morphological characteristics of the rhizome (posses- 
sion of very short iriici nodes with imbricate scales) and leaf blade (thin, flat 

blades w irh low k i; ill v idth i mo [\>hl ( Ji nngui I eel tfu pccn h .in 
others in the genus. However, these same characteristics are seen in M. 
califormca, a species of the mountains and valleys oi southern California, and 
many other species common to the soul h western United States/Mexico. 

Morden and I latch (1987, 1996 J invest igated the anatomical and morpho- 
logical variation within the M. rcnenscom pie x, which consists of six species in 
North and South America. Anatomical data supported the placement of M. 
squarrosa (Trin.) Rydb. as a synonym oi M. nchardson is, and supported the rec- 
ognition oi two varieties oi M. viUiflora. 

Peterson et al. (1989) and Peterson and Amiable (1991 ) investigated 29 annual 
species of Muhlenbergia and found 14 characters useful in distinguishing four 
major species groups. Species oi group ( 1 ) had flat blades, two tertiary vascular 
bundles between primary vascular bundles, vascular bundles positioned on 
the median layer, distinctly radiate and compact chlorcncln ma cells sepauu i ng 
adjacent vascular bundles, and Ian-shaped central bulhform cells. Species of 
groups (2, 3 & 49 shared three characlerist ies: I > indistinctlv iM incom pleieK 
radiate and loosely arranged elilorcnehyma cells, 2) chlorenchyma cells con- 
tinuous between vasculai Hindi nl lucid shaped < nil I bulliloi m lb, 
(= M. subg. Muhlenbergia). 

More recently a biosystematic study investigating the anatomy of the M. 
montana complex (consisting oi 15 species) has been completed (Herrera- 
Arrieta 1998; Herrera-Arneta and Grant 1 994). Herrera-Arneta and Grant used 
18 characters to dil I e rent lai c among t hew species and louncl lour major species 
groups. Important characters appear to be the central midrib structure (similar 
insizetotheothci pium \\ ,1 lid, i pi rnceo a prominent central 
midvein), the depth oi rheada\ia and abaxial furrows, sclerenchymatous girder 
development between the parenchyma bundle sheath and the epidermis, and 
epidermal vestiture (glabrous or papillose). 

Leal anatomical characters within the Poaceac as viewed in transverse sec- 
tion have long been recognized as important diagnostic features used to deter- 
mine systematic relationships, and haw been critical in elucidating infragenenc 
relationships within Muhlen bergia (I lerrera-Arrieta and Grant 1993; Holm 1901; 
Morden and Hatch 1987; Peterson 2000; Peterson and Amiable 1991; Peterson et al. 
1989; Pohl 1969; Soderstrom 1967 J. A preliminary summary of our anatomical 
analysis is presented in Peterson (2000) where two subgenera, M. subg. 
Muhlenbergia and subg. Tnchochloa, and a possible third group, 'Clomenti are 
recognized. In this current paper we will give a detailed summary of the ana- 
tomical features as viewed in cross section oi 148 oi the possible 151 species 
within Muhlenbergia and present a subgenenc hypothesis that most closely 
reflects a cladistic analysis ol the data. I his is the first anatomical survey of 
nearly all species within this large variable genus. 


Over the last 16 years fresh field collected leal blades were obtained from North, 
Central, and South America, as well as China for anatomical study (.Appendix 
1). Five mm long leaf blade segments from the central third of" the mid-culm 
region were fixed in l T AA (.10 parts FtOI 1; 1 part glacial acetic acid; 2 parts 37% 
formaldehyde; 7 parts distilled water). A lew species (less than 5%) were studied 
from dried herbarium specimens because fresh field collected material was 
unavailable. Leaf blades were first desilicilied in 100% hydrofluoric acid (HF) 
for 48 hours in order to ease microtomy, then dehydrated using 30, 50, 70, 90, 
95, and 100% (twice) ethanol, graded into xylene (twice) and transferred to 
xylene: paraffin oil 0:1, steps 1 hour minimum). Blades were then dehydrated 
by using 2-2 dimethoxy propane iHMP). acetone, and tertiary butyl alcohol 
(TBA) series while in a vacuum. Infiltration was accomplished using two, six 
hour minimum changes of liquid paraflin before being embedded. The tissue 
was softened using 95% LtOH: Glycerin: lib (.8:1:0 to improve slicing (Foster 
and Gifford 1947). A standard rotary microtome set at 6-10 :m thickness was 
used and sections we re stained wathsafran m /last green or 0.05% toluidine blue 
(Berlyn and Miksche N76). Samples were examined and photographed on an 
Olympus BL1-2 photomicroscope using Kodak TMAX black and white or 
Fktachrome color slide film. 

A natomiciilclescript ions were coin pleledli>llowiiig the procedure for stan- 
dardizing comparative leaf anatomy in grasses as out lined by Hi Lis (1976). For 
purposes ol com pari son and standardisation, primary vascular bundles (1% are 
delined i oniainin i i icta vlem lem nt on either side of the protoxy- 
lem elements with additional ksigenous cavities, and are usually associated 
with sclerenchyma girders or strands ( 1:1 lis N70: Peterson et al. 1989). Secondary 
vascular bundles (11% resemble 1° vascular bundles by having distinguishable 
xylem and phloem but lack large metaxylcm elementsand Ksigenous cavitities. 
Tertiary vascular bundles (111°) contain indistinguishable xylem and phloem 
areas and are usually smaller than the hand 11° vascular bundles. 

A list of the anatomical characters and their states used in all ensuing analyses 
appears in Table ha list of specimens used in this study is given in Appendix 1; 
and a complete data set is given in Appendix 2. The 15 5 taxon by 16 character 
data set was analyzed with WmClacUOOO. Parsimony heuristic analysis was 
performed with NONA (Goloboff 1668; Nixon 1666). We used 10 random taxon 
order replications in NONA, with TBR swapping holding 20 trees, followed by 
TBR swapping (to completion) holding up to J 00 trees. A hard collapse of all 
unsupported nodes was selected to produce the cladogram m Figure 17. Since 
we were not testing I oi' monophvlvol 1 1 ic genus, the out group species were con- 
strained, i.e. .there were nosvnapomorpliicssuppoi t mg the Muhlcnbcrgia elade. 

1. Adaxial furrow depth: 1 = <- 1/ stud 1 I i ! , i u , I or more than blade avei 

2. Primary vascular bundle shape: I [minded I n II p i ! = v, uiihiui it Ihi i inluuilii in 

detenpinethe h tin l'i iin'i'l'- ill. i hi m tn in i I i in tt Hinrimg each vascular bundle. 

3. Vascular bundle outlined ' |i,i nh i il i tli i re as secondary and tert 
vascular oundiey; 1 = subequal, seconcar y and te-t'a'v v.e,; .1 ,i' :• r Vies -1 '5 the V/ iranar/vasculai honor, 

] eijuil ) 1 a i 1 1 a 1 1111 1 rt i| 1 ii ] ciilar bundles. 

4. Median (keel) vascular bundle structure: = t|. - 1 1,1 1 1 1 |nt i, . i . n t I 
pound keel, a sttiijie primary vasctiku bumile with 01 ly bviuddihniial letttaiy vom iiImi liund es, '] = complex u 
poind keel, with dime or more addlinnal p-nn-irv, -,er onr :. u v. .r:ti/u> unitary vne iiIji bundles Instate 1 theeun 
associated parenchym 1 I in 1 ti ml ( I m 1 d n m ith t| 1 I 

5. Vascular bundle position: 1 = one level, centered at j 1 1 ;■ il^nl r uln nl \uiface 2 t i >t tl 
levels, usually closer to the abaxal surface. 

two or more large metaxylem e 
and has distingu 

■1 ll''yij : v\i:i'i> 

7. Chlorenchyma arrangement: 1 nd b 1 uupi t m M, nt mqed 1 ib I mm p m 1 ill 

h i ih il n j n. tnn ti i mi ,t i l I iln mi i iihiln i || 1 1, ii 

PCK [ihi w bine the chlorencbyimi \ 1 obin hi I - - . 1 n I i i, nt vascular bundle. 

8. Crown of inflated cells (adaxial) in primary vascular bundles: 1 = absent 2= pit n Tl- * i, 1 

usually composed of parenchyma or (olk'ti , nbl Iruh' -m uim, lie ,<leienchyma cells. 

1 = circular orittei I 1 h | „ , tbti hi f ndtn h I 1 \ 1 

it 11 1 1 r idavial and abaxial si 

■cular bundle. 

mdary and/ortertiary vascular bundles b 

Allautapomorphic; were not includi ! in tin final < ladograrn because the\ add 
no additional information lor inferring relationships anion g two or more taxa. 


The results and discussion are interpicted in two parts: i ) a general description 

combining all species of Muhlcnbcr^uu and 1) results of the cladistic analysis. 

General Description of Leaf Structure. — Lamina (blades) are sometimes 

undulating, to more common h flat outwa li\ bowed I, , commonb in\o 
lute (Fig. 3), or folded (Fig. 4 J. The angle formed by the two arms is broadly V or 
U shaped to expanded, occasional lv loosely involute. Ada. via I kirrow depth in 
comparison to the leaf thickness can be slight, shallow (< 1/5 leaf thickness, 
Figs. 1, 11, 16), medium (1/5 to 1/3 leal thickness, Figs. 2, 13), or deep (1/2 leaf 
thickness, Figs. 3, 10). and m the form oi clelts located between all vascular 
bundles Adaxial tib irceommoiil on niopposii ill i -nil ai bundles (Fig. 
2), the same size to general b .mallei than ibaxial rib to less frequently absent 
with shallow groves opposite the vascular fmndles (lag. I). Primary vascular 
bundleshape vane Atom ion aho 1 1 i 1 J loobovan ellipiiclFig AlOi 
or rectangular (lags. 5. I 3, 1 4): secondare and tertiary vascular bundles also ex- 
hibit the same variation in shape. The secondary and tertiary vascular bundles 
are generally of the same size as the primary vascular bundles (Fig. 4), to about 
4/5 the size of the primary vascular bundles i, lag. 5), or very unequal, less that 
2/3 the size of the prirnai y vascular bundles (.lag. P.). Abaxaal projection of the 
median vascular bundle or midrib, msed b\ clerenchyma is omelimi laigs 
and witha protruding ridge (lie , u oncon picuous, often flat to round (Fig. 4). 

and adaxial fibers, and intercostal sclerenchyma. 3. M. dubia with an involute lamina, deep adaxial furrows (1/2 or 
more than blade thickness), obovate/elliptic vascular bundle shape, adaxial crown of inflated cells, abaxial inflated 
cells, simple keel with only a single primary vascular bundle, tertiary and secondary vascular bundles <2/3 the size of 
the primary bundles, with secondary and tertiary vascular bundles positioned c 

two layers of continuous abaxial sclerenchyma, and a sclerenchyma cap along the blade margins. 4. M. brevivaginata 
with primary and tertiary vascular bundles about the same size, a simple keel, rounded vascular bundles, an undiffer 
entiated median vascular bundle, centered vascular bundles, and 13 vascular bundles per blade. 5. M.pubigluma with 
secondary and tertiary vascular bundles that are about 4/5 the width of the primary bundles, two secondary or tertiary 
vascular bundles between each primary bundle, rectangular vascular bundle shape, centered vascular bundles, tightly 
radiate chlorenchyma,a column of colorless cells separating each vascular bundle, sclerosed phloem, parenchyma bundle 
sheath extensions, one or two layers of adaxial sclerenchyma development in the primary bundles, and discontinuous 
abaxial sclerenchyma development in the primary bundles. 6.M. nigra showing obovate/ellipticvascular bundle shape, 
tertiary and secondary vascular bundles that are <2/3 the length and width of the primary vascular bundles, tightly 


japonka with an abaxial enlarged and bulbous projection of the median vascular bundle, a complex compound k< 
consisting of a primary vascular bundle and two tertiary vascular bundles with associated parenchyma, with or 
mary and tertiary vascular bundles, loosely arranged chlorenchyma, and five tertiary \ 

The median vascular bundle or midrib is a simple keel (.big. In consisting; 
of a single vasculai bundle with oui issociatcd parenchyma a compound keel 
(Fig. 8) consisting of one primary and two secondary or tenuis \ vascular 
bundles with associated parenchyma, or; u o triplex compound keel (Fig. 9) con- 
sisting of three 01 mon priman secondare a\v\ or tertiary vascular bundles 
with associated parcnefn ma. All vasculai bundles are com mon 1\ situated w ith 
their position in tli. m diau lau i oi t In nib. anudui iiu nomtli. 
adaxialandabaxial leai urface(Fig: ' 1 >!l 1 3j. or are occasionally closer to 
the adaxial surface (lis > ), 10) at two or thn le\< Vascular bundle compo- 
sition consists of primar\ nd econdary only in thi am. blade (Fig. 7), only 
primary and tertiary in the same blade (bigs. 1. 8. 1 1 ). The presence of primary, 
secondary, and tertiary vascular bundles combined in one blade is not as com 
mon (Figs. 3, 6, 10). The total number ol primary vasculai' bundles varies from 
5 to 15 (four in M./t/sl i^niid nun in \/ puuciflont I >in M.^igantca). Secondary 
and/or tertiary vascular bundles are arranged m a regular fashion between 
consecutive priman vascular bundle and the number vans Ivi wen I- la 
3, 4, 6), or 4-8 (Fig. 8). 

The chlorenchyma tissue consists of two major types (arrangements). It 
can be composed o I mgl< radial la\ roftighrb pael d tabular cells that 
surround each vascular bundle [NAD-M b cen tri pel a 1 1 y ] nsttioned photosyn- 
thettc carbon reelru rum !'< R; cell i hloropl ist XyMSa- md F( V « II outliu. 
that are even in tran vei e section ,ec I laliei ley and Watson F>P] and i epa 
rated by uni-, bi- or tri serial columns ol colorless bulliionn cells (Figs. 3-6, 9, 
10, 12-15). Or it can be composed of tabular cells that are indistinctly radiate 
and continuous between the bundles [PCK type, defined as centrifugal/evenly 
distributed PCR cell chloroplasts (with grana). XyMS+ and presence of PCR 
cell wall subenzed lamella, in Hattersley and Watson's (1992) sense] (Figs. 1, 11, 
16). Colorless cells (lag rib, 13) an mailer oi imilat it ar.< md hape to 
bulliform cells and are often inflated. A crown oi ml lated cells is sometimes 
present over the primai y \ nl r bundk on tin. id nib urtaci md thesi 
inflated eel Is can be ion i id over the secondary vascular bundles as well 'bags. >. 
6, 7, 10, f 4). Inflated ill ometinu < n b found e| uat ing the primary vas- 
cular bundles from the epidermis on the abaxial surface 1 1 Igs 3, 6, 10). Strips of 

lore than the blade thick 

f inflated cells, abaxial inflated cells, and 

nchyma in the primary vascular bundles. 11 . M .schreberi wil h 4 - 1 1 primary vascular 

h tertiary vascular bundle, shallow adaxial furrows, centered vascular bundles, loosely arranged 

■shaped bulliform cells, and a single layer of adaxial and abaxial (discontinuous) sclerenchyma 

of colorless cells separting the two vascular bundles. Scale bars = 50 i^m; symbols as follows: b = bulliform cell; cl = 
chlorenchyma, mx = metaxylem; p phloem; ps = parenchyma bundle sheath; s sclerenchyma; sp sclerosed phloem. 

Figs. 13-16.Leaf blade anatomy of Muhlenbergia,adax\a\ surface uppermost in all pi 
adaxial furrows 1/5-1/3 the blade thickness, rectangular vascular bundle shape, c 
rosed phloem, tightly radiate chlorenchyma, a column of colorless cells separating each vascular bundle, three 01 

rectangular vascular bundles, sclerosed phloem, tightly radiate chlorenchyma, an adaxial crown of inflated cells in the 
primary vascular bundle, abaxial and adaxial interrupted parenchyma bundle sheath, abaxial girder of sclerenchyma 
m cells, three or more layers of adaxial sclerenchyma, and mam 

interrupted parenchyma 

well-defined and regular hullilorm cells are present in the epidermis and are 

distinct i nan normal epidermal cells. Bui I i form ceils can he closely associated 
with the colorless cell Bullilorm and i oloilcss cell iocel her lorm the urn . hi . 
oi tn-seriate columns which extend I rom the adaxial I arrow ,,, the ahaxial 


epidermis separating the vascul u buiulli (I ig^ 5, 1 ' 1 "b or the columns do 
not extend to the abaxial urla< (Fig. 2) f"he central bullilorm cell can be cir- 
cular to fan-shaped (Fig. 14) or narrower than deep, shield-shaped (Figs. 1, 11, 
16). Outer tangential epidermal cell wall are un ln« k ned to slightly thick- 
ened, with cells of si mi lai iz< Macrohairs have i unkery nonconstriered bas< 
and are embedded bi twe< n bulliform and/or colorless cells. 

The phloem of the primal s vast ul u bundles can be homogeneous or 
mi ,( li ii f (Ul n M M Ih.i inn iiupndvnli d u m liMiia or sclerosed (Figs. 
5, 10, 14) where it ad |( hi i tin iiumohh ,h.aih |\\ o enl need metawlcm \cssels 
are present adjacent to the phloem and one or two other enlarged protoxylem 
vessels are located adaxial I \ uuh< phlot tn U w ' 1 10 1 D Mi taw U m \ ( t 1 
ate mill not w id i th n the pamn< In m t In ,nh nil li htly thickened, and 
circular in outline K mestome heath ui rounds the xylem and phloem. The 
mestome cells are small with unilon ly thicl ned \ d 1 in ' b ndli h 2 
5, 10, 13, 14). 

Bundle heath intra prim ry \ cnlai aindk ometiiues include e.\ 
tension Fig. 5)andareentir< forma mplei circle) to adaxially interrupted, 
oradaxiallyandaba\iall\ inlet nipu d \\ igs I I L5)by a broad girder of a few to 
mati\ ,< In em n \ ma lib. i 1 1 ay 1 I Finn >knk s nit lar< de elk (Tig 1 I ) -hi 
ondary and tertiary vascular bundle parench ma lit u lis are mostly entire, 
not interrupted, toabaxially interrupted in ;ornc ;pecie: khe median vascular 
bundle parent h\ ma sheath i: most I \ abaxi lb interrupted, to interrupted on 
both sides, or less frequenth not interrupted t oinmonb 6-21 cells form the 
parenchyma sheath ol pinmun \ i t mai bundl il w ' > 10-12, 15, 16), with 
up to 24 cells found in som< ;peci< (Fig. 14 while 3-14 cells commonly com- 
pim p it m lo in i bundk ixath ol e< ondai \ (i igs > 10) and tertiary (Figs 
1, 11, 16) vascular bundles. 

\daxial aid ibivial ,t leu v li\ ma development is extremely variable! rom 
a few fibers (Fig. 1) to 1-3 layers or strands (Figs. 5, 10, 14). In a few species a 
continuous layer of sclerenchyma can form beneath the epidermis on the 
adaxial or more com i noii i\ ibixul mk i I io ) } (, 10) Sclerenchyma is 
usually present alom the margin o! thi i d. forming i 'cap" that ma) be 
rounded or pointed (Figs 3 4) flu clei leh i u > adjoins normal meso- 
phyll cells. Sclerenchyma is usually absent U h ecu ,k1u ,cular bundle where 
there are no continuous sclerenchyma layers. However, a few species, M. 
pauciflora (Fig. 2) and M. seatonii, form i m n ixiai irder ol" ini istal 
sclerenchyma. An abaxial pi o|. < ion a i ik i d In < leicnchymaissome- 

times enlarged and bulbous (Figs. 7, 8). 

