i ' i f A n m lérnati onal ournal to expedite ip/a n t s ys tem a t c; j , ' " -an d eco logiga ] pup/i ca fi on

' ‘ r C ONTENTS

i i LOPEZ FERRARI, New combinations i n Mexican

TURNER, Rebuttal

0 0 0 0 0 0 0 0 0 0 0

14

Inde u hors Ph tolo ia l e 1 x to a t y g vo um 8 a g “

Di s si ion of anuscri ts volu mes 80 e po t m p ; n w fi

. t l m a a Phy o ogia Me oiis vail ble . w PHY TOLOGIA ( IS SN pu b li sh ed. monthly witli two volumes per year : b y ‘ Mi el J 1 W l s a e cha 85 s rid Dri é un s i lle X 734 . S econd C ass o t ge v ; H t v , T 7 0 p t g e " aid at H un sv i l e - I 1996 ' H T ‘ I A nnual d mes i c i n di i du a l p t l ; X b P Y OLQG A. o t v j ' t i n 1 2 subs cripti on ( 12 i ssuesz z A nnual domestic . 1 nst1 tuti onal sub script o ( ' “ 44 i an / or a1rm ai l e s r s e : c ur r en $ Fore n d t ext . i e co a s g pos ag a S ngl py l t . " ' ' ' 1 ssue and. back i ssues V olum eJ 72 to present : $400; b ack i ss b es 1 (previ ous ; to v o l ume _ ' " ‘ 72 ) add $ 75 per copy postage and handlin US 5 0 er m m fo reignJ) J B ack “ g p ' ui - le e ssue sales by volume: $ 1 7 00 per Uvol ume . 42 71 . (11 01 all av ai lable as comp t " volu me s) ; $ 2 1 00 per v olume 72 mresgnt add $ 3 00 per volume po stage US p e r ' ' 1 - - r olum e forei n . . POSTMA STER: S end d ress an es to Ph olo i e l SS We st i d e v g ) a d ch g yt g , g ‘ r D ri v e Huntsvi lle; T X 77340 - P hytolo gic (Febru ary 199 7) 72 .

HIS TO RICAL NOTES ON LOUIS IANA P RAIRIES : CHANGES IN P RA I RIE FLORA IN HALF A CENTURY

MacRoberts MacRoberts M .H . B . R.

ar h 740 lum i a S hr rt ouisi ana 7 1 104 U S A . Bog Rese c , Co b , evepo , L

H r ari um Mus um i S i n s uisiana S a ni rsi i n Shr r e b , e of L fe c e ce , Lo t te U ve ty evepo t,

hr ort ui siana 71 1 15 U S A . S evep , Lo

A BSTRA C T

r n r Unpubli shed field notes and herbari um specimens of C A . B ow we e used to reconstruct the 1939 flori stics of the Keiffer prairi es in central Loui si ana These histori cal data were compared with current species

m i i n r m the same rairi s. Al h u h the rai ri es are smaller da co pos t o f o p e t o g p to y , x A we found that they have not lost species reported si decades ago . compari son of the species composition of the Keiffer prairi es wi th rnidwestem tallgrass prairi es indi cates that they are relati vely hi gh quality and have a ri ch n array of co servative prairi e species.

KEY R S : Prairi oui si ana flori sti cs i sat hi a i nal F r s WO D e , L , , K c e N t o o e t ,

INTRODUC TION

In rma i n on lan uni l ain Thi s i s fo t o p t comm ties spanni ng decades i s not easi y obt ed . particularly the case with prai ri es (Collins A dams Their rapid demise has n d um n d and sin so li tl i s l da an ir in r mnan i s hi hl riz ed bee oc e te , ce t e eft to y , y v g e t g y p (Kucera 1992 ; Whi tney This is especially tru e in the S outheast because the small is la d rai ri s hi h r n r mm n r ui l d str or o te p e , w c we e eve co o , we e q ck y e oyed degraded by early settlers (DeSelm Murdock