Cladistics. — For the overall analysis ol 148 species (pi li two infra pcciiie 
taxa) of Muhlenbcrgia and four outgroup species representing four genera 
(Eragrostis acutiflora. Erioncuron avenaceum, Lcplochloa virgata, and 



I , , r , lit n n i , , , i 1 urn, in, ..ii K ii h i I 1 til. ,.iii1mii u o i 

All possible combinations were obtained by changing the order of each 
outgroup listed in the data set anil sequentially eliminating one, two or three of 
the outgroups. These 200 lives I torn the single overall anal\ sis are 97 to 99 steps 
long, with a consistency index (CI) ol 0.30 and a retention index (Rl) of 0.89. 
Twenty of these 200 rives were only L )7 steps long and therefore one of these 
was randomly selected lor illustration (lag, 17,). As indicated in the methods, 
[here were no synapomorphies supporting monophylv ol the Muhlenbergia 
clade, therefore the outgroup species were constrained. These 16 anatomical 
characters are not, b\ themselves robusi enough to test lor monophyly within 
the genera o\ Muhlenh. rgiinac. hragrost ideae. or the entire Chloridoideae. 
I In m l hull ivsoh in tin tncl consensu r. loi lln < t ill .w^ id i 


(using the 200 trees). However, the strict consensus tree separates a clade (Fig. 
17A) containing M. aurca, AT brcviliyula, M. distant, M. distichophylla, M. 

emersCy/.AI lydidcd, M. ruuadrv, AT omapudi VI ai\,^'nUi M n uhfolia M 
lehmanniana M lindhcimcri \i lonyj lumh \i Cms/byaC \i pilosd \1 
pu/vsccn.y Al. recckTm urn. A I. robust a, M.>copa> at, M spa M.xanthodas 
(bootstrap value of 100%); a clade (when rooted with Erioneuron avenaceum in 
a separate analysis at 0..W CI and 0,88 81 j or a grade (B), when rooted with 
Erayrostis acutijlora in a separate analysis) containing the following 37 species: 
M. alamosac, AT cuubTiu. Af appicssa, M. M. bi\'ndcyci, M. bushii, M. 
californica, M. ciliata, M. curtifolia, AT x curtisetosa, M. curviaristata, M. 
diversiglunus M dunn ■ \\ am; ( . /,..., \! >..,,« \l.glomerataM-hakoensis, 
M. himahiycnsis, M. hueycln., M japomca, M. mexicana var. mexicana, M. 
mcxicami var. jiiijormis. M. micmspci irui, M puiuifliTii. AT pectinata, M. 
polyandis. AT ported. M racemoM. M. M schreber'r M. setarioides,M. 
sobolifera, M. spiaformiy M. sylvaticcy M. tenellci M. tenuiflora, M. tenuijolia, 
and M. thurbcri (bootstrap value o\' 10080: and a grade ol all other specie- m 
the genus. The formei group ol 'I spe( ic: all appeal to be members of M. subg. 
Trichochloa, sect bpi.m)i/\ i mdi i rinm i Ha I', n i on 'n HU fwoapomorphies 
support a clade (big. clack \ J: complex compound b« I nth hi i i 

additional primary, secondary and /or ternary vascular hnndleslcharactcr It Oj 
and median vascular bundk that u lib rentiated from oth i primary vascular 
bundles fcharacter I [(?.){. I low-ever, complex compound keels [character 4(3)] 
appear in five additional species in two clades. One a\' these clades. containing 
M curviai istata VI. hakocnus \i himaiayi a h an \1. jd'po i, a i compos* 
onlv of species endemic to southeast -Asia. 

We must point our that Ai. nicxtuoia var. mexicana and A/, mexicana var. 
/i/i/m jm.saxaaar in rwoscixiratecladcst I le 17 -a tdeA).Both ipecies have identical 
scores for the data set. 1 lowevcr. abaxial sclerenchyma development (charact 
1 1) is am biguoLisI y scoret i as hav i ng one or more layers ( si ate 2) or th ree or mo 
layers (state 3) for each ol these raxa, 1 am not completely familiar with the c 
gorithm used in WinChuL'OOO but it appears that it selects cither state 2 or 3 
when reading ambiguous character scores. That would account for the inclu- 
sion a\ AT mcxiLdiid var. //A i/nna/.s after the node supported by churactci I y 8. 
whereas M me\uunu\ a in \u maw i <1 n 1 i I n ' ] ha tsjust one of the 
20 shortest trees and most of the other trees include both varieties of M. 
mexicana in the same clade. 

I lira tpoira rph i ] | 8 id an row i m r than 1 2 the blade 

thickness [charact ei 1 u| \ isculai bundle pa uioncd in t wo or three levels 
A h u u U i " ' ' uul ml 1 in d a II lo< i 8 (daw > lal > d < pi nnai \ \ i ( u- 

1 u bundk charact r 1( I i| uppoi i coi m no a T | w u I i 17, clade (J 
that correspond to members ol AT subg. Tr'n hochloa, sect. Podosemum (M. 

muytoMM. M duhia. \l pcpsopliiht, Al lalisnina, Al. lnridu. Ai m.n idiird, ,\l. 
nuu:rofu/td,M nana M palnuai Vi w. us VI r^it/i/ and \J s u/wristata). These 
12 species plus M. arlnuhita, M capillar is. and A/ sir it iu form a clade charac- 
terized by having vascular bundles composed ol primarv. secondary, and ter- 
tiary types [character 6(3)]. 

\UthUailhiyjti (iipiUin "is. ,\f. evpiiiiMi, and M. jilipcs are problematic since 
in other trees these species com prise a single clade or in separate clades. In 70 o\ 
the 200 trees these t hree species form a clade with the 14 previously discussed 
specicsteimii ivek placed in Ai.subg. Irn Mur/i Tut, sect. Podoscmum. In all other 
trees they are aligned with species of M. subg. Trichochloa in a grade containing 
clades of each section iT.pauaipc.s and l\hloscmuin). Therefore, placement in 
either section o\ AT subg. irn Inn hloa is premature. Interestingly, these three 
species were treated by Morden and ! latch ( 1080) as a single species with three 
varieties. Although rhev can be somewhat difficult to distinguish using gross 
morphological features there appears lo be sufficient differences in habitat, 
flowering time. and anatomical structure to warrant recognition at the species 
level. We believe (Ik etfirei .pea Icarlv belong in Al ubg // u hochloa since 
they form a clade with other mem hers ol sect, dpi rum pes and sect 10,/n.symiim b\ 
possessing a crown of inflated cells just below (abaxial to the primary vascular 
bundles [character 8(2)]. 

SoderstronH No'/ J delineated A I ub; /; t hochioa(;\ \i if- I'Oilosciutim) 
based on possession ol sclerosed phloem, caespitose perennial habit with erect, 
usually stout and robust culms, and glumes vemless or 1-veined. The single 
apomorphy(symplesiomorphyT of sclerosed phloem icharacter 1 5(2)1 appears to 
bean important character aligning at least II additional species in our analysis: 
M. clongata, AT jaimc-hinlonii, AT moaliina, AT miilica, M pubigluma, M. 
pungent M. purpusii. AT rcvciclioiiio M. sctijolia. M. versicolor, M. virlettu. All 
these species except A/ punpcus have the morphological characteristics that 
Soclerstrom described for Al subg. h a Inn hloa. Mnhlcnhcicai panccas is tin 
zomatous, decumbent near the base, and short (culms 20-70 cm tafl). Based on 
our morphological observations AT pungciis appears rehired to M. arcnacea, M. 
arcnicola, and M. fonryi. The last two species are the only other taxa in out- 
study that have sclerosed phloem [character 1 3(2)f however, these two species 
with M. sctijolia always form a separate clade. There are no obvious morpho- 
logical characteristics that align. Al. sctijolia with either M. arcnicola or AT 
torrcyi. Therefore, the evolution ol sclerosed phloem within Malilciihcryia ap 
pears to have occurred twice. Oven though M moniaaia exhibits some individuals 
with sclerosed phloem, others lack thischaracter state. Muhlcahcryia montana 
was aligned with M. straminca and M. vircsccns in about half of the 200 trees 
and therefore should not be included in A f. subg. ii a T>, hloa at this time. These 
three densely caespitose species all have 5- wined upper gl nines that are usually 

3-toothed as well. It seems best to tentatively place I hose eight species (excluding 
M. montana and M. pungens) in M sit by. /r?e/iei,. /j /<></ without further afTinii aw 

These 200 trees in the ovci all mab i ippear to support a group (Fig. 17, 
grade B) of 37 species with apomorphiesol loosely ai -ranged chlorenchyma[C4 
PCK type; character 7l2j] and Ian toslucld-shaped hullihi n cells without for- 
mation of a sclercnchyma girder from t h< icla iahoth 1 1 )axial surface [char- 
acter 9(2)]. These 37 peci< con pondtoM ul> WhM. mC gin \ll ohlu 
species except M. ursenci and M. phvrmdi.s have an additional apomorphy of 
four or more secondary and/or ten larv vascular bundles between consecutive 
primary vascular bund I e [Character 1 Oil )}. Howcwr. lour species (M. curtifolia, 
M.glauca, M pain ijl i and \i in hi hi i f i i i i.u t, i 10 

,A homofilaaousM m m tliest *., ,p ( i i< i t h» mi m u m i of only primary and 
tertiary vascular bund h lehai ctcrO Olalso haredwith nnualorshort-lived 
perennial* (Fig. 17, clades D & lis species (M. an nun. M. brcvis, M. breviseta*, M. 
lapillipcs. M. ! nspiscld. M. de/>o ii,mo aw Ai. c/nhmv Af. jilijcn ni/'s A?, /hv;,h 
h. /rtigibs. M. na/hiedfu. ,\/. hwhnh Ai. ma/hrcr/os. Al. iiniminMrio. Ah. 
peruviana, M. ram u losa, M. sch mitz i i, M. soi uosm M. s£ nC mr, M. tenuissima, M. 
texana,M. vaginata > llouoo I'O.-Mh i^Uim m < i ( dm i I ( m 
i.e., derived from a si ngle com in on ancesl or. lor the deri varion of this State [char- 
acter 6(2)]. 

The evolution of C\ photos\ m hesis in grasses is a complicated subject. 
ho\u v< r. itse< in o P ai i hat the | li i ha »ngmated at ! ilouitnm ml 
and Kellogg 1996) or more (seven or more times in Brown 1977). One of those 
origins appears to h Ik ubia i! < ilori > i c h 1 u l nvoize and Clayton 

1992). Our study suggc I th tm \lul nlm m In Pi \\ uh\ oc ol photos) n- 
thesis was a single evolutionary event [character 7(2)]. Since the occurrence of 
the PCK subtype isfoundin thu i ol th, o a, uoup |ho Cw ml/ men 
flora, Leptochloa virgata. and S/vwCh/o mrouhsj. it is not surprising that in 
all 200 trees this stati tppea ph lomorpfi lien rooted with t lu i t is 
HattersleyandWat MiiohXC j h\ pudi iml h iih l'< U nbi\|.ii ol mIIohi 
NAD-ME since in thcC4 acid cycle I'C'.K subtype isaneiiliaiicementof the NAD- 
ME subtype, and PCK is only known in grasses and may therefore have evolved 
subsequent to the N A I > M F. t y pc w 1 1 1 c h i s k 1 1 ( nv 1 1 i 1 1 ol lie to n onocotyledons and 
dicotyledons Jacob tlh i I i hat rh< l< 1 subtype is perhaps 

primitive since it is found in othei groups, i.e.. Panicoideae, whereas the NAD- 
ME subtype is restricted to Cfdoridoideae. We agree with Hattersley and 
Watson^ assessment and preier to vii v th levelopment oh Tic P' I nb p i 
Muhlenberg] a as tin derived tan \n alternativi h > jtoi Pn hi u I tin 

woufd require additional morphological or molecular evidence, might be that 
the PCK species or M subg. Muhlcnhcrgia actual I y represent :a separate lineage 
and deserves generic status. 

The remaining o4 taxa [M. apuasudicntcnsis. Al. annua. M arcnacca. Al 
arcnicola, M. avgentea. M anzonua. M. aspcrijolia. Al. btc\ijolia, AT brcvis, M. 
brevisettp M.aipiUipcs, M.uixamaiicnsis, M.ciccju. M trispiscla, hd.cualcnsis, 
Al curvula. Al cuspidaui. Al depaupcrata. Al duranycnsis. M. cludcns, M. 
jlaviseta,M-flcxuosa. Mjiapilis. Al hintonii. Al implicate;, M/<meszi,M. ligularis, 
M. majahcnsis. M. muhiscnsis, AT minutissima. Al. monnnu/. M. orophila, Al 
palmircnsis, M. pcnivianu. A/. /Wh/h /n-cf. A/, /'unpens, Al. pnrpusii, M quadri- 
dcntatci Al ramulosa, M. repens, Al. > ir/ian/sums. Al. si limhrui, AT stYJtonii, M. 
sinuosa, M. sinuosa, M. straminca, M. sinclior, M. tenuissi?na, M. texana, M. 
torreyanaM- torrcyi, AT utilis. M vagmuta. M. villiflora var. vi!!i/Iord, M. vil/i- 
/[ora var., M. vires. ens, M iva/smiiana. and M. urig/ilii) seem to contain 
sympleisiomorphies, i.e., they lack anatomical synapomoi pines I hrsyspn. ics 
all exhibit radiate, compact chlorenehyma or the classical NAD-MR subtype 
characteristic ol many chloridoid grasses Thai aciei n.\ )| contain primary vas- 
cular bundles without sclerosed phloem (character I 3(2)1, although ]-)resent m 
M.arenicokp M ptmgrns.and M lot rcyi: have rounded vascular bundles [char- 
acter 2(1)1, although A I. toi in \ ana. and M ju myc a > 'have ohovate/eUiptic or rect- 
angular bundles; have simple keels [character 4(1 .»[, although AT torrcyana has 
a complex compound keel like species m Al. subg. 1 ir/id iilaa sect. Hpuampcs; 
and have circular or irrcgularto Ian shaped bulhlorm eel k -[character 9(1)]. Fven 
though the clad istic analysis using these lo anatomical characters does not sug- 
gest a monophyletK lineage lor these 3 species, we prefer to recognize them 
inlonnallv as the 'Uomcna' complex (Peterson 2000). 

Within "Clomena' there exists some resolution, for instance, a ciade con- 
taining annual or short lived perennial species dag. 17. chide IT is based on the 
occurrence of primary and secondary vascular bundles (character 6(2)]. Other 
annuals (M. annua. Al. brcvis. Al. enspisefa. Al. depaupcrata. Al cludcns. AT 
/ragilis, Al minutissima. A I. sin mesa, and Al tcxana) occur as a grade. A grade 
(Fig. 17, grade FJ within T'.lomena' containing A J. apuasu.dicntcnsis. Alcua/e/isis, 
M. curvula, M. durangensis, M. flabcllata. Al hintonii. Al quadridentata, Al 
slraminca, and M vi rcsic ns. along \\ it h the remaining sprues in the analysis is 
alsodepictcd.Thcse species along with A I. urge n tea. M ( n.s/wsrfa, M. criophylla, 
M.filiculmis, M.jlavisela. M.pnesii. AT nuehisensis. AT montanaM-peruviana, 
and M. vvcflsoniufx/ haw been rclcnvd to as the Muhlcnbciyui montana complex 
(Herrera-Arneta 1998; Merrera-Arncta and De la Cerda-I.emus 1995; Herrera- 
Arrieta and Grant 1 993,1994). This complex consists ol high I y caespitose species 
usually with a three waned upper glumes, and in our analysis, leaf blades 
mostly with three or more layers of adaxial sclerenehyma in the primary 
bundles [character 140)|. The Muhlcnbcrgia re pens complex (Morden 1985, 
1995; Morden and 1 larch 1987, 1996) which includes Al /nsnginin, AT plumbca. 


M. repens, M. richanhoms. M. utilis and M. villijlora) is nor monophyletic, i.e., 
these species are found in two or more clades or as a grade with many addi- 

One of the two least homoplasious characters in the analysis is chloren- 
chyma arrangement (character 7, consistency index = 0.50). All species in M. 
subg. Muhlenbergia appear to be IX K leharacter 7(2)] whereas all other members 
of the genus are NAD-ME [7(1 )]. Only two species are ambiguously scored for 
character seven, M.glauca, more t han I i kely a mem hero! M subg. Muhlenbergia, 
and M. capillaris, clearly a member of ,\f. subg. 7nY/i<n iiloa. Median vascular 
bundle structure (character 11, CI = 0.=)0) is the other least homoplasious char- 
acter. All members ol \I subg /nV/iorMousect Epicatnpes have median vascular 
bundles that are differentiated from other primary vascular bundles [11(2)1 If 
you choose to disregard the outgroup species in the cladistic analysis, then 
within Muhlenbergia there is no homoplasv (CI = 1.00) for these two characters 

In conclusion, our data support the division of Xhihlcnhcrgia into two sub- 
genera (M. subg. Muhlenbergia and Trichochloa) and two sections (M. sect. 
Hpieampcs-cincl Podosemum)m M.subg. Trichochloa. Preliminary investigations 
of Muhlenbergia and relatives based :^o internal transcribed spacer region se- 
quences of nuclear til )osomal h)[\'A provide support lor containing on I v 
PCK species (= M. subg. Muhlenbergia) and another clade containing only M. 
subg. Trichochloa (Peterson, Columbus. Cerro Tlatilpa, and Kinney 2001). We 
prefer to view this partial classification based on anatomical characters as a 
work in progress and realize that with additional morphological and molecular 
data our understand i ng of t he evol i it ion of t his gen us will i m prove. We feef it is 
important to present this anatomical information since it is the first time the 
entire genus has been surveyed in 1 his manner therefore this serves as a foun- 
dation for further taxonomic research. 

Specimens used in this study, all housed ai I he Imiicd Stales National Herbarium 
(US) unless otherwise indicated. Those marked with an asterisk * appear in the 
figures. Collectors are abbreviated as follows: A - (. !.R. A unable: AC = S. Acevado; 
B = S.M. Braxton; C = A. Cortes O.; CA = MA. Carranza; CV = A. Campos- 
Villanueva; D = C.H. Dietrich; DC = M. De la Cerda-Lemus: DU = WC. Dunn; G = 
M.S. Gonzalez-Elizondo; H = Y. Herrera-Arrieta; J = E.J. Judziewicz; K = M.B. 
Knowles;KI = R.M.King;L=J.l.inl<ins;LB = RJ.LeBlond;M = O.Morrone;P = P.M. 
Peterson; PO = M.H.Poston:R= N. Relulio-Rodngucz; S = R : ].Soreng;V=J. Valdes- 
Reyna; VI =JA. Villarreal; W = A.S.Weakley. 

Hentensis Y.Herrera^ 

& H&DC1185 

MEXICO. Aguascalientcs: San Jose ( 

MEXICO.Chihuahua: 76 mi W La Ju 


P„ 7 

MEXICO.Chihuahua: Parque Nature 

[■ i 5 -. i:l'J I 

P&A 4982 

- ' .lit .i i. an Benito Co. 9.8 r 

RA&P0 8817 

ECUADOR. Provmcia de Chimbora; 

/:->-.■'. i S/.'dlle" 

P&A 4036, 4053 
P&A 4 102 

MEXICO Chihuahua: NW of Hernar 
MEXICO. Durango:Navios 


Holmgren& Holmgren 70 

j (Buckley) Hitchc. 

P&A 5703 

1 " "-M li hi Tl ill lit 

PAW 10(133 

MEXICO. Coa-.uila: 29.2 mi S Saltilk: 


P&A 5521 

>., 10 mi S f 

1 ■ ■ 

MEXICO Coahuila: 29.2 mi SSaltillc 

RA&H 8044 

MEXICO.Chihuahua: 15.3 mi SMex 

: IT , ^ 


M.EXICO.Baia California: Sierra San 

, , rih> 



:T Road, 1.2 mi E Dragoon 
Rucker Canyon on Tex Can\ 

:?-■ E F..urn 

A.S.Muller 1853 


RuXF 1 XoOO 

L i'. ; I r- • r 

RVVoSF 14236 


F'-oorl MEXICO. 


MEXICO. Baja Calif >rnia ir:lsla Catahna 
MEXICO. Baja California Sur: Isla La Partida 
Ml ■ < Chihuahua: 12.6 mi NE of La Bufa and 
GUATEMALA. Guatemala city 
USA. New Mexico: Grant Co., NE of San Lorenzo 

iE -I Lun mil t Hernandez Javales 

ME- ^',~raciuz: Orizaba 

r mil n mi E ( i i I I iimi, i i 

Mt MMnnj- Gni_ofLharcosonroadl 


I M t v i| ,n ,1 1 F'i Cajamarca,18 l< 

up load to mmh n i Sanchez Vega 
MEXICO.Chiapas i, i E 1 ,n n t LaM 

MK^CG. Chihuahua: I 2.1 mi N[ o' H Vorael o 

F I I I 1,1 i 1 ir NE of El Verge! o 

5 Imi Wof Navios,42miV 

zona:CocoinoCo., Oak 


P&A 5886 


SE Saltillo and 8 


/\m< it • 




SE Saltillo and 8 



E- ' i -> 

MEXICO. Durango 

nri:. , - r 

MEXICO. Durango 

mS of Durango o 

La Flor 

MEXICO. Chihuahi 


MEXICO. Durango 

mW of Durango 


mi E of La 


.VEX ICO. Durango 

miWof El Salto.S 

1.2 m 

MEXICO. Durango 







MEXICO. SinaloaD 








Ionia Cumbresd 

lea, appro 


-4 45, N 

MEXICO. Chihuah 

a: 2 

.6 mi S Creel on r 




SE San Antonio de las 

nas & SI. ( 

end of roa 



MEXICO. NuevoLe 

i i 




MEXICO. Durango 

S Charcos on roa 


MEXICO. Durango 




Santa Rita Mts. Be 


/on, 7 mi 


n Forest 

i:( "oemse Gxalono : -ii icier Creek 

'E lee Dut I I,, . 

MEXICO. Durango:: 
MEXICO. Chihuahu. 

Boyce & Godfrey 1 


: Co., 

MEXICO. Chihuahua: 35 mi V 

MEXICO. Chibun i lnr 

mi S Basigochi 

MEXICO. Durango: Arroyo El 

MEXICO Michoacan:CerroL 


A. North Care 

ZEl.TiM Pi 



Valley, along Hwy 67 

;CAColorar!u:'',j l iujrhi.'Co.,NWnl Xiouoi ho, 14 mi oof ■>■.. I o ■ 1 1 > f >o pa c e o; 
F CntMHiHv.-ylH 

rizona: Apache Co., E of McEMry 

XoEjOOE k\ ill 


H&AC 982 (CIIDIR) 

ME/ 10 ,ira 

E_E t [ 1- = : ..n tl - i 

a Reeder & C. Reed< 


Ear I 3-4 1 

i"E> ■_'_■ ■ iliuitiud 13.5 miWofParral 
MEXICO. Sinaloa: 34.5 mi NE of Mocontc 
MEXICO. Nayarit: 8 mi E of Compostela c 

MEXICO. Sinaloa: 3 mi SW Estado de Durar 
,6 mi N ofWasi 
MEX!CO.Chihuahua:76 mi W La Junta a 
Cascad Basaseachic 
MEXICO. Coahuila: 1 7 mi SE Saltilio and 7 

I : rr : 
ne at Bosque d 

MEXI >. 
V1E> 7 ).| 

; mi SW Estado de Durango a 

T.Tateoka s.n. 

JAPAN. Hakone:Kanagwa-ken 

S&P 5666 

CHINA. Xizhang (Tibet) Prov,Markam Co.Ningjing Shan,Markham Pane 

between Mekong SYantze 


O'NAb <:h,Ac.\< OovDvYenchman Cn.VV" --n j f _- of fieri rancie NW ot Ochuan 

5&P 5344 

CHINA S'cnua,n. Oiungla 1 Slamca 40 km WVVo/iou am; ca. 120 km NWC on 


MEXICO. Mexico: crucero Temazcaltepec 


MEXICO. Chihuahua: 0.7 mi W of Nuevo Majalca, 8.5 mi W of Hwy 45 

MEXICO. Durango 2 ) Hi - 1 ui m t i i 1 to.vards La Flor 

P&A 4095 

M.PX CO. Durango: 2.1 mi W of Km Chico crossing, 21 mi VV of Diming 

VENEZUELA. Trujillo:Quebrada de Dun 


T |. ■ f , |, m l"i„itiHliin t ,0 Imni of Laguna de San. 

1 T 1 ii u i H ) )nTex473 


MEXICO. Jalisco: 50 mi W of Ameca on road to Mascota 

V&C 2560 

MEXICO. Nuevo Leon: La Joya, Cuesta Blanca, 1 5 km S of Arambern 

P&A 6137 

MEXICO. Jalisco: 50 mi W of Ameca on road to Mascota 

P&A 6149 

MEXICO. Jalisco: Pass above Talpa de Allende,3.6 mi W of Rio Mascota 

S&P 5240 

CHINA. Yunnan Prov.: Fugong (N 1/2 Bijiang) Co. W slopes of Bilou Mts. 

S&P 5301* 

CHH I- 1 i in nmshdii d 1 H n, F ut 1 >ongchuan,ca 120 km NNEof K 

P&A 4857 

1 ' i i in, i f n ml F let Q 5 mi SE Hwy 

< O ^2' 

PANAMA. Province of Chirigui: between Rio Quebrado 

P,A&P0 8884 

ECUADOR Pio ii h< mu - , LA Faz on the Panamerican Hiv 

km N Ona 

r--.. i 

MEXICO. Coahui a: 260 mi Se SaltHo and 2.7 mi SE Jame on road to Sieti 

F i hi mif. ' imtl, , , h It 1( i L. Volcanes 

P&A 5408 

mi mi, into ruz Zo Patanonia Ml ilong , >ad t H d Mt - n 


RA&DU 30£ 

P*A 3990 
P&A 4048 
P&A 4097 

USA. New Mexico- Grant 

Co no e :ng Ho.o lornWof Mul- n • 



MEXICO. Chihua 


MEXICO. Chihuahua: 10. 

MEXICO. Oaxaca: 11.4m 

i W San Juan Mixtepec and 1 .5 mi E San Isidi 

MEXICO. Michoacan: 8.4 

mi : - E Zacapu oi Me> 15 towards Quii ; 

MEXICO. Chihuahua: Eo 

f Cumbres de Majalca 

USA. Idaho: Lemhi Co., 2 

.5 on E of Salmon River on Vva-m Son-gs Co 

CANADA. Ontario: Lake" 

7oh i mi ii i I'ii : n 

MEXICO. Durango: San J 


ECUADOR. Provinciade 

Az U ay:10.2kmNOnaonthePanamencanF 

ECUADOR. Provincia de '■ 

ochmcha: 1 ? h- N Calce^on on the D anamcr 

MEXICO. Baja California 

Sur: 25 km S of La Paz, W side of Isla .a Partis 

MEXICO.Sonora:18.2 m 

i E of LosTanques on road to Milpillas 

USA. Arizona: Santa Cru; 

: Co., Sycamore Canyon 


Co.,W of Flagstaff 

m SvV of Madera off h 1 towards Cuair 

MEXICO. Durango: 3.2 m 

W 7 loo Chico . toosing, 97.1 mi W of iJumiu 


MEXICO. Chihuahua: 25. 

MEXICO. Chihuahua:33. 


MEXICO. Oaxaca: 4.8 mi 

N7/TlaviacoonroadtoSan Jn.^n .V : >tcpec 

i Laegaard RA&PO I 

MEXICO.TIaxcala: 5.2 mi N Tlaxco on Me 



- -t: M mta Cruz Co. Patagonia IV 

ts„ 12.3 mi S Patagon 

MEXICO. Chuhuahua:40 mi W of Balleza 

and 19miEofGuach 

,E>r i mi.ruz Orizaba 

MEXICO. Mexico: 28.3 mi NETemascalte 

MEXICO. Mexico: 1 5.6 mi E Amecameca 

nd 2 mi N Paso deC 

MEXICO.Tamaulipas:63 mi SW Cd.Victor 

MEXICO. San Luis Potosi:45 mi NE San Luis 

USA. Arizona: Cochise Co., 3.1 mi above 

pper Picnic Area, For 

i in Mi i inta 9n Its.! Canyon, 7 mi H 

rCUAD03. ihovincia cic COmbora::o:S.9 km N Palmira on the Panameiicar Hwy 

a Hwy 62 (180) 
ECUADOR. Provincia Cotopaxi: Lago Limpiopunga 
MEXICO. Durango: 1 4 mi SW of El Vergel on Hwy 24 
MEXICO. Durango: 7.0 mi W of El Salto on Hwy 40 
MEXICO. Mexico: 5 km NE of Teiupil . n , I it ,i i Mmascaltepec 35 mi F ilO d m,d 2- mi E Guachochi 

O: a mania 

f in On til 11, <irandes on Mex 2 

MEXICO. Durango: 4.5 mi N of Borbollones, N of Hwy 40 

iEXI lahuil , L t ii I l i u mi H 53i i I F i nlla d. I arm. > 

MEXICO NuevoLeut, mi I I ,i u, Oi I d del Potosi 

: <:':!'-:■: i". i d- : 

- : :of,!CIIDR.. 
_ -i ~ ■ '"MLjIP 
/.Tall C814 

RG&K 1 vo •; 

■Li ' 1-i- 

P&A 5422 

- -/', rnr 

■. i 

P&A 6273 
P&A 4056 



'" 9 ' d °' Kun,h,Tri 

RA&PO 8895 
RA&PO 8820 
P&K 13300 

PSo 1330 H 

E i iuhpas63miSWCd. 

' E- i I n mjo 40 km WDurai 
MEXICO. Mexico: 28.3 mi NE of Temascaltepec or 

i'.'E'h i'.i. .ico ParqueNacional NevadodeTol 
MEXICO. Oaxaca Ladera SW del Cerro Pelon, 500 
M.EXICO.Puebla:Ladera E del Popocatepetl, 10 N 


VE-lv" T'linn: ,', oi f,,i. . ■ 
VEXICO. Durargo:VVof El Saito 
MEXICO. Michoac- 
VE- r: hunngo 56kmWDuranc 

I! E 

S Charcc 

VEXiCr..-;.i.-^ioa:3 rri SW I Oooy r)e D.raro i .-md 03 rV 5 El Paimim on Hwy 40 
USA. Arizona: Santa Cruz Co. Patagonia f 't i r c '-o Service Road 1 35 8W 

USA.Texas: Bexar Co. 5 mi N of San Antonio, city limits on Hwy 281 
MEXICO. Chihuahua: 10 mi Wof Cuahutemoc on Hwy 16 

MEXICO. Chihuahua: 5.6 mi N Cuahutemoc on Mex 23 

' El i ih ii iid r r ii [ 1 I ii-nii' it , i I 1 i i i dda de Basaseachic 
MEXICO. Chihuahua: 52.5 i SE Villa Matamoros and 1 mi f I Ejido Pevolucion 
ECUADOR. Provmcia Ic^ih rin N na on the Panamerican Hwy 
ECUADOR. Proving I I i_ -~h -hi ni I r o- > - 

I h a- i E i I n i I n i 1 mi c Temosachic 

E i Nu Sml 4niiIH ~ nH it i ml t t 

' E Nu Leon: 1 3.4 mi E Hwy 57 on 1 1 58 at crossin I , t 

'/L-COSmM, . r i EOV Cd. Sictora on "- IS' t h I m L r t 

P&A 5455 

P&A 5434 



P&A 4631 

P&A 6079 

P 14231* 

P&A 9946 


pap 133^. 

P&A 5716 


APA 1 2590 

O<0 -152:1 

P&A 4054 

' lEXI ) Oaxaca 5 mi NE la del m ulaf in 31 d mi NETt tonqo 
MEXICO. Oaxaca: 5 mi SWTeposcolula and 2.5 mi NEYolomecatl on Mex 1 
USA. Arizona: Cochise Co., 2 mi SW Sunnyside along Forest Service Road 2 
\Arizoi banta Cruz Co. Patagonia Mts., 12.3 mi S Patagonia on Forest 

MEXICO. Mexico: Durango: 5 mi E of Mezquital on road to Charcos 
MEXICO. Michoacan:6.1 mi W of Ciudad Hidalgo on Hwy 15 
MEXICO. Nayarit: 29 mi SW of Tepic on Hwy 1 5 to Guadalajara 
USA. Maryland: Montgomery Co. Bethesda,4520 Cheltenham Dr. 
' I F H I i in __ ' . rdanonMex140 


V of Hwv 45 c 


M'- AAA. Dmaryo: "" m vP or Mezquital on road to Charcos 
MPXICO.Sinaloa:3 mi SW Estado de Durango and 2.2 mi S El Palmito on 
MEXICO. Coahuila: approx. 20 mi SE Saltillo on road to Los Lirios 
MEXICO. Nuevo Leon: 3.8 mi S Allende on Mex 85 towards Montemoi 
MEXICO. Oaxaca: 1 .4 mi E Ayutla on Mex 1 79 towards Mitla 
MEXICO. Puebla: 3.5 mi SE Cd.Serdan on Mex 1 40 
MEXICO. Coahuila:85.5 mi NW Muzquiz on Hwy 53 towards Boquilla del 
MEXICO. Iamaulipas:55 mi SW Cd. Victoria on Mex 101 towards SanLui 
MEXICO. Chihuahua. 

MEXICO Aamauhpa r i in A I i n i , I I n hi ■ 

MEXICO.Chihuahua:21.1 mi ' tH v 4 r i>4 mi E of Cumbres de Me 

MEXICO Chihuahua 3 1 m 1 r H, inand-^ Pi des,32 mi SW of Colon 

; __ I' Pi id i l r n ', )t La Junta on road toTomochic 

P&A 4059 
■ < Ann'- 
P&A 4092 
I -..-. | 3514 

VI HCO.ChTuarua:54Ai-ni N Farrai m '/._■/ 24 to Chi- aah ja 
t I till nti i n il L if i H JStowardsChih 

; 'i ' liihu itm 1 - mi E t f I . i ml on Hwy 24 

f Durangoon I 


E 1' ' I In, t,ii- 10miSWLaJunt c 

P&A 4545 

E' lnl at h uladeBasaseachic,; 

P&A 4045 

MEXICO. Chihuahua: 5.0 mi 5 of Hernandez Jav 

P&A 401 9, 4021 

— ii r i int, ru -miSWofCanelc 

P&A 561 8 

j m - 1 r f hpe House Overk 

P&A 56 19 

USA. New Mexico: Rio Arriba Co., on Hwy 84, at 

USA. New Jersey: Burlington Co., 0.1 mi N ol Atlar 

i L i IN- F in i i ii 4f Gn EofCachic 

ARGENTINA. Mend , ir ir t -it E i 

Hwy 40 & variant 40 

P&A 11418 

ARGENTINA. Mendoza: Depto. Lujan de Cuyo: a 
ARGENTINA. Saita; Depto. Chicoai 
Carril and Cachi 

- EMTI ' ilta, Depto. San C 

UiA.New iV'K'>ico: Grant Co., IE 
JSA. New Yoik: Oneida Co., ' i 



;. Ed> :> hhi.dh 

P&CA9709 MEXICO Oa.,1 , 1m„ dE . C 1 GTun izulapan and 5.5 mi 

P&CA9723 MEXICOOa i inilG I il ion Mex 125 

P&CA9724 MEXICO. Oaeev rE-'- ET-e ■ Nn!a on Mex 125 

P&CA 9729 MEXICO. Oaxaca: 5 mi SWTeposcolula and 2.5 mi NEYolorr 

Hernandezs.n.(HUAA) n F - 1 ■ » hi i lGi, m. dm Jose de Gracia, Sierra Fna 

P&A5592 USA. New M< -i ml m r I tit T Y h n iE- 

[M]',r !"1! 'I 

12|l 121 22 I 221 1 

12 II 2. '122 11 1232 

2 2]2 M.jdfX) nit.ii 



212112212[1 2]121322 

A; /,in<j//i'i /u'm j 


23][14]2111 1111111322 


i! ..,,,/;, ■! 

[24]2[23]1 1121 1222322 




?>]?]?> 2 






211 '123 " 



21241231 11 



113 22122 





M 1222322 





241231113 1 

1212 12222 

3J312H '1!M ' 1 

ill 121 1212222 M.tenella 
1211112|23]22 M.tenuifiora 
111111 I ' ' MV.n, 

243123171 11 12[23]22 

1212322 M.uniflora 

M.setarioides 1121122122111222 

We thank Stanley Van! n k i lot lvi< in preparing th n t n i il pivpara 
tions; Carol R. Annable and Marjorie B. Knowles for preparing a portion of the 
anatomical slides; Robert J. Soreng for assistance with CLADOS; J. Travis Co- 
lumbus and Robert I ■ vVeb lei loi -reviewing th in hip cnpt the Smithsonian 
Institution's Fellow hip md Gram i( prw dins i hort term Visitor Grant 
to YH, the Scholarly St in lies Grants Program; and the Natural Museum of Natu- 
ral 1 listorys. Research Opportunities Fund. 





unouveaux genres des 


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ntlescaracteresde tousles 

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id J.P.MlKVHI 

1976. Botanical m • < 

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versity Press. 



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Guy L. Nesom 

Boha )/,' hit ' .' f '7e> 

509 Pecan Street 

h-)H Worth. i\/6!0.> .lOotl! ISA 

InstudyofGnaphalieaeofMexicoa dadjacentl nited ltat< ;, various undescnbed 

taxa and t uiyec n unions h ise t om 10 li'du I li pi nr | > pu u pui i mi \ 
species whose geographic range is centered in near-coastal and adjacent inland 
localities in southeastern 'it i omc lo. I i t i « more than lOOairline milt I i >i 
the coast (Fig. 2). One collection of the new species was made from the out- 
skirts of the city oh Monterrey Mtcvo heme Mexico. 

Pseudognaphalium austrotexanum Nesom, sp. nov. (Fig. 1 ). Tvrir UNITED STATES. 

1 1 \ \ |un \\ list o 10.1 tin ol \lu i iilh.ul n In oi y v ill i i it itiudisPiM 
ion nl Kin Ranch, open I > i m 1 1 n I now iiisIi-l>ui\ el.nt \ cml lo.un ' I N>« 
1Q-34, MCJohmton vf2/iV ( l IOU vi vni-:: TEX': 1SOTYPE: SMU!). 

tentibusbicolei'ibusglandulaiibusqiu umli E.imIi i n .0 n Jul. . i I -1 i n i I i in 

aipitulisminiinbusilotibd-paik lniib i , i pli II mi ■ innin idj[Huin rot undalisapiculatisdiHen. 