’ In the la 1930 s lai r A . r n r s r f an at ouisiana te , C B ow ( 1903 P ofe so o Bot y L S a ni rsi a s ci al i n r t te U ve ty , took pe te est i n Louisiana prairi es and made numerous tri ps s u h m r n 1941a 1941b A m n the m an rairi s r n isi to t dy t e (B ow , , o g y p e B ow v ted were the Keiffer Prairies on the Winn Ranger Distri ct of the Ki satchi e National Forest 66 P H Y T O L O G I A February 1997 volume -72

Th s are a r u of a u 45 na ural ni n s surr un in central Lo ui siana. e e g o p bo t t ope g o ded by Martin S mi h an sh rt l a ine/ oak/hickor r s Smi h et a l. 1989 o e f p y fo e t ( t , ; t L d ’ survey plats from the 1830 s and 1940 aeri al photographs show that these Openi ngs were larger than they are today .

' i neerin r i n r s i n rairi s aned and the i f r Prai ri s A fter Brown s p o g wo k, te e t p e w Ke f e e were vi rtually forgotten until their rediscovery and subsequent surveys over a half All n MacRoberts MacRoberts 1 ur r Smi h et al. 1989 1993 996a cent y late ( t , ; e ; , the las e a s u i s ha e n un rta n on h the rerrman l 996h) . During t d c de, t d e ve b e de ke bot t coastal prai ries and on the isolated prairi es scattered through north and central

a Smith et a l. 1989 Smeins et al . 1992 All n 1993 All n V idri n 1989 Loui sian ( , ; , ; e ; e e ;

MacRoberts MacRoberts l 995 a, l 995b, l 996a, l 996b) .

l alm s no rmal re ord hi s rai ri r u lishi n nl hr B rown eft o t fo c of p e wo k , p b g o y t ee r n l 941 a 1941b He di d h r l a a su s an i al i n rmal abstracts (B ow , , , oweve , e ve b t t fo ld n es and lle i ns hi h eli r id a d in i a i n of record i n hi s fie ot co ct o , w c we b eve p ov e goo d c t o n histori cal compositi o .

’ l n s and h r ari um s e im ns and urr n flori stic da a Using B rown s fie d ote e b p c e , c e t t that

ind nd n l er hal a e n ur la er Smi h et al . 1989 All n 1 were compiled epe e t y ov f c t y t ( t , ; e 993; MacRoberts MacRoberts 1996a l 996b we h d rmin ha if an , ) , ope to ete e w t, y , urred in a flori stic changes have occ five dec des.

’ We ex tracted all flori stic inform ation for the Keiffer Prairi es from Brown s field “ ” n and a summar h s n s la el d Prairi e Re ords r n n d otebooks y of t e e ote book b e c (B ow . . ) s r d at the LSU h r arium . On A ril 8 une 2 0 Se . 9 1 1 and 30 1939 r n to e e b p , J , pt , , , , B ow

i si d the i f r Prairi s and ll d and li s d lan s r m s ral h m . H e v te Ke f e e , co ecte te p t f o eve of t e “ ” als mad a i si the i f r Prai ri A r a on er 2 5 1941 but nl c lle ed o e v t to Ke f e e e Octob , , o y o ct

i h s i s all r m the surr undin lands . This isi w as r a l sh r e g t pec e , f o o g wood v t p ob b y o t in e als i h r ar as ha i it a rairi s ce h o v sited ot e e t t day . He did not cons der p e visit since it “ ” n r n a ar n l dis ntinu d hi s i si s the is ot li sted in the Prairi e Records. B ow pp e t y co e v t to

Keiffer Prairies at thi s poi nt .

hil r n i s the l a i n n 6 l N W h i o e rairi Sec . 2 T I R5 e s m l W e B ow g ve oc t o of p e ( , ) , p y “ ” designated others by such descripti ons as T win Prai rie (now called Carpenter Road '

Prairi S . : ec 1 3 T 1 1N R5 W and Small la rairi on sand um r ad. All e , ) b ck p e [ ] d p o these openi ngs appear to have been withi n a short di stance of each other and are part of ha w h an w t as t en d still rs a group of closely associ ated prairie remn ants .