Annual herbs from i woods tapi i a u m dm lall d< nsel> and closeh 

white-tomentose 1 hn < o ,. Mile n I ulnLu 1 1 n tl\ eiect, unbranched 
until m rthcinl'loi cenea >r rai I more highly branched and "a large bushy 
herb" (fide comments on Runyonl954). I eaves < onge ted on very short intern- 
odes, continuing congt ted limm liatcl \ lacn ihtlieh 1 
to strictly ascending, the lower (.moribund ) loosely spreading to deflexed, the 

blades cpetiolaie, linear to linear-lanceolate, 2 -5 can long, I -3 mm wide, rela- 
tively uniform in size, shorter immediately below the heads, subclasping but 

not b i ill\ nupli in i,n i< ul in >i a ui n ml iptcalb at tron d\ bi< >l 
ltd li hod i uti re< .1 n in (an gland ul nherwise glabrate, 

th< lower surlaei dense!) u id closed \ e bn romeniose I In mat in rc\a>luti 
sometimes closely sinuate. Oapitula 1. 5-5 nun high, in tight glomerules, the 
glomerules usual 1\ borne n i I'lai topped inflon eerie* 10-30 cm wide, the 
ultimate branches 1.5- 1 cm long; ph yl lanes narrow '1 \ ovate to oblong or ellip- 
tic, persistently wooly at the base, the inner with an elongate stereome, gland- 
dotted near the apex, the outer ca. 2/3 as long as the inner. Outer, pistillate flo- 
rets (in Mexico, 46-)76-l02; inner, bisexual florets (6-)8-ll. Cypselae oblong, 
brownish-yellow, 0.5-0.7 mm long, 4 ostnate-ndged longitudinally, minutely 
n ip II * i | i| | u ( t li oli >i i i 11 id i mis b ,i b« Han bi istb , 

Monterrey on the Momemoivlos highu ay gravel pit ot thet u. Fundidora de Fierro y Acero, 2000 ft, 

270ctl9b7 Fleetwood^' II \i .mil I i Ja n HKt ta4 (,-M llfoialti Bra:os Co.: 
College Station, 3 Oct 1940. 1 FX 7 Brooks Co.: sum N ol 1 allurnas .18 Oci 19/ W:"vmli mi. 
(SMU). Frio Co.: 13.6 mi NNW of Dilly on Hwy 1 17. scrub pasture, reddish soil, 31 Oct 1981. Mahler 
9225 (BRIT). Harris Co.: Seabrook. 18 Oct 10 R Falter 29028 iTF.Xj. jini Hogg Co.: 9 mi SW of 
Hebbronville, sandy loam not plentiful 10F\s FFU Ifom vtia'l I \) Kenedy Co n. n I udolnh oi 
Nonas, sandy knoll along highway. 3 |an 190 5 ( ,>rreU 2^'->2h (\C\ '. TFX;; 4.4 mi S of Armstrong. 
sandy sacahuiste prairie ai edge ol caliche t lat. 0> Nov N34. Johnston 742dO(HTEX); Sof Armstrong 
on side ol llw\ tmh lorn, sod. scan e, on ly.i lew plants seen. 17 Oct N 78 Funyon 1954 (TFX-2 
sheets. US); Nonas, highway right-of-way. dune -and. 4 I \s M 18. I harp.Johnson.and Webster 48- 108 
(TFX). Matagorda Co.: Matagorda, 14 Oct 1930. FalierdOOl (.ARIZ. TFX, US). San Patricio Co.: 8 mi 
SW of Taft, near shore of Nueces Bay but above high tide level, soil not saline. 10 Nov \956Jones 1261 
(SMU);ca.2miSEol e)dem,m sanciv open paauie local U abundant. 31 Oct 1959. Jones 3610 (TFX ). 

These plants are similar to fa/ay an/dad i ami viscosum (Kunth) A. Anderb. in 
general appearance and previously have bean identii ied as that species. Plants 
of both species are taproot cd annuals w 1 1 1 1 w 1 1 1 1 e foment ose, strictly erect stems 
mostly unbranched until th< ml la acnes leaves linear to linear-lanceolate, 
strongly bicolored (green and glandular above, w hite-tomentose beneath), 
loosely to strictly ascending, crowded on shorl internodes and continuing to 
uiiiia diaieb la low the head md b li til I pin but not tron I ut u i 
late, phyl laries si 1 very, t lun-h val t ne, and achenes minutely papillate. They are 
distinguished by the following contrasts: 

t i i - tni.liil a ii i, ' t it. glandular; leaves 

i 111 MUM ,11 III tit Ol 111) mi 



mi i h hi i| i il i hm) ) n i i i I h I ipiculate from a thic!- 

ened and slightK mi i Inn lnl I ^i i , 1 i i I I i il i 1 I 

not decurrent. Pseudognaphali 

Pseudognaphahumvisiosuiiw distribun i hrough Mexico nd Central America 
uid il o apparent ly is i omrnon on the Cai ib 1 .. an i land ol I lispamola In the 
United States, it is known only from sout h western Texas, widely separated by 
distance, habitat, and climate from P.austnHcxanum (lag. I) 

i I nt of I udo naphaliumau tivl< <auum\\ n inel ided by ( orrell uid 
Johnston (1970) as "rare in s.e. Tex:' with plants identified as Gnaphahum 
macounii Greene, a nam that ha ometiim b< n mi nnh* d to Pseudognapha- 
l in mvesrn.simi. The other Texas plants ol "( Kiuacounif ("local in Rio Grande Plains 
and Trans-Pecos ... and the Llano region ol the Edward Plateau") are Pseudo- 
gnaphalium viscosum rather than Pscudognaphalium(.Gnaphalium) macounii 
(Greene) Kartesz, wine 1 1 does not occur in texas The only known Mexican local- 
ity for P austrotexanum is separated by about } 00 kilometers Prom the closest 
Texas site, although the geology and topology arc general ly similar. The Nuevo i rnajoi high wen and i fruit may hae< been accident alb 
transported (vehicle-dispersal) from Texas. 

The distinctiveness o\' Pscudogiuiphalium austrotexanum also has been 
recognized by several other botanists: Marshall Johnston (by notation); Billie 
Turner (pers. commj; and 1 larvcy Ballard (by annotai ion and pers. comm.). 


I am grateful to the staffs at BRIT, GH, TEX, and US for help during visits, to 
Billie Turner for arranging for the fine illustration by kinny Heagy, and to an 
anonymous reviewer for helpful comments. 

Book Review 

Kay Yatskievyci i. 2002. Field Guide to Indiana Wild Flowers. (ISBN 0-253-21420- 

3, pbk.). Indiana University Press, 601 North Morton Street, Bloommgron, 
IN 47404-3797, U.S. A .(Orders: i upress.i nd! n, 800-842-o7Qo, 8 1 2-855- 
4203, 812-855-8507 lax). $17»5, 572 pp, 040 color photos, line drawings, 5 
1/2" x 8 1/2". 

icvisal during I In- past leu decades- 1 1 seems Ininr licit the importance o| held suidc ;;o.'s t;u he 
yond their immediate practu ,il use! illness ol idcutiU mg a spec ics o! plain thai might attract ones 
interest iorashorl moment 1 lere are some ol mv leelmgs aboui ihc Simula aiieeoC wild I lowers and 

plated that these springtimes have been rh\ahnnealK' undergoinj; their ever-ehan S in R complex phe- 
nomena lor at least 10.000 years in I he glaciated parrot Indiana. 1 personally know that the places 

and oak-hickory forests in some, it the 'Mate paths and Male loresisol Indiana are quickly vanishing. 
What tl an unexpected, terrible disaster an k as a loinado earl [uptake I ire. explosion, or even 
a terrorist act destroyed one ol the line buildings on the campus el Indiana University or Purdue 
University? By using die blueprints I were used to build the original sii :.u. mre, highly sophisti- 
cated, well-trained Indian,! arc luted-, engineer- and oilier si enttsis, , ml, I draw up plans to replace 
t he dest roved building exact I \ asii was even to the minutest detail sue k as t he determination ol the 

■vclopmetil destroyed n 

.1 architects, engineers 
nest detail ol the tm I 

lessly des 


into the d 


ded us in 374 pages at 

ings, colo: 

r photos ol examples 

species, h- 

er guide will help pb: 

much greater detail 1 lopclulls 

deeper int 

o the forests and uncc 


Guy L. Nesom 



i M ll ill uiiilml miii l, i pi K in \H« il.'lllli l C . I, his \ Ih-iiuh 1 i is 
iisjn u I i ihui llt» ii in . i. ii mi m ilnn iih i|m i omo Keysseria S( 

. sus ilosculos bisexuales y I en lies (vs. luncionalmei 
s dilerencias moriolngK i en las eorola eentrales \ perilen. 
demica de la lsla Vanua l.evn Fiji se exclme cle KevwTi.i v 
<<!■( i i. h riii i ii .in om ii. . ilrec. un i p i. 

n Kev.vseih! y I'vd ni. </i />.< se divideii en siele '. eiiirales- v t m 

Kt ]\soia\ auterb u n I ill rhacli 19I4J i en lis i 1 peu< divided be 

tween Indonesia 0' p v , i - p| u ml nv pe< il n i,r> i Im kt I'lmw a in ! 1 lav im ' 
species: Mill l°')(i l ( )Oi) , |),|,, U ) \1I ul l In Indmi. hi |k .x.iiiin 

Guinea; K.radicanslT Muell )M -.111' a] ;oi: knov n E"rom( 1 Lebes and K.gibbsiae 
(Merrill) Cabrera 1 \ Stceni also ex < urs on Ml I- inabalu in B01 nco 1 h. m 111.1 
was treated as a tm in bet >1 \s!crea< ubtribe I agcniiei 111a \ s 1 0111 ! ' ! 
2000) and a recent morphological and taxonormc overvnw is a\ ai 1 able (Nesom 

The three Hawaiian pc< e were transferred to RVys.s '< ml rom Ldgcnijcra 
Cass, by Cabrera (19po). Mill (1990, 1999) maintained diem in a broadly con- 
ceived / ((emu/i km 11 1 n 1 t h, \ , no v 1 lob 1 1 m 11 
SwensonandBremei "' 1 oexplicitly n irded K 1 1 1 vnonvmoi 
Lagcnifcra. Perspective on morphological distinctions among these genera is 
provided below in the key to the "core genera" o( 1 .ageniferinac. 

The Hawaiian plants o( Keysseria arc similar to the Indonesian ones m 
habu,u. I nihil ami morphological derail hut dill i a lol lows: disc florets with 
fertile ovaries (vsd line riona I lvsta mi i niie). ra\ eoroll, is usual I \ deeply and asym- 
metrically 2-4-lobed at the apex (vs. apieallv entire), and disc corollas some- 
times 5-lobed (vs. consistently 4-lobed). Mill (1990) noted that the Hawaiian 
species apparent!}' are "from one founder;" in this summary of supraspecific 
taxonomy o\^ Keysseria. they are treated as a separate section. 

Keysseria Lauterhaeh. Lepcri. Spec Nov. begin Yeg I CAT 1914. Myriactis Less. 

suhg HcidUntis L Much.. Trans. Royal See \ arena I, 2:11 1880. ilccatactis [V: 
Muell.H. Maell.ex MuttL, Bot.Jahrb.Syst. 62:407. L929. Keysseria sect. Hecatactis 

(.I: Muell.,) Maul.. Bet. lahrh. u8:2s0. L87 Tilt: Kevssei id gmiiaaa lauterhaeh ( = 

The nine Indonesian species constitute sect. Keysseria. 

s Nesom, sect. nov. Typk: Keysseria eiici 

Differt a Keysseria sect. Keysseria floribusdisciiovariisfertilibus,corollis radii plerumque ad apicem 

.'. 4 lohans.ctcorolli.discnaliquando-i lobatis 

Species included: K. erici, K. mavicnsis (II. Mann) Cabrera, and K. helenae (C.N. 
hordes Nr I ydgate J Cabrera. 

The "core genera" of Lageniferinae 

Lagenifera Cass, and genera similar to it have been grouped together as the 

Midi rihe I agcnilcrhiac I Table I J i Nesom l99Ta). 'I hree genera have been a.dded 
to the Lageniferinae since the recent elassil ication and ovci \\c\\'\Lagerioeypsela 
U. Swenson and K. Bremer (Swenson & Bremer 1994); Pytimcarpa Nesom 
(Nesom 1994b); and Pappochroma Raf. (Forbes & Morris 1996; Nesom 1994c, 
1994d, 1998; synonyms = Pagenopappas Nesom and / mvmdn i \ Nesom > The 
12 genera and approximately 71 species eA Lageniferinae are distributed from 
India to southeast Asia, Australia, and various Pacific islands, except for the 
nine species of Lagenifera and Mvnacd.s native to South America and Central 
America. Plants of the seven tore genera" of Lagenilei mac Liable 1) are prima- 
rily characterized (with exceptions) by a herbaceous habit, leaves mostly in a 
basal rosette, heads borne singly on sea pose or scapilorm si ems or few in a loose 
panicle, pistillate florets with reduced lamina and m several series, a tendency 
to produce functionally stammate disc florets, and flat, 2-nerved, epappose 
cypselae, commonly with a glandular apex or beak. The remaining five "pe- 
ripheral genera," while similar in geography mav not be closely related to the 
others; they differ in com binat ions o! \ anous features, particularly in subtercte, 
multinerved cypselae. 

A key to the "core genera" provides perspective for the positioning o\' 
Ke ria and its disi inction within the group. 

/ oaenocyv^chi '- ' Vwrisun & K. Brci 

I ii ' p< h \usti ■ 
Synonyms i niinioihippus N. 

• , » t ' i | > i- Austral 

"peripheral genera" 

Pytinicarpa Nesom (3 specie 

>s; New Caledonia and Fiji) 

Rhamphogyne S. Moore (1 S| 

Rhynchospermum Reinw.(l 

p ci I pin Km,, i ! ut Lhl vi 

r'nlldll- Ii III ' I, 

lamina of ray floret 

. i:i hvllaru'v.'llip1ii t >U >vale io oblong, I ilunllv founded Ioo.oIumo ipiiuLidoi 

. Cypselar beakrudim 

. subclasping but not basally dilated oi • , 1 1 .. jilrmi, 

ttl il ng unreduced 
ipletely up the stem. Myriactis 

Transfer of Kr\s^ ■ pick • <ji to Pytinicarpa 

Keysseria picker ingi'p originally close -pi bed In G ray (.18(31 4 is endemic to Vanua 
Levu Island, Fiji. Smith and Can (1001, p. 502 ) noted "it is not possible to sug- 
gest a New Guinean relative of KVvsserui fhlvi inyii. whieh is remarkable lor 
its very small heads and its eostate aehenes." 1 his species, which has a basal 
rosette of narrow leaves loosely villous si rigose on both surfaces, 
monoeephalousand minutely bracicate,scapilorm stems.small, radiate heads, 
convex receptacles, functionally staminatc disc florets, and subcylindric, 
eglandular cypselae 4 mm lone, \\ ith 8-10 longitudinal siiongly raised nerves 
and a truncate, epappose apex, is a member ol the recently described genus 
J'yi i in, ,n jui Nesoin (Nesom 1004 b), which originally included two species from 
New Caledonia. Some features of I'yiniiuirpa Oleography solitary capitula on 
scapose stems, functional!) staminatc disc florets epappose cypselae) suggest 
that it shares close ancestry with genera of l.agenilermae. but the conical-con- 
vex receptacles, ray florets in a single series, and multtnerved, subcylindric 
cypselae are unusual in that subii ibe. 

Pytinicarpa pickeriu^ii vA Cruv) Nesom, comb. nov. / a^cnophora pickcringii A. 
Gray, Proc. Amu Acad. Arts 0141. 18(0. Kcv.s.svnO pickcnn^ii (A. Gray) Cabrera, 
lOumca 14:307 Neo. Tyim:: 1 I ] 1 : Vanua Leva Island. Marhuaia Mis.Jca. Jul 18401. 
Wilkes Expl Exped. s.n. (hoiotyph: US!). 
Pytinicarpa pii lu ) in it dilli • i rom both New < dedoman peci mil cypsclui 
surface, which is in i mil el \ papillate, the cent ei ol each cell abruptly raised into 
a sharp point. In the New Caledonian species, these epidermal cells are simi- 
larly quadrate but the whole surface o\ each cell is convex and the cypselar 
surface has a "frot In" appearance. The relatively broad leaves of Ppickcnngii 
are more like those ol P. sarasinu (see below) rather than the linear leaves of P 
ncocalcdonia, but those oi P. picka ingii are obovate-spatulate, abruptly nar- 
rowed to a petiolar base, and the margins are shallow 1\ ' crenulate-mucronulate 
(vs. oblanceolatc, without a distinci petiolar portion, with margins coarsely 
toothed only near the apex). 

There is some indication that the narrow "'coronal rim" ol the cypselae in 
Pytinicarpa pickcringii might have been sticky when fresh, which is a general 
feature of the Lage inter mac. This run is similar to that illustrated for thecypsela 
of P.sarasinii (Nesoin 1 004b), where stickiness was not evident. 

Nomenclature of New Caledonian Pytinicarpa 

Lagenijera neocalcdonica S. Moore predates Brachytcomc sarasinu Damker, 

which was the basic in \ m lor one oi lie two original spec u in Pyi i/nourptKNi soin 
1994b). Cabrera (1966), in his list of species excluded from Lagenifera, recog- 
nized that these two names relet to the sunc species, But because the epithet 
"neoealedonica" al read v exists in l>\iinicarp,i lor the secoi id species, P.sarasinii 
is the correct designation of the broad -leaved New Caledonian species named 
by Moore and IXamket The two New C aledonian species are as follows. 

iinii (Camkei ) Nesom, Ehvtologia 76:138. 1994. Brachyscomc 
iker, Mitt. Bot. Mus. Univ. Zurich 142:479. N U \n NEW 

Lagenifera neoealedonica S. Moore. J. linn. Soc. Bot. 45: 34 x 1921 Inon Pytincarpa 
{Brachyscome) neocaletlonua (Guill.) Nesom BAH). Two NEW CALEDONIA: 
Taom, bare reel serpentine earth. 2i00 It, rare, 2 Dec 1914, R.H. Compton 2.305 IO- 
toni'i-: I5M. photocopy!). 

2. Pytinicarpa neoealedonica (Cm 1 1 ) N( om lOvtologia 70.1 >M E-NH On ' 

neocaletlonua Guilk, Bull. Soc. Bot Prance 84:61. 1937. syntypes' NEW 
CALEDONIA: Gatope. Yidlanl 2N23 c\A' GH!j: Nehoue. Puncher 425 and 
Deplane he 425. 

I am grateful lor help from the stalls at CI 1 and US during -event visits, the 
staff of the Couch Botanical Library 1 1 fN( -( hapel 1 [ill) in obtaining literature, 
lohnStrotherand Mark Wetter for commeruson the manuscript, and Roy Vick- 
ery of the BM staff in sending a photocopy of the type of Lagenifera 

I S.J. and [J.I. iV I " it 

1 ii - i ' N lw - I dfirl Ropeit Spec. Nov. Regni 

. In: Wagner, W.L, D.R. Herbst, and S.H. Sohmer. 
'i.Vol. 1 , pp. 329-331 . Univ. Hawaii Press, Honolulu. 
hytologia 76:1 93- 

Phytologia 76:136-142. 

Ni m!'.i, G.L. 1994c. KvxDiionm el tspt^rs.i I ol Austialian h 

Phytologia 76443-159. 
Ni mj | | )] I > I ilit tin i oirectgeneric 

La bill. Phytologia 76:426. 
Nesom, G.L. 1998 [2000]. Full constitution of the Austra 

(Asteraceae: Lageniferinae). Phytologia 85:276-279. 
N '.; w.G.L 2000.t".enerk v onspectus of the tribe Astereae 

Central America, tne Mitill" , and Ha aim Sin i,Bot Mi: 
Smith, A.C. and G.D. Carr. 1 991 . Asteraceae. In A.C. Smith. Fl. 
Si i ,U md K R i l ( < ' I I hi- I'-nu , ,< 'm m > , 

Guinea. Austr.Syst. Bot. 7:265-273. 





PNG I ores! Authority, POB 406 

Rabaul, East New Britain 614 


around the clmandnum. and by the differently shaped li 



The genus Saccoglossum is present lv comprised o\ lour taxa: S. lanceolatumL.O. 
Williams, S. metadata Schltr., S. /wpudnum Schltr., and 5. verrucosum L.O. Will- 
iam The laitc pedes is re<. 1 nl h mil m Java \ icreas the others are 
known primarily from Papua New Guinea (PNG). Chyme (1994) suggested 
there are about 5 species and that Mane new ones could he expected i tout the 
PNG Highlands. In the orchid treat merit s for Win uat u. t lie Solomon Islands, and 
Bougainville (Lewis & Cribb 1989, 1991) no records are provided for 
Saccoglossum. The new species extends the generic range to New Ireland and 
apparently marks the easternnu i limit lor th • , nu I lie Hans Meyer species 
is most closely related to S. lanicolatum L.O. Williams but differs in the size 
and shape of the pt t d th tmai , ih inn ndi um uidloaf ei i in 
the shape of the leaves. 