sin urr n lis s the i r ra Smi h et a l. 1989 All n 1993 MacRoberts U g c e t t of Ke ffe flo ( t , ; e ; ’ MacRoberts l 996b and r n s lis s s ar h the LSU h r ari um for his ) B ow t , we e c ed e b s im ns. Th r r urs the n rm al r l ms: s im ns on l an pec e e e we e , of co e , o p ob e pec e o , ’ mi s la l s n r n r or d s r d a t r r n s d a h man of his lan p ced , o t , eve e te ed , e t oye ( f e B ow e t , y p t ll i ns r und i n a sh hi n hi h u i n a e m s n co ect o we e fo ed be d s o se d co po ed co diti on) . M acRoberts MacRobert s: Chan ges in Louisi ana prairi es 67

limi ati ns a m l r ns ru ha r n un and 2 Given these t o , we tte pt to ) eco t ct w t B ow fo d ) i compare it with what currently grows in the Ke ffer Prairies.

RES ULTS

r n The di r nc n Table 1 lists the plants collected or listed by B ow . ffe e e betwee hi s li st and current lists produced fifty years later can be compared by simply j ux taposi ng MacRoberts MacRoberts l 6 89 A ll n 1993 99 b . h m Smi h et a l. 19 t e ( t , ; e ; )

T a l i r Prai ri as ular lan s i n 1939 . b e 1 . Ke ffe e v c p t

' i L al n S is . ex R s A GA VA CEAE Manfi eda virgi ca ( .) b o e

A NA CA RDIA CEAE Rhus copa llina L.

’ r n i ma u A PIA CEAE E y gium yuccrfol um Michx P olyta e na tta ll DC .

x id A Q UIFOLIA CEAE Ile dec ua Walt .

A S EPIA A EA E A s le ias vi ridi o ra Raf . A . i ridis alt A . erti illa ta CL D C c p fl , v W , v c L .

A STERA EAE Ambrosia silosta ch a Aster oolentan iensis Ridd ll Aster C p y DC g e , seri ceus V nt oreo sis la nceola ta L. Echinacea a llida u . u . Eri eron e , C p , p (N tt ) N tt , g tri Eu a t ri um em r s osus Muhl . ex ill . a illa dia aesti alis al o s iserra tum . t. g W d , p DC , G v (W ) H ni m . le u autumnale L . H elian t n a li R . H e hus a us o us L . H elia nthus hi rsutus ock , , g f , ' Lia ' Raf Iva annua L . ms cnosta ch a Mi ua a . hx . L. s n ulos Mi hx . Rudbe kia , , py y c , q c , c ' h l hi um la inia u i i rta L . Si t m L. ol da o ri ida n ia ur a S L. Vem o miss rca R f. , p c , g g ,

RA I A EAE Onosmodium e a rie v ar . his idissimum a BO G N C b j nse A . DC . p (M ck )

L . Turn r B . e

R A EAE Ca m us drum M e r mo ndu C . M . o ida . CO N C y . , C. fl L

PRES SA EA E J un i erus ir in CU C p v g ia na L.

S TA EA E us uta CU CU C C c cuspida ta Engelm .

E E A EA E Dios ros vi r inia n B N C py g a L .

FA A EA E Astra a l i us rass a r us u . Ba tisia alba L . nt Dalea andida B C g c c p N tt , p ( ) Ve , c

ill . Dalea ur urea nt Desma nthus illinoensis Mi hx acM W d , p p Ve , ( c ) M . ex B . L .

R i ns . F rn leditsia riacan ho s Mi sa tri il L . losa T rr mo s A . ra ob e , G t t , g o ey G y , ' Ne tum a Iu tea a n p (Le ve worth) Benth .

E TIA N A EAE Sa ba tia a n la r is L u Pursh . a m . S estri s N ut G N C g ( ) , c p t 68 P H Y T O L O G I A February 1997 volume - 72

A Li uidamba r t r i u W CEAE q s y acfl a L.