Species nova ad Saccoghi 

Hpiphylcctxxt tosuberect, up toca.7 cm mil. Rhizome creeping, concealec 
sheaths. PscitJohulhs obovokl, 1.25 to 1.5 cm long, up to 0.75 cm in diam< 
unifoliate, surface smooth, yellow green, leaf blade elliptic to lanceolate. 

v r > . 1.0 1.7cm. apicalk olauisc.baM< duplicative, sessile surfaces glossy green, 
texture firm.costa impressed above, hi/larcsi enrcs more or less fasciculate, lateral 
I mm base ol the pscudobulb. erect, single 1 lowered, peduncle short, 3.1-3.8 cm 
long, 0.2-0.3 mm in diameter, provided with a single sheath, glabrous. Perianth 
purple, glabrous; dorsal sepal ovate lanceolate, 1.05 x 0.33-0.5 cm, slightly 
hooded, subacute: lateral sepal obliquely ovate. 1.18 x 0.66-0.70 cm, apiculate, 
glabrous: petals broadly obo\ ate u > obi iquel v quadrate. 0.28 >. 0.24cm, apically 
truncate, attenuate at base, glabrous, median nerve slightly raised and thick- 
ened; labellum saccate, 0.55-0.66 cm high, 0.55 cm broad, 0.78 cm long, front 
part incised; column arched, glabrous thickened to the base and with a short, 
thick foot, ca. 0.24-0.25 cm long: clinanJvium in two segments on either side, 
the lower with one cirrhose stehcl on each side, the upper subpalmate with a 
subulate margin; ant her cordate in outline with a raised, thickened keel, 0.07 x 
00N , in ovai v with glahiou pedicel, ca. 1. > cm lone 

Distribution -Known only from the 1 lans Meyer Range in New Ireland, 
Papua New Guinea. 

Habitat -Mossy montane forest on trees at ca. 1 175 m elevation. 

; ■ a vm ii i /( y v —The new species has been named for Wayne Takeuch i, the prin- 
cipal collector of this orchid who submitted the type specimen to me for iden- 
tification, together with many other orchids from New Ireland. 

Notes-The description and illustration ol Saeeoolossum takeuehu was 
made from spirit material. The new species differs in I lower color from its con- 
geners (based on Takeuch is fielclnotes) and I lie I loral parts are also smaller than 
those from previous!} described axvk's 


1 wish to thank Wayne Takeuchi and Joe Wiakabu for providing the specimen 
of Saccoglossum takeuehu and John Pipoly lor his help with the Latin diagno- 
sis. To John Ohanaa verv special thanks lor his assistance with the Melanesian 
'I ok Ihsin abstract. 

ea.SNP Publishers, Mo i, i| oir. 
a-Neu-Gumea. Fedde Repert. Spec. 
1 982, D.F.BIaxell,ed. The Australian 

Book Review 

Drni Bown. 2000. Aroids: Plants of the Arum family (ed. 2). (ISBN 0-88192-485-7, 
hbk). Timber Press, 1 33 SW Second Aw.. Suite 450, Portland, OR 97204-3527, 
U.S.A. (Orders:, 800- 327-5680, 503-227-2878, 503- 
227-3070 fax). $34.95, 468 pp, 108 color photos, 50 line drawings, 6" x 9". 

Contents -Foreword tot he Scumdhdition . Peter C 1W e: Foreword lot he First Edition. Simon Mayo; 

san amacmg lainiK -so muel 

00 genera and 3200 species Fi- 

nd jack in the pulpit ', hivio 

na\ dumb cane UV//enKi l /ii«n. duckweeds i 

olliosand relatives), caladtum ( ( , ( /udnon ). go 

calla(CaHfl), taro iC.Wm asm J. water lettuce 

ie and Fugciunht/uJ. titan arum cAni.u n/i 

^»,icdcvhn,).... even those wit 

h onlva passing interest in hoitu ulturc know 

conveys so much intci 



Joseph K.Wipff 

Pure Seed Testing, Inc. 


nisctum Rich para el Manual ojGrasscs fori 

There has Ix-cii considerable il L 'b;Ui' concerning the generic Limits of Cenchrus 
L. and Pennisetum Rich. The predominant character traditionally used to dis- 
tinguish the two genera is fusion, or lack of fusion, of the bristles (e.g., Henrard 
1935; DeLisle 1963; Clayton & Renvoize 1982; Filguenas 1984, Clayton & 
Renvoize 1986; Watson & Dallwitz 1992). but its variation across the two genera 
is continuous, making the placement of numerous species arbitrary (Webster 
I) j. DcL isle ik)o3j.thoi hi in his treatment on the traditional criteria of 
bristle fusion, recognized the difficulty in the interpretation oi this character, 
and refined his generic criteria with the addition of the follow characters. 
Pennisetum has bristles that are seldom more than 0.2-0.4 mm wide, and the 
base of the fascicle rarely exceeds 0.5 mm in width; whereas in Cenchrus, "the 
spines usually 0.5 mm or (more) wader, and are generally united for a consider- 
able distance above the base of the bur, with the base itself usually at least 1.5 
mm in diameter. These characteristics, although admittedly arbitrary, ai e used 
in the present treatment of the genus Cenchrus" (DeLile 1963, p. 269). The in- 
crease in base diameter is probabb a sti uctural response to the fusion and thick- 
ening of the bristles and is closely correlated with bristle fusion. The more fu- 
sion and thickening of the bnst les that occurs t he wader the base of the fascicle 
must be to support them. 

Filgueiras (1984), using criteria similar to that of DeLisle (1963), separated 
the two genera as follows. Cench rus has fused bristles, at least basally forming 
a basal disc at least 1 mm in diameter, whereas Pennisetum has bristles to the 

base, not forming a disc. Webster U L >87) used only the presence of this disc or 
callus to separate the genera; in Cenchrus the callus is pronounced, with the 
apex llaredtolorin a discoid receptacle, whereas in Pennisetum the pronounced 
callus is absent or, when present not differentiated as in Ccnch rus species. Web- 
ster went on to say thai this character allows lor the separation of the species 
along traditional grounds, which is based on bristle fusion. In addition to fu- 
sion, Clayton and Renvoize (1986) and Watson and Dallwitz 0992) also men- 
tioned that Cenchrus usually had 'spun" bristles. I lowever, Chase (1920) sepa- 
rated the two genera by bristle type, in addition to fusion: Pennisetum has 
bristles that are usually wry slender not rigid, and are free or rarely united at 
the very base; whereas Ccnch rus has rigid bristles that are united below. 

Webster (1988 J stated that even within a number oi species it is open to 
interpretation as to whether the brist lesare fused or the callus flared. In regards 
to bristle fusion, Pennisetum ciliare(L.) Link is extremely variable and has been 
treated in both Cenchrus and Pennisetum. Hignight et al. (1991) evaluated 800 
accessions of P ciliarc collected in South Africa and selected accessions based 
on extreme differences in morphology, including differences in bristle fusion. 
Thirteen of the most diwrsc morphological types were studied for morphology, 
cytology, and lertility. Fiveol these diverse morphologically topes were used in 
hybridization studies with a sexual genotype (Bashaw 1969) of P. ciliare. Though 
they lound most accessions to have at least some lusion, two of the accessions 
studied had a complete lack o! bristle I iision. These plants were verified at Royal 
Botanical Gardens, KewUx) to be Pri/mrc. 1 Ivbridization studies with the sexual 
genotype showed a close relationship between t he plants. Some of the F t prog- 
eny from the hybridization studies segregated for union of bristles similar to 
the bristle fusion found in Cenchrus setigerus Vahl. I hgnight et al. (1991) con- 
cluded, "that bristle union is an arbitrary character that varies with genotype 
and is unreliable for the taxonomic classification ol buffelgrass [P. ciliare]? 

Read and Bashaw ( 1*0 j hybridized the same sexual accession of P.ciliare 
with an apomictic accession o\ C setigerus. The resulting progeny represented 
a complete intergradation in morphology between the parents. Read and 
Bashaw concluded that the chromosome homology and cross-compatability 
ol P. i Hi are and (.. setigerus, plus the high lertility and morphological intergra- 
dation, observed in the b| progeny provided overwhelming evidence of a very 
close relationship between the species and concluded that they belonged in the 

Sofms U9 L n)exam i nation ol fascicle organization in eight species o\ Cent hrus 
and six of Penniset urn suggests an add it lonal differentiating character: whether 
the axis of the fascicle is prolonged as a, usually prom ment, bristle {Perm iset u in) 
or terminates into a spikelet and is not prolonged (Cenchrus). Unfortunately, 
the prolonged bristle in Cenchrus setigerus and P. clandcstinum, although 
present, is less prominent than in the other species ol Pennisetum studied and 

was overlooked by Sohns. This m,iv he win most subsequent taxonomists con- 
sidered the presence or absence of the prolonging bristle not to be of generic 
signiiicancc. Also, this character has historically been evaluated as a secondary 
character in conjunction with bristle fusion (e.g., Dehisle N63), which is known 
to be arbitrary in its separation of the genera, and would explain why Sohns' 
character has appeared to be ol little laxonomic value. 

Avdulov (1931) and Nunez (1952) reported that the genus Cenchrus has a 
base number of x = 17. Pohl (1980) used chromosoi ne base [lumber as part of his 
generic criteria. 1 le distinguished the two genera on the lol lowing characters. 
Cenchrus has inner bn.stles that are spmed ike or pungent, are usually retrorsely 
scabrous, and usual!)' have a base chromosome number of .v = 17; whereas 
Pcnnisetum has bristles that a. re not spincdike m pungent and are antrorsely 
scabrous; and have base chromosome numbers o\ 5. 7. 8. or 9. However, despite 
these observations, Pohl later (Pohl & Davidse 1994), without explanation, fol- 
lowed Dehsle (1963), kilguemas (1984), and Clayton and Renvoize (1986) in his 
generic concept o\' Cenchrus and Pcnniscum. 

brom the examination ol specimen ol t he lol lowing species of ( cm /mis and 
C. distichopyllus Griseb., C. brownn Roem. & Schult., C. echinatus L„ C. 
C. platycanthus Anderss., C. spinijex ( a v.. C. trihuloidcs I..: Pcnnisetum advena 
Wipff & Veldkamp, P. alopecuroides (b.) Sprengel, P. annum Mez, P. 
hambusijormc (Fournier) Ilemsley. P. ba^edowii >uinmrrli. P chilense (Desv.) 
Jackson, P ciliare (L.) Link, P clandcstinum Hoch. ex Chiov, P. complanatum 
(Nees) Hemsley, P crimtum (Kunth) Sprengel, P. distachvum Rupr., P divisum 
(Gmel.) Henr, P domingense (Sprengel ) sprengel, P durum Beal, P elymoides (F. 
Muell.) Gardn., Pjlacadum Munro ex Griseb.. Pfrutescens Leeke, Pglaucum 
(b.) R. Br, P hohenacken Steud., P Jwdeoic/es (Lam J SteucL P intectum Chase, P 
karwinskyi Chase, P /conUn/n Klotzsch. P bui/nhmn Sprengel, P macrostachys 
(Brong.) Trin., P macrourum Trim, P mmssmaun Stapf, P mczianum Leeke, P 
montanum (Griseb.) Hack., P nervosum (Nees) Trin., P occidentale Chase, P 
orientate Rich., P. pauperum Nees ex Steud., P. pedicellatum Trin., P. 
pennisetiformis (Hochst. & Steud. c.v Steud. J Wi pl'f, P peni vmn urn (Doll) Trin., 
P petwlare (Hochst.) Chiov., P. poly stack hm (L) Schultes, P prieurii Kunth, P 
prolificum Chase, P pu rpureum Schumach.. P omasum (1 lochst.) Schweinb, P 
rigidum (Griseb.) Hack., Prupesl re Chase. INuwUmniM Henr I' setacc urn (.Fovsk.) 
Chiov,Psetigeri ( m(Vahl)Wipff,Psot ) i ( dcnts'is(C:lavton)\Vipff,Psphace?atum 
t Neest Dim Sr Sch t nz. P.squa mu lut ti/n Prescm. P mm/ Pohl, P.thunbcrgi t 
Kunth, P tnstadiyon (Kunth) Sprengel, P. uniscium (Nees) Benth., P villosum 
R.Br, ex Fresn., P vulcanicum Chase, and P. wehcrhau ri Mez; as well as cytologi- 
cal examinations of L ) species of Ccnchrusand 2ospeciesoi Pcnnisetum, and in 
addition to the cvrolojncal work already published (lor a review see Jauhar 1981; 

Wipffl995;Schme] :er isconcluded that thcgcncru interpretation thai 
Pohl adopted in 1980 is correct phvlogenctically The decree of fusion of the 
bristles is generally unreliablcat the generic level and should not be used as the 
primary character in separating the two genera 

The following characters are considered the most important in delineat- 
ing the two genera: 

Pennisetum.l) bristles are not spine- like or pungent and are an trorsely scabrous 
(one South American species is both antrorse/retrorse); 2) the axis of the fas- 
cicle is prolonged as a, usually, prominent bristle: >J inner bristles free or fused; 
and 4) haw base chromosome numbers ol 47.8, or 9. 

Cenchrus: Dinner bristles are spine-like or pungent, and usually retrorsely sca- 
brous (when antrorsclv scabrous, the inner brist les are fused and not grooved); 
2) the axis of the fascicle terminates in a spikelet; 3) inner bristles are fused, at 
least at the base; and 4) haw a base chromosome number ol x = 17. 

Though, there are still species in Com lincsand Penmsetum whose generic 
placement still needs clarification, bor example, C. myosu nudes Kunth, which 
has a base number of x = 9 or 10 and a fascicle structure very different from 
Cenchrus s.s., as well as some South Pacific taxa. The process of obtaining the 
materials needed to resolve these problems has begun. 

New Combinations in the Punniseh'm Ciuare Complex 

Penniselum setigerum (Vahl) Wipff.comb. now IvWiono M: Cxiuh russcl igerus Vahl, 

Fmum. Pi. 2:395 180vl\auu>rt(un nililii Kunth. nomollcg.. Rev. Gram. 1:49. 1829. 

/VMMiNvlinMci/i t MrU..)^n(mlVahbLeeUe./..Nalur\viss.7^:22.1907. 

( \; h h > us,:, inn is I .. se! iiynu l Vahl ) Maiiv M Wcilcr. El, Air. Nord. 1:342. 1952 

Typf.: Arabia. Forsskal (HOl.OTYPE: C!). 
Fisher etal. (1954) reported that the type of reproduction was identical between 
Pcnnisctumcilian and ( .rue/i i is and that ihen was continuous variation 
in morphological eharaeierist ics between the two species. I 1c concluded that the 
two species are members i)\ a single agamic complex. Snyder et al. (1955) also 
reported that these two species had similar reproductive behavior. Bashaw (1953), 
alter studying tin. morphologv.cvrolog\ md modcof u production ol Cenchrus 
setigerus, concluded that (7 sen ernes and Penmsetum aliare were "much more 
closely related than our preseni elassil icat ion indicates, perhaps even varieties ol 
the same species." Oelasle U9o g). alter examining speci mens of each taxon from 
throughout their range, only observed a lew specimens that could be considered 
intermediates and recognized the two taxa as distinct species of Cenchrus. 

Read and Bashaw U9o9j hybridized a sexual genotype of P.ciUare with an 
apomictic genotype of (Tsc/igcnes. The result ing k population consisted of both 
sexual and apomictic plants that represented a complete intergradation in 
morphology between the parents. They also stated that some o^ the hybrids were 
so different from either parent that populations from them might be mistaken 

tor new species. The hybrids were highly fertile and had fewer quadrivalents 
and more bivalents than either parent. The\ concluded that the two species 
wen Mii.inl uhuj. [inn nrljio ii'h ow [miIh Ih \ noted, however, that 
"They have been effectively i olaied in natun b\ obligate apomixis and their 
morphological distn it tno w,« aiilicient to permit valid taxonomic treatment 
at the species level. It is also apparent that with sexuality in buffelgrass 

different hybrids. At present it would be convenient to retain specific rank al- 
though we feel it would be justifiable to merge the species.'" (Read & Bashaw 
1969, p. 806). Although they recognized both taxa as species of Cenchrus, Read 
and Bashaw stated that it might become necessary in the future to reconsider 
the generic rank of this entire agamic com pi i -, 

JVntiisiium [h musculo! iih I U h r n ml i Wipll lomb no\ IFigs. 1, 

2) U h l ( , ii, i r U p, . i and . x a, iul s\ n ['I Uunoo 

L:KH 18>7. Ho: Saudi Arabia: jedda. "In dcscito pr. oppid. neschedda.," 28 (an 
1836, Schimper 973 [lectotypi heie deseuiaiol L iwi o i. -ni-i s- K! (3 sheets)]. 

a l(K I I ISi | l v 1 . Huiioi I , 'i IHI , ' M I I II i p I, | ] 

of these collections were examined from P and louiul lobesimilai. .0 ImmpcrsPo o.o 
chosen as the lectotype because duplicate sheets are known to exist at K. whereas 
presently there isonlv one sheer o\ iilniipo l, 77 know n to lie in existence 
Delisle (1963) consief ud (. ,, do u< naiv ///mmoa p ,\ i ol ( c thai is Clayton 
(1982) reported that the boundary between (.. i iliai is and C. pennisetiformis 
was indistinct, but that th peca could b< eparated follow G i t s'i n 
pennisetiformis has inner bristles basally connate for 1-2.5 mm of their length, 
is usually annual (.short- load perennial;, smallci in statttivand found mostly in 
sub-desert grassland ,. w hen a. ( ( ilums has tin inner bri ales basally connate 
for CO )0.5 ha mm o\' their length, isastout perennial, witli or without rhizomes, 
usually forming a Inn d knottw ;ometimi dmost wooch b i ncl i toumlin 
deciduous bush land and wooded grasslands. 

s(Cl a ton ) V\ ipii . omb no\ L.oa re, ir In io on l( |, n j 

Clayton, Kew Bull. il:Y 8)77. Tvrr: SOMALI LLLUBLK7: Loeavo, sOOQ-7000 It 

[IW I 71 » oni undo huh oi huh nd tn Nov 1938 \ WcKinnon S221 (ho- 


C day ton ( 1977 J reported that C. somalensis and C penniseiilor.nis were closely 

related, but that ( sonw.lensi i . lensely tufted perennial with in rolled leaves 

about 1 mm wide: whereas, ('..pennisetiformis is an annual cm short-lived pe 

rennial, with flat leaf blades, 2-5 mm wide. 

I am grateful to the following herbaria lor loaning specimens: t La . \|py \y p 
US; and to K for providing photographs of type speed mens. I would like to thank 
Mary E. Barkworth (.UTQ and Kathleen M. (lapels U.TO lor assistance while 


Ed. II. Holu ■ ■■ ■ 

UTCs assistance in obtaining loan oi i p< j'uinm md to Stanley Jone 
(BRCH) and two anonymous reviewers lor their insightful comments. 


Bot.44,Suppl.:1 19-123. 

Bashaw, E.C. 1953. An in\e. tn nil n Ml n >n I I I rvtology,and mode of reprodm 

tionofCenchrth r ,,•' i It I nun I f 1 l Jniv^i >ity, College Station. 

F ii bo C't ' I'nuhti iti t )t I tli u 1 , I- ni | I Lin ih'p So. 9:396. 
CHASE, MA 1 920.The North American species of Cenchrus.Contr. U.S. Natl. Herb. 22:45-7 

( i \A D and SHli 1 <t i , i „ ,- ft 11 \ I, F .1 Polhill, ed. Flora of 

Iropual hast Africa.AARalkema,Rotteidam. 
Clayion, W.D. and S.A R ii u 1 8n i „ n. i i ii imiimm in . it the world. Kew Bull. 

Addit Pei 1 ; . Hi i f 1 i|( n l t.iti.'h. p i M 1 1 l i> [ > mill m 
Delisle, D.G. 1963. Taxonomy' and distiihutmm ot the genus ( cncivus. Iowa State J. of Sci. 

Filgueiras, T.S. 1984.0 Genero Cenchrus L No Brasil (Gramineae: Panicoideae). Acta 

Amazonica 14:95-127. 
FiSHER,W.D.,E.CBASHAW,and I .( .Hon. HGAI vidence toi upomixnin Pennisettimciliareand 

Cent hit, < I genu Agron I 46:401-404. 
Hi i n IT I I I i iti ti t i i i ii i i < [ i imi i tr ,| I 

Hignight, KW, EC B i .aiulM-l I "' d i J n i md mm phological diversity 

of native apomictk PnlfelgiasAV/mnepmmaGpm (I .» I ink. Rol.Gaz. 152:214-218. 
Jauhar, P. 1981. Cytogenetic and breed u m of pi ill mUei and mimed .pedes. Vol.1. Progress 

and lopu int-tm-iH'i n AA audU in e I -'don P I is*, Inc., New York, NY. 
Nune/,0. 1 952.lnvestigai mnes cariosistematicas en las Gramineaes Argentmas de la tribus 

"PaniceaedRevista I am Agron. Univ. Nam I a Idata 28:229-256. 
PohpR.W.,l amilvPi d-.lidWiiiiain Ruigei.ed.l lot a Costaricensis.Fieldiana 

I' Mn i h > Man No 1 I h I i f lu - inn f N ituial Hi l a ( hi igo II . 
Pom , R.W., and G, Davh oi , 1 9P4. Cenchrus I . Pp 374-375. In: G. Davidse, M. Sousa S., and A.O. 

i h iter, ed lora Mesi »americana, Vol 

Rfap, IC and EC B i I ' ', m i [oi , mi. m i ii i aimul \< md the role of apomixis in 

4" ' lalion in huff. ! in , md laid I ;m ( i, [ P i ■ or i)n 

Schmfizfr,G.H. 1989 / 5e tu i . ; i « t t 1 Morphological and 

gen; tn anation mm mam |t - rig « lid I tt.iti i ^andbouwuniversiteit 

Sohns, E.R. 1955. Cenchitr, and Penniseltinr. I ascicle moiphology. J. Wash. Acad. Sci. 45: 

Sn i ii I A, A R H i i I I I I i i i ii mi ii Pi i mi m mn t /> 

c/7/are.Bot.Gaz. 11 6:209-221. 
Steudei, E.G. 1854. Synopsis pl.mtar urn gluma. eumm.PaP hi .lainineae.J.B.Mca/leraStuttgait. 
Watson, I. and M.J. Dai i wm-.(lP92+). 'Grass genera o( the work Id inscriptions, illustrations, 

id* mill iti 1 1 m i ii it in a hi a il it i ti, i n in i i p h i i m 1 tn 

physiology, phyto< d. man t I i i ti ti i \ ,ti i u i d nd local distri 

bution and reft it i - lag G It mm- In f i , m i u lath August 1° <" 
Webstir, R.D.I 987. The Australian Paniceae (Poaceae). J. Cramer: Berlin. 
Websiir, R.D.1 988. Genera of the North Ameiican Paniceae (Poaceae: Panicoideae). Syst. 

Bot. 13:576-609. 
WipffJ.K. 1995.A hit i init ti t e ti ih ,t , i ,i tu species of Pennise- 

tum (Poaceae: Paniceae-) and their hybrids. Ph.D. dissertation, Texas A&M University, 

College Station. 


Jose M.Bonifacino 

°aseo del Bosque s/n, C.P.I 9G 

o/v.'os l.'iS.Moin^Kk'oJiRUGUA: 

Gisela Sancho 

,an< ho.niwkmNMNH.SI.FDU 


Permanent address: 

iltadde< lenaasNaturalesyMuseo 
Bosque s/n,CP. 1900 La Plata, ARGENTINA 



. /\\,'u'« l >/>ll:H 1 

a (Cabrei 

w) Nesom and NanlophyJlum pdtd^onna 
mean Patagonia that were originally desi 
cacpappusm l°o3and No 1 -), respectively. 1 
•ithin the genus rvi/rdnprivlhun. arguing i 


etweeu the pah u , o , p ippu "1 h i\n ,'/ lie 1 .1 u 1 Pdidd, I p | 11 mi i ih p p'|" 
I I ll [ 1 I 11 

II, II 1 lll| 11 |> ll I. hill III u HI 1 1" Wl /, / l| H [ , I Mil I I I U I mi " ' 

,, ,P uhUmIh in 1 pi mil 11 pih u ol \unhplnlhu,nh unmlaand \anh< In Hum r ,i 
re wide and enclose the I lords, while in Ndulophs tlu in species the paleae. if present.; 


mnnoiipii o \\ I 1 pt !,, 1, I ' d, ,n poppio. 11 ! l h i \ P> (1 > n , [V , n mi 
Nesom iih lu\ o . \v,',i, op/iooi \ dolompMppio dent ro del genero \di\hph\ ! 
dis, imiiiiuiiLulis cut re el papo palaeeo de MThophdv.i v PiiluipM 
Xli ; J,,, 1, (Pin, 110,11 niu m, on id, 1 n u toscomogi n, 10 111, , pi lulu 1 

Aylacophora y Paleaepappus, respectivamente. 

Endemic to Argentinean I 'at agon ia, Nardophyllu mdcscrlnola (Cabrera) Nesom 
and Naidophyllutn paiayouu urn (Cabrera) Nesom are 2 of the 1.0 species recog- 
nized inside Nardophylluin I look. & Arn. by Nesom (1993). 

Ncm/o/'/n Hum de sen uola and A. paUigoiueum were originally described 
by Cabrera under the monotypic genera Aylacophora and Paleaepappus. 
Aylacophora (Cabrera b)"C>) was characterized by its paleaceous receptacle, scaly 
pappus,and compressed cypsclae wit h 1( })ciliate ribs Paleaepappus (Cabrera 
1969) was defined by its paleaceous receptacle and its pappus of 7-8 paleae. In 
contrast, Nardophyllu m sensu Cabrera 1 N54 ) has receptacles naked or with only 
3-6C-13) paleae, a pappus ol bristles, and terete, more or less pubescent cypselae. 

According to Nesom, the discontinuities between the paleaceous pappus 
ol Aylacophora am] Palcacp<ippus;uu\ ilic bristles oi Ntad.ojdiylluin pappus do 
not support their distinction from Nardophyllum because there is a tendency 
for the pappus bristles to be somewhat flattened m Nardophyllum. 

However, according to our observations and the interpretation of the data 
gathered (see below), Ayhuophora and Palcacpitpus should be considered as 
independent genera from Nardophylluin as follows: 

Aylacophora Cabrera Pol Sex Argent Hot 4:200 19=) 3 I n Spkcies: Aylacophora 

deserliuha Cabrera. Hoi. Soe. Argent. Rot, 4:208. NSy \,udophxUum\lcscrtueld 

u abrera.) Nesom. Phviologia 75:3o2. 1003. Fig. 1 A-D. Tu-'i-:: ARGENTINA. 

PkoviNOiA Np.uqufm Plaza llunuul, 12 Apr 1052. A.I. A ahvni /I252 U mi otyh-: 


Shrub 50 cm high, densely branched; old branches aphyl Ions, bearing furrows; 

new branches with sparse nodes; leaves 1 1 near; capit ula discoid, solitary at ends 

of branches; receptacles paleaceous, paleae wide, apically 

pubescent, each palea enclosing a floret; cvpselae compressed, 2(-3) nerved, 

pubescent only on the ribs; pappus of 10-11 oblong scales no longer than 1.2 

Ecology- Ayhuophora desertuola inhabits semidesert areas in Patagonia. 
The very few herbarium specimens o\ this species lead us to regard them as 
very narrowly distributed., According t o lug. SteibeKpers.comm.), A. deserticola 
Cabrera grows on edaphic communities in the Monte biogeographic province 
(Cabrera & Willink 127 >), where it is present on sane! dunes with very sparse 
shrub cover, associated with Larreadivaricata,Atriplexlampa,Prosopisflexuosa 

Fig. 1 . A-D, Aylacophora desertia 

patagonicus. E. Capitulum. F. Receptacular 

bracteolatum.l. Capitulum.J. Receptacular palea. K. Cypsela with pappus.L. Pappus bristk. M-O.Nardophyllumbryoides. 

M. Receptacular palea. N. Pappus bristle. 0. Cypsela with pappus. (A-D, based on Cabrera 11053, LP; E-H, based on Rio 

Chico, 1900, Ameghino s.n., LP; l-L, based on Sena 77, LP; M-O, based on Ruiz Leal 27011, LP) 

solbrigii,Fabiana patagouica i\ne\ Larrca cuiu'ifolia. Aylacophora deserticolahzs 
been collected in Argent i na, Prov. Neuquen, Dptos. Conl I uencia, Cutralco, Anelo, 
Pehuenches, and Zapala. West ol Rio Covunco and south of Rio Neuquen in 
Dpto. Zapala, A. descrtuola is the dominant species regarding surface cover. 
Flowering in the fall. 

Paleaepappus Cabrera, Bol.Soc. Argent. hot. 11:27 >.ls>o9. i Mnspn irn J'dcac/><i/>/nis 
/HJfdgcmcu.s Cabrera. Bol. See. Argent. Hot. 11:27 11*0 Nurdoph\Uumpata^onieum 
K .ahivrajNesom.Plnaologia 75:302. 1001 iMg.l li 1 1. 10 n : ARGENTINA. Chubut. 

Shrub densely branched, lateral brandies sharp ended' leaves oblong to spatu- 
l.iie, eonaceous; eapiiula discoid, solitary at the ends of branches; involucres 
campanulate; receptacles paleaceous, paleae wide, apically pubescent, each 
enclosing a floret; cypsclae terete, densely pubescent; pappi of 9-10 elliptic 
paleae 7 nun long, in 2 series. 

Ecology —Paleaepappus piitagonicus inhabits semidesert areas in 
Patagonia. Knowledge of the ecology ol Paleaepappus is scarce and speculative 
because the only record o\ this species is t he tvpe itsell . and there is no ecologi- 
cal information on the label. The conservation status of I his species could aptly 
be recorded as endangered. 

Aylaeophoraand Pa Icac pappus are stronglv segregated from Nardophyllum 
by characteristics of the elements of pappus such as number, shape, and num- 
ber of series. The paleaceous pappus o\ AyUuojmora and Paleaepappus con 
trast with the bristles m Nardophyllum. Intermediate states of pappus shape, 
like narrow paleae. ha\c nor been found m Nardophyllum. but occasionallyflat 
bristles have been found in Nardophyllum. 

Nardophylhun circumscription 

Nardophyllum I look. & Am. (1830) was described by Cabrera (1954) as shrubs 
densely branched, leaves alternate, small: heads solitary at the end of the 
branches; discoid capitula, involucre campanulate; receptacle small, convex, 
naked or with few paleae; cypselas turbinate, 4-5 ribbed, hairy; pappus com- 
posed of several brist les. The circumscri prion ai Nardophyllum adopted here is 
the presented by Cabrera (1954) in his revision o\ the genus, where he included 
7 species: Nardophyllum armatum (Wedd.) Reiche, N. hraeieolatum llauman, 
N.genistioules (Phil.) Gray, N. hryoides (Lam.) Cabrera, N. chilwtruhioidcs 
(Remy)A.Gray, N. /minima (MeyenjCabrera.and N.obtusi folium Hook.& Arn. 

Palraepappus ,\ad \anloph\ lliun 

Contrasting with the pappus o\ Paleaepappus (big. 1 II), the pappus ol 
Nardophvllum is com posed olea. 30 bristles 5-10 mm long (.Fig. 1 L),sometimes 
flattened, especially ar the apex (lag. 1 N), arranged in 2-3(-5) series. 

Involucres and shape o\ Palcticpiippus cvpselac arc similar to those of 
Nardophyllum. Differences in number and shape of receptacular paleae are also 
found in these two genera. Palcacpappus has paleaceous receptacles with wide 
and apically pubescent paleae enclosing each floret (Fig. 1 F). The receptacular 
paleae of Nardophyllum are narrow, do not enclose the florets (Fig. 1 J and M), 
mid vary from absent cV. armalum ) to Vo> (rarely more numerous, 9-13 in N. 

Aylacophora and Nardophyllum 

The pappus of Aylacophora (Fig. 1 C and 1 D) contrast highly with the pappus of 
Nardophyllum (see above). In add it lon.ot her characters distinguish Ay /acophora 
from Nardophyllum: the involucre ol Aylacophora is globose I Fig. 1 A); its cypselae 
are compressed, 2(-3). and pubescent only on the ribs (lag. I C). The involucre of 
Nardophyllum is campanulate toobcomcal (Fig. 1 band the cypselas are terete, 
(4-)5-7(-8) nerved and uniformly pubescent (Fig. 1 K and 1 O). 

Aylacophora has a paleaceous receptacle with wide and apically pubes- 
cent paleae enclosing each floret. The number and shape of receptacular paleae 
of Aylacophora (Fig. 1 B) contribute to set this (axon apart from Nardophyllum. 

Aylacophora and Palcacpapus were placed in the Chihotrichum group 
(Zhang & Bremer 1993; Bremer 1994) that includes shrubs with mostly densely 
set. coriaceous, ami often abaxial \ tomentose leaves, and for most of the genera, 
paleate receptacles (Bremer 1994). Within this group there are intermediate 
morphotypes for pappus elements that range from terete bristles in 
Chilliophyllum, through narrow paleae in Pcpidophyllum (Cabrera 1971), to 
paleae in Aylacophora and / \i lee epop/arv In addition, the number of receptacular 
paleae vary within the Chiliad ichum gmup. In i defence to this ehacu tor the 
variation observed wiihin Nardophvlluiu isalso present when comparing other 
genera of Chihotrichum group, such as Lcpidophyllum without paleae, 
Chiliotrichum with lew paleae. and ( '.hilioplwllum and Chiliotrichiopsis with 
fully paleate receptacles (( labrera L971, 1 978) Because the gradation in pappus 
and receptacular paleae is inherent to t he Ch Hint richum group itself, this varia- 
tion cannot be an argument againsl the recognition of both Aylacophora and 
Palcacpappus as distinct from Nardophyllum. 

Genera o\' Chili ol rich tun CI roup i ensu bremei F-W4) can v identified as 


in 2-3C-5). 



1 1 i iiif ii . 1 ill, h, 

in ,i,-,n n, 

;d to the ribs. 


?ceptacles epaleate. 


Nesom's reinstatement of Nardophyllum scoparium Philippi (Nesom 1993) is 
not accepted here. Presence oi pistillate ligulate florets in the periphery (5-6) 
casts serious doubts ;il km n posh ionmgthisiavon inside \ at doph villi m. The taxo- 
nomic placement of this Chilean species and its possible status as an 
undescribed genus allied to some genus inside Chiliot richum group as Nesom 
suggests (pers. comm.) is being reviewed by one oi us ()MB) and will be pre- 
sented as a more comprehensive study of the (dwliol ruhum group (in prep.). 

According to Bremer ( .1 W), Tvlm ophoi a and Pakacpapput. ;are very similar 
and possibly sister groups. Ayldcoplh'/u was related to Amhcpliv! I urn and Crnliofri- 
ohiopsis by C Cabrera \ l l > : > U Palcitop>opj >m w a-, i elated lo \m< I. >pliyllum and Avldou 
p ho ra (Cabrera 190Q) Ouropinion, based on observations of evpsclar morphology 
number of series of the pappus elements, in vol uc re shape, leaves shape and plant 
habit, favors a closer relationship between Palcacpappus and N'mt/onliyllimi. 

Nesom (1993)suggest that the narrow, internally tomentosecaulmesulcae 
oi Aylacophora could be homologous \\ ith t Inwo hu.incl in somespecicsol Nardo- 
/dn'l I am and so denoting a closer rclationshipol Avlmuplamc! to Nardophyllum. 
To complicate the matters further, the paleate receptacle i^i both Aylacophora 
and Palcapapus, would favor a closer relationship o( these two genera with 
Chi Hot richiopsisX'.hi I iophyllum and Cli iliot nr hum instead, which are, as Nesom 
(1993) points out the closest relatives to .VmlnplivlImM and all oi them have a 
paleate receptacle X'hi Hot rich topsis, Ch i '/it>p/n I hi m I ul I \ paleated,Chi/iotnehum 
15-21 paleae). Evidently, as Bremer U994) states, the elucidation of the phylo- 
genetic relationships among these genera demands a more comprehensive study 
oi the Chiliotrichum group that is beyond the scope ol this paper. 

Segregation ol Ay/a. nia'i.M a and Adem />np/)m Irom Nmdn/'liyllum on the 
basis of pappus shape is supported by other characters such as the shape ol 
both cypsela and capitulum in Aylacophora, and the shape and quantity ol 
reeeptacular paleae in both A\ lu< ophoro and Palcacpappus. 

Although Aylacophora and I'alcacpappas are closely rehired to the rest of 
Nardophyllum species, the discontinuities basically observed in the shape ol 
the pappus elements, with no defined intermediate states, justify the consider- 
ation oi both species as t wo distinct genera. 

Otirconclusion issupported b\ Nesom el al.Un press). These mil hors. based 
on features of the involucre and mainly on pappus morphology have included 

a new Peruvian spec a so! uu Wi cu inside On /ml n«;hiopsis Cabrera 
(OnliotrichiopsLS pcrin'iciiirt Nesorn, Robinson & Granda). Nesom et al. (in 
press) found that the pappus moi phologj i good < harai ter to separate gen- 
era inside Cihiliotrichum group and concluded that Cabrera's narrow generic 
concept of Nardophyllum w \ b< tt< i d< < iption of the diversity inside 
Chiliotrichum group then fore ha\ ing incf pi ndcnth an ived at the same con- 
clusion we have, regarding the consideration of Aylacophora and Paleaepappus 
as distinct from Nardophyllum. 

Additional specimens examined of Aylacophora deserticola Cabrera. ARGENTINA. Neuquen, Plaza 

Huincul, 11 Dec 19* L hi II, H il 1 ][ h i , H Troiani etal 12504 

(SRFA);PasodekoslndiosaCutrako 5h [ 1 < - , ft i n ) RFA), Paso de las Bardas, 

2 Feb ]999, H.Troiani\ -e, " I I, II Indtos, 5 Feb 1999, P.Steibely H. Troiani 

/4069(SRFA) ; Paso del o liulio oIihImh it ,1 HP" ' \ H Troiani 14073 (SRFA) 

Nardophyllum. Nardophyllum armatum (Wedd.) Reiche. ARGENTINA. San Juan: Iglesia, 
camino a el Fierro Cam h d I i _ it \ h 1 >i I P) Pampa de Pauacan, 

entrelasAguaditasyChepical, 12 D - , , LP). La Rioja: Gral.Sarmiento.Rio del 

I i I i { 1-jn ii d t , i 

Salta: San Antonio delosCobres, 2^ l.n I n 'I P).Jujuy: 1 km al Wde la Quiaca, 1 1 

Feb 1 960, Meyer & 

Esquinas Blancas, 22 Ene 1966,Gv/vm < - If . H i , Pampa, Feb 1937, Cas tellanos 20229 


Nardophyllum bracteolatum Hauman. ARGENTINA. Mendoza: San Carlos, El Pedernal, 25 
Mar 1916, Sanzin 1810 (LP);Tunuyan, Paso del Portillo, Cuesta de los Afligidos, 29 Jan 1 934, Ruiz Leal 
2052 (LP); San Carlos Pnu >nd I > L n I I it I 41 ~ ' (I r i an Carlos, Arroyo de la 

Qda/'Casa de Piedra," 1 7 Jan 1 952, Serra 77 (LP). 

Nardophyllum bryoides (Lam) Cabrera CHILE. Magallanes: I ii |u Naciomll ti III i 
Lago Paine, 1 7 Jan I At uLNTINA. Santa Cruz: alrededores de El 

Chalten,10Feb2000,P<»/i i a i il L i i i m. rlinero.Cabo Virgenes, 

Mi l'4ho//n (I PI Riod illt-go r h 1 , H r i E i le las Vizcachas,Cerro de 

(LP);LagoArgentino,ParqueNacionalh< J i t hi I Ml i 1 I -crmW.25864 (LP).Tierra 

delFuego. Est.Cullen,52°44'5,08° > in i ^ , - -, ,> il P) Bahia Lee.52° 52' S 
70° 1 6' W, 6 Nov 1971, Moore 2339 (LP). 

Nardophyllum chiliotrirl- ii I-.- ARGENTINA.San Juan:Calingasta,Qda.Los 

avestruces (oeste de Cerro Castanet F I I i • > m .4 (LP), Rio Mauriquc a 

Pmt -II, |, I, ' tl tilt. . Iltl 1" 1 • I • hi. Ine , I hi, lIF ,1 II I, I, h 1 

1947, Soriano 2786 (LP); Ravs. m, 1 ' I in il f ft, I I, 1 1 40, i astruvie,o & Lopez 23 13 (SI). 