AMIA Mona rda istulosa L . P runella ul a r is L . P na n the m um L CEAE f , v g , yc tenur lium S hra Sa l ia a urea Mi hx . ex Lam Sa l ia I rata L . S utellaria fo c d v z c v y , c i pa rvula M chx .

A llium anadense L . H ox is hirsuta L . ill Nothos ordum LILIA CEAE c , yp ( ) Cov e , c

ri . bi va lve (L. ) B tt

LIN A CEAE Lin um sulca ram Riddell

r v A . ra MA LV A CEAE Ca llirhoe papave (Ca .) G y

a ura lon i ora S a h. Oeno thera s e iosa u . ONA GRA CEAE G g fl p c , p c N tt

i ta e PINA CEAE P inus ech na ta P . Mill P . da L.

P A EAE A ndro o on erarda V itrnan Andro o on lomera tus alt. O C p g g , p g g (W ) ’

i an tha Mi hx A nstida ur uras ens P i r . Era rostis s e ta bilis Pursh A ri stida ol g c p p c o , g p c ( )

S u ani um ex ile a t . S ri n . P as a lum oridanum Mi hx P as alum te d P c fl (G t ) c b , p fl c p ' hi a h ri um o a ri um Mi hx ash eta e setaceum hx . S s S na nicula ta Mi c , c z c y c p ( c ) N , g

Beauv Sor ha strum nuta ns L. ash Sor ha strum un eus B eauv . un h. g ( ) N , g j c ( ) K t

hl x i POLEMONIA CEAE P o p losa L.

RAN A EAE Del hinium a rolin ia num alt Ranun ulus as icula ris UNCUL C p c W , c f c uhl l M . ex Bige ow

'

M A EA E B erchemia s andens ll . h ean othus amen can RHA Hi us . N C c ( ) K Koc , C L

R t OSA CEAE Cra aegus spp.

R IA EAE Diodia teres alt H ous tonia ur urea . var . ca l cosa A . ra UB C W , p p L y G y

SAP TA EAE B umelia lanu sa Mi O C gino ( chx ) Pers .

' S R PHU A RIA EA E A a lini li o h lla P nn ll Buchnera n s o ame cana L . C O L C g g p y e e , , ‘

P edicula n s anadensis L. P ens temon a ustra lis Small c ,

MA EAE lm UL C U us a la ta Michx .

VER E A A E Glandula ria E anadens L . erbena ut B N C c is ( ) Nutt V x ha Lehm .

VITA EAE A m elo sis a rborea C p p (L .) Koehne MacRoberts MacRoberts: Changes in Louisiana prairi es 69

MacRoberts MacRoberts ( l 996b) li st 137 species for two Keiffer Prai ries. ’

r n r rd d 86 s i s 80 of hi h are on the MacRoberts li s . B ow eco e pec e , w c t

B rown lists or has vouchers for six species not listed by the MacRoberts : As le ias erticilla ta Astra alus rassicar us Oenothera s e iosa P an icum ex ile c p v , g c p , p c , fl ,

Ran unculus ascicula ris an d Verbena x u tha . F ur h s are r rt S mi h et f , o of t e e epo ed by t , l r n The r mai ni n s i s Ra nunc ulus as icula ris an d a . ( 1989) o A lle two e g pec e , f c

Verbena x utha are id l s a r d i n o uisiana . , w e y c tte e L

DIS CUSSION

A lth ough the sample i s far from ideal (we can say nothing about relative abundance of s i s l arl duri n the as 60 ars the s i s lis of the i r Prairi s has pec e ) , c e y g p t ye pec e t Ke ffe e ’

n li s n ai ns no s i s missin a . Thi s i s an in r s i n not declined. B row s t co t pec e g tod y te e t g fi nding since it appears to be generally accepted that a process of degeneration has “ ” been occurri ng over a wide range of island type communities (Hoehne er ai nl rairi d n ra i n urr d for m s uisiana rai ri s hi h ha C t y , p e ege e t o occ e o t Lo p e , w c ve as d xis al h r and h r s ies n e urri n in uisiana for xam l ce e to e t toget e w e e pec o c occ g Lo , e p e Eus toma randi orum Rat Shi nn rs ha e not en und for alm s 2 00 ars g fl ( ) e , v be fo o t ye