CHILE. Santiago: Cotdlhtad, I it Ft it i hi it, u> I in huh UCr/ermcmn 621 (LP). 
Nardophyllum lanatum (Meyen) Cabrera ARGENTINA. Mendoza: Malargue, ruta 40,20 km 
L > alf,u - 1 h I, 1 i- ' I NPi.qi.on: I , ■ ■ I u 

Calmuco,15 Feb 1942 Ft j n I i lm Pio Barranca y Buta Ranquil,8 Feb 

1 950, Boelcke 4235 (LP). CHILE. Colchagua fen 9 Feb 1 96i <ner833 (LP);Vegas del 

Flaco, al E de la Qucl>t 1 I i I I It hl<h L II t III s >•" (LP). Valparaiso: 

Cerro Roble, Cordtlk n I I, n h l t I I I Ovalle: Ge/sses.n (LP 60269). 

Nardophyllum obtusifolium Hook & Am. CHILE. Magallanes: an Gn gorio, 76 Oct 1 968, 
Cekalovic 33903 (kP). ARGENTINA. Santa Cruz: Pto.Sai i luliai i I M i n (LP) Corpen Aike 20 

hnHIVHidRiK'm.Kut.i o \ ! >< ■. I" l/l^t^fi < ', ■' >M L),L,.in|cnL b astillo.11 Feb 1936,5coff 

Nicora3638 (LP); Pueito [ >. < nio I in 1 ^ A/bo// V il I ) I gu. I ' Apt 1 946, Scotnik 306 (LP); 
Leleque, 13 Jan 1947,5ooono ?3A I (I P); hst. Pcpita, Alto Rio Viuior, 1 i Fob 1 947, Soriano 2585 (LP). 
Neuquen:Charahui!h A,, , I ,, , I I I 1 I I i I i inn 1 1 lm a Q Jan 1966;camino 

aNinhuau, 16Jan !■•- .-. . ■. ' L P Santa Cruz: olt alios Pellegrini, Est. La Flora, Dec 

I ' " " > inqen 2 (LP). 

It is a pleasure to acknowledge Lihana Katinas and Jorge V. Crisci lor helplul 
critical comments on the final manuscript; tiny Nesom lor his helpful critical 
comments on this paper, making available lo usihe inanuscript "A new species 
ol ' Chiliotrichiopsisi \ teraceai \ tereae ) from Peru. (m\ Nesom, Harold Rob- 
in on no! • 1 1 u ro t ,i unl.i Patu ai ' Bi ilioni i.mpn i m I In en< mi n i nit m 
towards publication; John Strother for his corrections on the final manuscript; 
lng. Agr. Pablo E. Steil x I for t he i n formation regarding the ecology of Aylacophora 
and its geographic on me the< on .ejoNai uMialif InvestigacionesCientificasy 
Tecnicas de Argentina (CONIC.LT) and ( . '.omision Sectorial de lnvesfigacton 
(dentil ica(CSIC), Uruguay, ioi the financial support on tins investigation, and 
LP, LPAG, MVF, SI, SRFA and US for loans of herbarium specimens. 


Br[mer,K. 1994.Asteraceae Cldcli th it I 1 ti bit hi il i I i Portland, Oregon. 
Carrfra, A.L. 1953. Un nuevo genero de astemn , dt a bepubl a Argentina. Rol 5t> Ai 

gent. Bot. 4:266-271. 
Cabrera, A.L. 1954. Las especies del genero Nardophyllum. Notas Mus. La Plata, Bot. 83, 

CABRbRA, A.L. 1969. Compuestas nuevas de Patagonia. Bol.Soi Anient. Bot. 1 1:273-275. 
Cabrfra, A.L. 1971. Compositae. En M. N. Correa (cd.), LI Pataqonica, Colecc. Ci. Inst. Nac. 

Tecnol. Agropecu. 8( 7): 1 45 1 . 
Cabrfra,A.L. 1978 ( on if it). In' Cabreia LI I. Pi uju\ le Ci.lnst.Nac.Tecnol. 

Agropecu. 13(10):! -726. 
Cabrera, A.L. and A. Wn no 1 ' I i i p oi ill i It m ' t ili'io ' E a bene de Biologi'a, 

Monografia 13. Washington, D.C 
GrauJ. l \ , toman P > m: In M IL \ *>, I | I Lirl mine, and B.L.Turner, eds. 

The biology and t bemisliy of the Compositae ). At aclemic Press. London. Pp. 539-576. 
NtsoM,G.L 1993 laM in mi inn i i ' i i v eae Astereae) with 

observations on the definition of Nardophyllum. bbytologia 75:358-365. 

Astereae) from Peru. Bnttonia (in press). 




Richard Spellenberg and Sergio R. Rodriguez Tijerina 1 

Department of Biology 


ices, NM 88003, U. 




sionol Mi, 









n/is,»l W ai 


ndl.)J.H Mac-bride 

\l , whayhouh 

s (A. Gray) 

A. Gra; 

/, and M.I en 

lally classified. Mi 


(Benth.) Curran, a 

, n,G, 


as varietie 


laevis van villosa 

<\ ellno.MSpelKn 

ov). M. 

locvis x 


..v) speller 

, bo com 

b. nov), M. iaevis - 

car kicvis and M 


a (Helh 


lin- 51 G J and M.oxyhaphoidc^lu 10.. > 


America. Mi refhi Ji.s o/igurK /ui (Standi J | T M.u bride \(. cvGcj/Goi ,1c. '. A t ,i ,iy) A. Gray, y M. tcnuiloha 
S. Wats, permanecen tal conin sc cla.-ai k aban t rada lonahr.enie. MirGGiA GeGcvn A. Gray. Af 

unicaespecie M laevi.s \ reconouda conm in d.ade < VI laevis' u \illosa (Kellogg) Spellenb. 

\ 1 1 u I | I | I I 1 i G', 

earaeicres. I'l estudio de eslos mapas hie inny i mpr.ri;mtc paia Miliar las decisioncs taxonomicasGe 
of'recen listas de colecciones miportanics Se ciian pm pnmcra vc. nunn a os cromosomaticos de M. 
hicvis var. viUosa (2n = BO,). M. GrviO car in mr,. i 2u - 11 G .. v M <> xvhuphaidcs ;(2n = 30„). 

Mi rabi/is Lis primarily a New World genuscoinprising45 60 species distrib- 
uted from southern Canada to southern South America, with one native to 
southern Asia (Bogle 1974; Heimerl 1934; Le Due 1995). Species have been sus- 
pected of hybridization (Shinncn, 1GVI j. In addition, some are known to be au- 
togamous and even cleistogamou < rudenl973 stamens and style curl tightly 
together in flowers of i in peca ri red lieu i in ol herand land Mi m./imii 

southwestern North America, probably effecting sell pollination as observed 
in Boerhavia (Chaturvedi 1989; Spellenbcrg 2001), species in other Mirabilis 
sections (Cruden 197 3; I lemandez 1990), and several other genera (Spellenberg 
&Delson 1977). Coupling hybridization with autogamy may produce individu- 
o.l I \- raihei nniloi in poj i I n ion bin e,< oeo iphii ll ompl \ \ riation pain i n 
(Stebbins 1957). 

Such complexes provided fertile ground tor the description of" numerous 
entities under taxonomic traditions ol early in the 20 lh century, in which, be- 
cause of locally uniform populations but widespread variation across a geo- 
graphic region, taxonomic decisions mav be subject ivc and [perhaps utilitarian, 
following a philosophy ex pressed by] ew is U 90 0. 1 lere, lor example, more than 
40 synonyms apply to our concept of Mi o o/u/os Mcvi.sgBenth.) Curran and va- 
rieties.. I he laxonomic problems associated with Mi rdbi lis were commented 
upon by Shinners (1951, p. 173) ("Mirabilis is surely one of the most trouble- 
some of Southwestern genera, in nomenclature and taxonomy both.") and by 
Standley (1931a, p. 73) after several decades ol' study in the family ("1 know of 
lew groups of plants \Ncca, lorrubui, Minibilisl in which specific differences 
are so unstable and so ball ling|;| ... no single character seems to be constant.") 
Turner (1993), conversely, in a rather refreshing approach to the taxonomy of 
the genus, noted that if emphasis on vegetative variation were minimized, and 
fruit characteristics were emphasized instead, i he genus in Texas was taxonomi- 
cally tractable. 

Mirabilis was divided tntostx sections by I lennerl (.1934: translated in part 
and reviewed in be \Xw 1995). one of which. Oxxbaphoidcs A. Gray, was char- 
acterized by slightly accrescent involucres and I nuts that are comparatively 
small and unornamenied (big. 1). Heimerl included in it the North American 
species M.oxybaphoidcs ;(A. Gray) A.Gray, M.ialijornica A. Gray (and close al- 
lies), a number of South American species, and one southern Asian species. 

\Uial>ili\o\\ bny/iniGw has presented little taxonomic controversy at the 
species level since its description by Gray l. hS 53) in the genus QmwwcUdhm. It is 
sufficiently distinct from or her species ol Mi ra/>i / the genus is now generally 
construed) that it formed the monotypic genus Mlnmiclla of Rydberg (1902). 
This classification was followed by Stand ley U909, 1918) in several treatments 
of the family, but he was apparently unaware o{ its presence in Mexico, as it 
was not included in his treatment of the family for that nation (Standley 1911). 

The remaining taxa of the section in North America were placed in a new 
genus Hespenui i ub\ Stand ley (1 909). w ho emphasized differences of fruit form, 
shape of the perianth, and number oi flowers in t he involucre. Standley recog- 
nized eight species and several subspecies, em phasizmg shape, color, size of the 
fruit, and vegetative characters such as plant size, leave size and shape, and char- 
acteristics of vestiture. Jepson (1914) treated this as subgenus Hcspcionni 
(Standi.) Jepson, including M. i Mi/nnih n and ;U iYimi i/nbn S. Wats., noting also 


Fig. 1. Variation! in fruits of Nyctaginaceae, section Oxyoap/jo/fH in North America. Fruits are grouped by taxon [Mirabilis 
laevis,M, oligantha,M. oxybaphoides,M. tenuiloba). Those above the line"leavis" refer to varieties within M. laevis. Fruit 
above letter N is 7.5 mm long. Each letter refers to a fruit from a different collection. Collections, fully cited in Appendix 

D f£ llrnb(„>^.;<,t >,i, "-.■.• •i.'.j^.F '•/■ '<,-. - ■. , ■' ■ •>• -...'.. H , v/ « '>■ ,tf .. I '. // ' ,/ 

12332; J, Barneby 18303; K, Spellenberg 12342; L„ Speilenberg 12329; M, fusf wood 785 73; N, IV/^/ns & IW^/ns 75940; 0, 
Gentry & Fox 11731;?, Moron 23808; Q, Johnston & Muller 603; R, Waterfall 12142; S, Columbus 637; T, Correll & Johnston 
245 16; U, Powell & B. L Turner 1708; V, Wiggins & Wiggins 15863; W, Brandegee s.n. 

that M. /an band At - , imnI.M hk.II Jep on wc n < lo ( h related if not the 
same as M. califormca. Standley (1931b), upon completing studies of South 
American NyctaginacMi mm d i h u i lui urn tu m,.dn>di im;mi li'madi 
American genera all i In VI i /hi Ms did m o so and a. I so ho tounm ill in 
an inclusive Miui)>ili id ilu it ion lollm i I >\ most I >i i i inci then 
Of those taxaeaiK ])! u ( d M >\ mm, i dim u nit m d heie M.oligantha 
(Standi.) J.F.Macbride remain .oorl !n> n \l. t mMnhbha pi< ented .i 
few problems, but A 1 /m u > h. been a om col plethora of nann n ono 
mists have attempted to deal with the variation presented by populations in 

the complex. The high pom ol tl ma mcand tings are discussed 

under each of the taxa below. 

Recently Le Due ( 1 993 J c I e sc r 1 1 md MinibilisrusscUiW .e Due from the west coast 
of Mexico, placing the new spec u intln ci i< >n ( Ixybaphoides because ol itssulli u- 
tescent nature, campanulate perianth. and mm. NaginousanthocarpCwhen wet). 
It rests very poorly in this section pnmank because of general habit and ant ho- 

carp morpholog\ lie ingle i in mat i | mailable to us on the paratype 

at NMC generall) u cmblt mtlux irpso loth in \n Minibi lis such 

\ \f m ,iith >• I k mil ii \J i i n i i mi I i 'in ml / i i I i lii 

in Le Due's (1995) plate 11. For the present treatment of section OxyhaphoiJes 
we exclude the spec i . no ngg« tit In moiccomfoi rabl\ in section Mirabilis. 
In this paper ve < vunna dn ^vmhIii \uutioii ind ta\onom\ ot the 
most complex species in the section in North America, Mirabilis laevis, and we 
provide a key, descripi sons. and disi ribuiion maps for the other three species in 
i In cation G\ yhupliimle.sin the I n i ted Stat t md M uco Mirabilis laevis and 
its component taxa hasa i.tortuou ra oni)ini< i torvth r lias resulted in many 
names published at d i. piiln induilii ps.ili I. -I i \\, luguez 1992,) based 
on differing generic, specific, and ml Vaspecific concepts in the group. Generic 
concepts emphasized primarily die importance ol the number of flowers per 
involucre, the shape ol the haul and thedegiet of a nan i ice of the involucre. 
spec i he or \'arietal decisions have primarily eir pfia.sized Iruit shape and sur- 
face patterns, color ol perianth, and rial Lire ol pubescence of foliage and stems. 

For this study more than 3000 herbarium specimens were examined from A, 
UTC (abbreviations from Holmgren et al. 1990 J .From these specimens, 256 from 
the M. laevis complex were selected that had mloianalion about perianth color, 
possessed ripe fruits and at least midstem, and. had adequate data re- 
garding place and dale ol toll uon lh, i |x linen i pr< cut I lis na pin 

i d variation and gci i aphic range ol the taxa. The\ applied data for mor- 
phological characteristics plotted in Figures.' and hind deseri bed in treatments 
of taxa. Taxonomic decisions were made after study of specimens and the in- 
spection ol maps generated by plotting morphological characteristics geo- 
graphically Types ormiciof ie In oir\|> ( loi a hhimh w i i e ntiinnhi 
possible. From this information taoa \vi e delineated thai ,ecmed to um. 
morphological, ec of u'e a nd geographical n ih fhosediat howed consid 
erable intergradation were ivcognized at the varietal level. The order of taxa in 
the treatment is bast don pcrceivcel habit al spec iali at ion and reduction in num- 
ber of fruits as generall}' compared to other Mi rabilis. 

\\\ li iv< epai h I del ul« ddi i i ion of vai a itaon based on Huh ol pi < i 
mens in the Mirabilis laevis com plex from the main taxonomic treatment and 
have included that in Appendix (.Appendix Vonsistsol standard citations of 
representative and/or cited specimens., including those that voucher chromo- 

Le Due (1995) provides a key to the sections of Mirabilis. 

bilis sect. Oxyhaphoides A. Gray in Torrey, hot. Mex. Bound. 17 3. 189a. 
MlioiurUd R\ Jlvig. Bull. lonvv !hu.( lab /OoK7. 1 002: Hes/vmiiM Mam! lev. Contr. 
U. S. Natl. Herb. 12AO0 1000 loi Ah i\i hi bsd\ ydu/s'ieajo (A. Gray) A. Gray. 

[ lerbaceous to suffrutescent or shrubby perennials; root (.of North American 
taxa; others unknown J long, cylindrical, cordlike: sterns erect to decumbent or 
prostrate, densely to sparsely leafy, hemes more or less evenly distributed, basal 
leaves larger, pet 10I. in distal] r\ mallei fiort-petiolate or sessile, margins 
plane. Inflorescences axillary and I rminal in open or eon; ted. lew or repeat 
edly-branched cyni< • involucre: bell shaped lightly accrescent, with 1 or 3 
flowers inserted at base. Perianth broadly funnelform, abruptly flared from 
narrow tube, deeply 5-lobed; stamens 3-5. Fruit ellipsoid or obovoid, base not 
or slightly constricted, apex rounded, truncate, or somewhat nipple-like, sur- 
face with or 5-10 indefinite or prominent lines, often somewhat furrowed, 
smooth or very slightly rugose i nail Labrous, mucilaginous when wetted. 
A poorly understood section of about U » '< ' |vcu f n th America. South 
America, southern Asia. Heimerl (1934) suggested there were about 23 species 
in the section, but considerable redefinition and consolidation of taxa in North 
America has reduced that number. Diversity is greatest in South America 
(Heimerl 1934). 

I. Involucres 3 flowered. I.M.oxybaphoides 

, ' mm l> urn tin li iln tn inn il hi I ii - I I it ill- 

n I n i in. .in i i i • inin i 2. A 

l.MirabilisoxybaphoideslA ( iy)A.< .m in [on ' & Vlex Bound. Bot. 173. 

1859 Out/mm ('. Iiono^ vbaphonU s \ ( ira> Am* i |oui Sei >.. E> J20 L85 S Ulionia 
oxybaphoides(A CrayjKunt c Re\ Ccn PI !?a)l 180! \i lion did oxybaphoides 
(\ t„,n J U'\«lh hell hatm liot < lul.2')(,87 Nit', l\ pe it tin- toot ol mountains 
eastoi I I Paso in ilu h uli . In I m< I U C .1 n>t\p I d mm «! 
CEP nght suli ..1 In. n (H n am ilWiiv'ht Ii Id iimik i / ' ' T mountain, in u 
hi Pino mshiJ [»i imili m - |.l 1! r'S md ' u n oot uul ui tin h id 

ofhighioOvs tl piupU t'i i I I lo I 1 iniK.lon i I | mlebb Hueco Tanks). 
At this time it i ami a be determined from which iw cither of t he two specimens 
on lin ah. t originated Die one on the right is the more nmtim mul repn enta 
tive; a second specimen originally at the Boston Society ol ttui tl Hi tor) trans 
ferred to CI 1! in EH I, very closely resembles the plant on (he right of the type sheet, 
is in a imilai una o! matunm mil i i probabh i oh . mm p. 
Oxybaphus wrightu HemsE, Biol. Centr. Amer. 3:3. 1882. Tver:: NORTH MEXICO: 
h i ih in fountain Wright tin mi' linn n nl n i ill ility 

and collector, without number or date Cray ( hSa d dies U a ie/P 172/ ltd I!), from 

Ctiadalupc Pass in die U luricaluii Mountains; - the colleelion probably seen by 

I Icmsley. II from the present dav tmadalupe Pass, the collection originated in the 

Peloncillo Mountains in New Mexico. 
MirdbiUsowhaphoiili's var. yjabntli! I lemierl, Ainuiaire ( .onserv. ]ard. Bot. Geneve 

y I SO. \M I Ml ; 1 )/ii l -// [ (,)\vKi/>li,ii l /csv.ii-.y,| 1 (/>i 1 (i</ U leaned) standi.. Contr. U.S. Natl. 

Herb. 12-357. 1W Tvns NPW MPXIt a\ I iv 01 \i .cPU .apitan Mts. d A l^ 1000. 

HS.6>E.S.Earie.399uioi otvpis IPs'; [Sotyh-: NMQ). 
Plants usually loosely clump lorming. herhaeeous basally, the stems often 
intertangled and clan i 1 vi mg through oilier vegetation. Stems ascending, spread- 
ing or decumbent, 0.2-1. 2 m long, repeated I v branched, green throughout, pu- 
berulent in lines or ihroughotu. glandtdai or nop the pubescence denser dis- 
tally. Leaves thin or slightly flesh}-; petioles up to 15 em long on basal leaves, 
becoming progressively shorter distal iv. the distal leaves subsessile or on peti- 
oles to 4 mm long; blades oi the basal and midstem leaves broadly deltoid or 
ovate, 1.5-8.0 cm long, 1.0-7.5 cm wide, glabrous or puberulent, and then often 
glandular, the base cordate, the apex usually acuminate or acute, sometimes 
rounded; distal leaves from broadly deltoid to lanceolate. 5-15 mm long, 3-10 
mm wide, the base cordate oi lounded bi//ore.v cmcc loosely and narrowly cy- 
mose or narrowly thyrsoid. Involucres solitary or loosely clustered at the ends 
of branches, or solitary in forks oi branches or axils of leaves, on slender pe- 
duncles up to 17 mm long, glandular-puberulent. widely bowl-shaped in fruit, 
much broader than deep, 5-9 mm long, the 5 bracts united by their margins 
1/3-1/2 their length, the lobes approximately equal, broad I y triangular, 4-6 mm 
long, about as wide at the base, the apices acute. Pei'u nith cam pan ulate, purplish 
pink, pale pink, or occasionally white, sparsely viscid puberulent externally, 
5-9 mm long, about as wade, strongly constricted above ibe indurate base. Fruits 
3 per involucre, olive or dark brown and black molded nrcvenly black, broadly 
obovoid to nearly spherical, ca. 2.5-3.5 mm long, the width ca. 70-90% of the 
length, smooth or very slight!)' rugulose, sometimes faintly marked with 5 shal- 
low grooves tbig. 1). In - )0 M pSyc/lcnbepa, eCSo/eny bSOoP 

Dns/n but muting. ■{).- Southern Nevada, southern i a ah and southern Colo- 
rado, south through Arizona, New Mexico, and western Texas to northern Chi- 
huahua, western Coahuila, and western Nuevo Leon, in open woods, on banks 
in woodland, among brush or boulders, usually where somewhat moist, 1500- 
'aHH.i i ' owcringUiuu »Aus ust Octobei 

The species is readily recognized by the distinctive shape of the leaves. At 
the apex of the petiole the base of the blade is broadly cuneate within the sinus 
of the overall cordate base, the curve at each side oi the base of the blade revers- 
ing in a. sinuate manner before joining I he petiole. I he apex of the blade is usu- 
ally acuminate. Vet \ glandular pubescent plant- and glabraic plants ma\ oc- 
cur m the same population (Spellenber^et al.%81). Plants may be sufficiently 
viscid to "catch little birds" (label data, Vestal &■ \ rstal 50). Leaf shape is consis- 

tent throughout the range except in N'uevo Leon, where leaves on some plants 
are cordate-truncate at the base, rounded at the tip. On these plants the stems 
are little-branched and apparently ascending. 

Plants were used by Native Americans to help heal "broken o\- bent" Lone:-; 

(label data YesialOOt suddddd 

2. Mirabilis laevis (Benth.) Curran, Proc. Calif. Acad. ScL, ser. 2, 1:235. 1888. 

( Ixxlh'phus la, vis Brndi.. lint. Vov. sulphur 44. 1844. Hespcronia hicvis iBenth.j 

scanhl..Conti:U.S.Narl.llrrlU2:k 1 ).L)0n.gii l nM ( . ( /i L l,<>Ji( 1 (evriIVnihjRvdL. iiull. 

lortxv Bot. dub hO :( >87. 1002. Tyit: B,\|A CM IIORN1A: M.u;Jalena Bav 1841. 

Hinds .s.n.(Htil.OTYi'i : K!; photos ol holotvpe NMG). 
Plants few-stemmed and clambering 1 1 i rough other vegetation to inanv 
stemmed and forming clumps as wide oy wider than tal I; stems from the previ- 
ous year often presen I and skeletal wh i te. Sh c ms herbaceous or suffrutescent or 
clearly woody basally 0.1 5-0.8 in long erect or dec urn bent, repeated I y branched 
and appearing more ci lessdicliot >inou I bums, glabrate, puberufent, more 
or less scabrous, or viscid-villous. w hen pubescent, the pubescence denser dta- 
tally hairs spreading or retrorse; intern odes 8 I 1.5 cm long, Fcmrs more or 
less fleshy pubescent like the stem: petiole* 1-22 mm on basal leaves, becom- 
ing progressively shorter distally 0-4 mm long on distal leaves; blades of the 
basal and midstem leaves ovate, deltoid-o\ ate ovate rhombic, or subreiikorm. 
1-4C-5.5) cm long, 0.5-3.5(-5) cm wide, the base cordate, truncate, or broadly 
obtuse, apex acute (occasionally attenuate J. obtuse, or rounded, distal leaves 
lanceolate, lance-ovate, or ovate-rhombic, 5-17(-23) mm long, 3-1.K-26) mm 
wide, the base cordate, truncate, or rounded. Fi/Fururiuxxsuymose or, in west- 
ern races, more or 1< thyrsou iiy pvu iia.l uppn ion oi one ol the pair ol a\t 
Involucres clustered and neadv sessile ai the ends o\ branches, or solitary in 
forks of branches or axils oil ca\( any lunch H2 mm long, campanulate, 
3-7 mm long in flower, enlarging about 1.5 x in fruit, the peduncles elongating 
slightly and deflexed lulu ol uiw.lii. i d d loiu nwoaiallin ilnluitlua 
the tubular portion lightly mux pi I run < L to broadly triangular ot 1 1 lan 
gulai -lanceolate, tlie base 1/ 3 to equal to the height. Perianth widely flared from 
a narrow constriction toptheind i lei hm white with magenta win 

pink, lavender, or magenta, sparingly pubcrulent externally 10-16 mm long, in 
full anthesis usually slightly wider. /u'M!( sd (rareb' 2) per in ohtci i or dad 
brown to almost black, ovoid obo\oid.oi almost spherical, r'omin long, 3-4 
mm wide, glabrous, almost smooth to moderate! v rugose, ol ten faintly mottled 
with darker brow n oi bla< k ni i i u.t 10 1 I i ill I i i inal Im 
becoming mucilaginous when wetted (Fig. 1). 

Distribution (Figs. 2, 3 J. -United States from central California and east- 
ern Oregon southward through soul Invesicm \ hah and central Arizona, south 
to Mexico in west-central Sonora and west cent ral Baja ( ulilorma Sur. 


). Map simplified from Rodriguez (1992). 

nineties based on morphological differ- 
: or less distinct geographic races. For the 
th the variation presented b\ these plants, 
some taking a rather eonsei native \'ie\\ and planing most tonus in an inclusive 
Mirabilis lucvis, others splitting variants as species or in! raspecilic taxa. Even 
on one of the syntypes t,NY ) ol ,\ 1, nihilist n/t/nni u n, the first of the variants to 

? recognize lour mtergradmg 
itury authors have wrestled wi 

| /" 

\ > 

<3 var. LAEVIS 



a V^"'7k 

<6>- D r\ <^A K "\ 

o var. RETRORSA 


• var. VILLOSA 


"i& \ 


involucre lobes 

»► = or > than tube 
► < than tube 

and inflorescence 

< hairs conical or coarse 

- hairs long, often viscid 

y hairs slender, usually short, 

\^ 1 

\ I 


-29° >v \ 


i \ 



\ ^N ' 

N O^^T 

) ^ \ 

-°-f- N 

\\ \, 


vJ \ 

be split from M. laevis, the cpn het "icavis Benth. appears on t he collecting label 
along with "Ovvl.cip/)UNo| ( , /.),/«,?, ms Vahl." Curran (1888), in transferring 
Mirabihs laevis from ().vy/)d/)/in.s noted that plants recently brought from 
Magdalena Bay were "nearly but not quite glabrous the inequality of the in- 
volucral lobes variable and oi ten nor greater than is found in our Californian 

(onus." Wiggins h WW) placed all lorn 
Baja California Sur into M. iacvis, appt 
except lor Miru/u/hubi/nniiu/ van eedwsensis, which he noted to occur from 
SanClemente Island, halilorma, southward on the west side of Baja California 
to the Vizcaino Desert area tea. 27 Nl In that work Wiggins noted the inland 
specimens ol M. Inrv/'sto he "quite viscid-pubcrulcnt to short -villous and often 
have the coarser and almost retrorse scabrous hairs on the upper stems that 
occur m forms of M. californica." 

What was pre\ ion l\ o onsidered Wi <ahi hsi u/i/uj fin niiou com puses M 
laevis var. crassijolia w Inch usualK has an inflorescence with a more or less 
well defined central axis and shorter lateral branches, the entire shape being 
irregularly conical. We arc lenning this inflorescence form "thyreoid." This con- 
trasts to the much more openly and s\ mmetrically locked inflorescences of 
many of the inland populations. We call these "cymose." The distinctions are 


• ' : mm , h< ml.t h lulu fihni puh. ,., in iIih puh.-ie-n 
)t,il)lv viscid noi hMioim' (hut tuns htui stout and received .lit 

i. e usu.ilh," nui 

ith tetrorse hairs; inflorescences broad, cymose. 

i trassifolia 
2c.M.laevis var villosa 

h. Mitab.lh laevisUk-ntli.h m ran var. lacvis. 

Stems glabrous. Leaves glabrous adaxiallv, with a lew short straight hairs 
abaxially; blades of the basal and midstem leaves ovate or deltoid-ovate, 3-4 
cm long, 2-3 cm wide, apex acute: distal leaves lanceolate, lance-ovate, or rhom- 
bic-ovate, 7-14 mm long, 2- ) mm wide Anjloresi enee narrowly thyrse-like. with 
a long main axis and shortei' side branches bearing near their tips involucres 
borne singly or in small clusters, involucres 7 10 mm long, sparsely short vis- 
cid-villous, the bracts united 1/3-1/2 their length, the lobes lanceolate or ovate- 
lanceolate, acute. Perianth magenta I i nit almost spherical. 4h mm long, 4 mm 
wide (Fig. 1). 

Distribution (lag. 3)-Apparenrlv restricted to the vicinity of Magdalena 

Bay, Baja California Sui\ Mexico; habitai not recorded, C>-kkC m. Flowering late 

winter and spring. 

2b. Mirabilis laevis var. crassifolia (Choisy) Spellenb., comb. nov. Oxybaphus 

^(Jjri/d/insValilvar.cTcfssi/o/in.sClM.isvinlX .'. Prodr. MUM Ml. 1.846. TYPO NOYA 
CALIFORNIA: 1833, Douglas es.n. (IIOI.OTYPI ■:: G-DC [microfiche RSA!]). 
Mirabilis californica A. Cray ex Terr in YV I i, kmory. Rep. U.S. Mex. Bound 2(l):l09 173, 
plate 48. 1859. OwTtO'/nes ru/i/ewtnk us (A. C .ray J I look, m Bcnth. & 1 look. I'.. Gen. 

II >4 1880 H.sgmomm Pi/mnm \( lM srandl ( onn I s Natl Herb 12:364 
1000. Twr: CALIFORNIA" San Diego sand lulls. 1850, CO. Pm / v XHOl r< iwnn here 

1 i i I 1 I i I i I ' ( , tesikat Ai i u 

"dry hills, San Diego Gal Horn i a.." ami then indicate*, t he col retorOTarry Thurber." 
(On a sheel I rom the lorrex I lerbannin at \ V are t hiree speennens. Gne, at' the lop, 
collected m l.os \nw I. kv Mr. R u k i ol m Ouokcr eoncern. \r the bottom ol the 

hi , i jn i o) |ioi iiiui M pk m i muni iIm in k hi, I indicating the plants 
were collected by G( Parry on uulliill in in Diego in 18x0. Because the state 
rnent on tin lab. 1 mo nkk in n k. ih. h ibn ml. . nb, d aiu\ ik, p . in n 
at the lower right ol the sheet clearly is the source ol the illustration for plate 48 

i u i\ pi (i i pi u i in ik. i | n \ i I ungtlu nntui il at 

the bottom of tin >heet as the led i p. lOf m Diego. Wood Valley .May 

1852, Thurber 569(2 specimens. Id Ik: BocUr. YY hippie bxpedition, on the Colo- 
rado 1S>3 ,| Kill! NYklwluch u pr. eni i k. , u v,llo . Oclloggi pcllcnberg 
as delimited herein]: Bigeknv. 21 Mar 1954 Kill!). At the tune of publication Gray 

|iu n >n d w h rk i ( ) \ i i i in k mi In k ik i i in m | 

cies, noting that the species is commonk inon oi k pub, emu nidi i l\ 
glabrate." Torrey (U.S. Rep 1 xpl h I i n i -+ 1 >l 1 » i i i iu i i 1 i i 
in iiMiHonulk lo ( )\\ i <i i, i m'. 1 1 a * ihl One in iki pmtolngik ol \1. 

californica, cites Torrey V. listing,, an iccting, l he rnent ol I hese specimens i nay 
I id i m rln niotnlo in i \i ' km 11 m I, s in i kn Bigelow specimens and nidi 
catesthat this new nixon oeeurs ovrkags m oahlor ma on the Colorado." 
ticspcronia calijornua subsp. microphvlla '. Standi.. C.ontr. LPS. Nail. Herb. 12:365. 1009. 
lb or MkkXICO. 1 owt.r (.ho trokxi Y san Martin Island [oil the west coast o\ the 
state of Baja California!. 12 Mar 1867 /banelegee or OtOOUYPF.: DC!). PARA'lYPH: 
I .OWF.R CALIFORNIA: Fnsrnada "o Apr L893, T.S. Brandegee S.n. (UQ). 
Hcs/u-wniac-a/mscJisiNStandl..( mm 1 x Nai I Inrk. 12: M2, OOP. Mirabilisccdwscnsis 
(Standi.) Jepson, Id ( alii 450. Ibid \l . ek/mi ink e sir... cd ms, xi.o.o. grand In Mae In 
Con tr. Gray Herb. 56:24. LOS M. Gevirear , r, dose /ion Srandl.) Mum r Man. S.uii h. 
Calif. 151. 1935. TYPE: MkkXICO. B,\p\ C.Y1 lPORXlA: Cedros island. 3 Apr 1867. 45. 
Bmndcocc s.n. Oioi.otypp: IkCk. Paratypps; c AklPORNlA: San Clemente lcf, Oct. 
1602 OmskHOHSk i alilornia San ( lenient. Id Max log; PO xP 
Hcspcronia he i merit i Standi.. Contr H.S. Natl. 1 lerkr I k 1 12. 161 1. Mirabilis heimerlii 
gstandlego Macbnde, Contr. Gray Herb. 56:24. 1918. TYPE: BAJA CALIFORNIA: 
Guadalupe Island, S end ol island. 5 Mar 1886 /:. DdmcrSS'PO \o\ OTYPlr US!: isotype 

Mirabi lis laevis ear cordifohu Hun kle. Bull. SO . alii. Acad.! Sci. 40:1 08. 1941. TYPE: CALI- 
FORNIA: San Clemente Island, CluncttiCanyon. ? Apr 19 H Dunkle 7234 (HOLO- 

TYPE: RSA #3506851; isoiYPP RSA ^4646766. Both RSA have been 
transferred from I.AM;ol the two the holotvpe lias a hand written label with the 

Stems often glabrous basal! v, viscid-pubescent or more or less scabrous distally. 
Leaves puberulent, viscid-villous, or more or less scabrous, sometimes becom- 
ing glabrate with age (or occasionally glabrous); blades of the basal and midstem 

leaves ovate rhombic, subreniiorm, or deltoid-ovate, 1--IA cm long, 0.4-3.5 cm 
wide, the apex obtuse or acute, occasionally rounded: distal leaves lanceolate, 
lance-ovate, or ovate rhombic, '5 14 mm long,, 2-7 mm wide. Inflorescence often 
rather thyrse-li ke after t he first lew die hot onions branches, the branches shorr. 
the involucres in clusters along, a main axis Involutes a-4 mm long, densely 
>hoi [ \ im id-\ illon oi onu huh ,hghl \\ i abi i(! ilu hi hi unit <! [ ', ' Ohui 
length, the lobes ovate or ovate -oblong, obtuse or acute. Perianth pink, lavender, 
magenta, occasionally white C ah ovoid, >- c > mm long, 2.7 3.7 mm wide, dark 
to pale gray-brown and mottled with dank gray-brown, tan. or red-brown, some- 
times Lunilv and ii regular! \ pak striped U ig \) 

I h 'si i i hut ion U ugs. 2, 3) —West -central California south along the coast, on 
the Channel Islands, and in the Coast Ranges to the Viscaino Desert, Baja Califor- 
nia Sur and the axu tal islands. Mexico: coastal bin 1 1 road banks, coastal scrub, 
grasslands, cha pa 1 1, d oak woodland oltcnonroi cy outcrops, 0-1.830 m. Flow- 
ering most of the year, most vigorously in spring. 

A particularly difficult area with regard tovariation is around the southern 
in I oi tin in Nevada in < tliio m t.wlt ( hrc< ol dii n < u, an m cont u i 
Howell . 181 79, from the 1 .ake Isabella region in Kern I o.. ill ust rates very well the 
problematic classil ication oi some specimens It was lust left unidentified in 
Mi rabihs, then sometime later placed in an inclusive M. laevis; 9 years later was 
identified as M. rcl rorsa: 21 years later as ,\f bioelovPi van. hi^elovii, and shortly 
later placed in a van bit \ v era m/uIju I In o ' ncn com bines the spreading 
leaves and (light) villous pubescence ol win. biyelovir the pointed leaves and 
rather long in vol uca e lobes oi var. < rassijolnr and some retrorse hairs similar to 
van n'lmisu bkiwer color was not given In t he collect oi an ml is not evident from 
the specimen, but as judged from Spellenbergs collections from this area, flowers 
were probably white. Nearby, from the entrance to Kei n River Canyon, comes 
Howell 38142, a late season collection showing thyrsoid inflorescences of var. 
crassijolia, and fairly pointed leaves, but in other respects is the var. villosa; a 
mnl to latei oil lion ' Plow li CC, »') from rh nm i i note cab no l : oi 
plantslnoin the lowei reaches ol the Kern Riwr Canyon. Spellenberg's observa- 
tions note 1 lowers only pink to nose, vet il these plants were collected eastward 
thev easib would be placed m more consistently white-flowered var. villosa. 
"bvrssr/mmi 83x4, from the same area, is more or less villous and has blunt leaves, 
in these respects similar to the van. villosa. but it has a more or less accrescent 
involucre with proportionately longer lobes, more rem i n iscent of var. crassifolia. 
Also seemingly intergradient to var. villosa is Bedell 74-5 And Twisselmann 198, 
both from the north end oi the lemblor Range in western Kern Co.; they have 

thick flesh)' leaves that are blunt, short involucre lobes, but thyrsoid inflores- 
ceno Tw i elm urn no i - ill, it ! lo\ ei ire pm pie 

XUfahili dacvis vouuuu.ssi/obinuav al ointergrade with M.olivdnthain Baja 
' aliln ma ui ( Moo K J ./ ; b 

Much has been made of features ol pubescence over the taxonomic history 
of this group. Even within var. cru.ssi/obu there is considerable variation. Near 
the coast, and particularly on the islands of non h western Mexico, hairs are stout 
and conical, distinguishing Hesperonia ccd rose n.os a ncl subsequent synonyms. 
This pubescence type is thoroughly intergrad lent to I iner but still conical-based 
hairscommon within the var. crassijoiui. bruit characteristics, such as those used 
to distinguish H. hamcrln, also an island population, seem to be completely 
inconsistent from population to population. Other specimen-based discussion 
focuses individually on characteristics that have been used to distinguish species 
in this complex and is found in Appendix 1. 

The label on a specimen collected in Baja California I \ ioran 12832 I notes 
the indigenous name and use "Yerba del Empacho.-bueno para el estomacho." 
The vernacular name is repeated on Moran 23821 from Baja California Sur. 
2c. Mirabilis laevis var. villosa (Kellogg) Spellenb., comb. now. MirabiHscaUfornic a 
var. villosa Kellogg, Proc. Calif. Acad. Sci. 3:10. 1863. Typo CALIFORNIA. MONO 
Co Calif llw\ 182 10 m from Nevada borcK, U-mI ,(,at. . unmiul la I Wall < i 
One, s r Y[ t i,YoN0YY lijui IW(hi koii hoc! ivmi i i MM( 'no. i,p 
[!|: BYL.CAs, I: K. MP.YC, MO, NY, RM, RSA, UC.USi. Kellogg (18o 5j provides hnel 
but clear d< eription >l i plant that match the classic concept oi Mi mhilis 
hi e/m nN < pt that li nor in * , nhn n o 1 n, ( ml , c i miu.m ' i it hi 
than ii < i 1 ! in tin t on to< n \> i on the in