i a s ru i n aus d his l ss. A ri al h MacRoberts et al . H a d ra hs ( , b t t e t ct o c e t o e p otog p of the Keiffer Prairi es show that they have shrunk considerably due to woody ’ i n asi n Man r raz d and r raz ed in B r n s im and a r and s m v o . y we e g e ( ove g ) ow t e fte , o e

ha en ul i a d Smi h et a l. ve be c t v te ( t ,

H r h r is the u s i n the addi i nal 40 50 s ies not r r d oweve , t e e q e t o of t o to pec epo te by r are th acR be B own (comp e list in Table 1 with that of MacRoberts M o rts l 996h) . ’ H as the flori sti c divers ity i ncreased over the past fifty years or w as B rown s sampli ng incomplete"

r n ma nl i rr u lar i si s the rairi s: n a h in A ri l and un and B ow de o y eg v t to p e o ce e c p J e ,

- hr im s i n S m r. H e was not a dri n ll r or n a r and no id n t ee t e epte be ve co ecto ote t ke , ev e ce xi s s sh in he a the n ra : he e t ow g ttempted a total fioristic list of the prai ries. Q uite co t ry n ll t nl a ofte co ec ed o y few species and otherwise mere ly listed plan ts he encountered . H e paid particular attenti on to trees and shrubs (thi s was one of hi s specialties) and often listed woody vegetation adj acent to the prai ri es; he typically identi fied many h r a us s i nl Man n ri s remai n inc m l e b ceo pec es o y to genus. y of the notebook e t e o p ete and i nl a lan num r asi nall r n i s sh r g ve o y b k be or genus name . Occ o y B ow g ve o t “ ” s ri i ns su h as small hi fl rs but aus he did not r r his de c pt o c w te owe , bec e ewo k n s a r d rmini n s i s the n s al n do not r i a r lia l otebook fte ete g pec e , otebook o e p ov de e b e ui the id n i a i n or l a i n H e a ar ntl a i e g de to e t fic t o oc t o of herbari um specimens. pp e y vo d d the ra a al h u h he un a h Cype ce e , t o g fo d few of t e rare gras ses.

While it i s an intrigui ng possibility th at spec ies diversity in the prairies has ’ i n r as s1nce r n s im hi s i s un r una l not sta l i h the da a ha c e ed B ow t e , t fo t te y te b e w t t we ve a ail a l v b e .

H igh- quality prairies are identi fiable by their diversity and ri chness i n conservative fidel or n ar fidel s i s the a s n of x ti s e i s and la of r r w h of ( e ) pec e , b e ce e o c p c e , ck ove g o t 70 P H Y T O L O G I A February 1997 volume -72

brush and trees (Packard Ross While no one has developed a rati ng system f r ies for the s u h rn ni d S a es su h do exi s for the midwestem o prairi e spec o t e U te t t , c t tallgrass prairies (Masters 1997; Packard Ross 1997; Ladd T aki ng in to account such distri butional differences as would be ex pected between central Lo uisi ana and uri or Illi n is it is ssi l to es imat the rall rairi uali for the Misso o , po b e t e ove p e q ty s m ari n heir s ies i h h s un ls h r rem m ri n Keiffer Prairie by co p g t pec w t t o e fo d e ew e e , e be g , urs ha hi s e m ari s n is hi hl su i and n di r n of co e , t t t typ of co p o g y bject ve ope to ffe e t

i nterpretati ons.

K i r rai ries MacRoberts MacRoberts l In 1996 we surveyed two e ffe p ( 996h) . ha and n ain d 100 and 12 4 i s r s i Th and l . ey measured co t e Spec e , e pect ve y m nsid rin the iz of the rai ri These are relatively high nu bers co e g s e p es. Very few

h as a rex microdo nta T rr H . H li tr x i . Man su e o o ium Species were e ot c y , c C o ey ook , p hes ma ni am mm S hevi ak u . T rre S iran t c o and H ous t nia tenellum (N tt ) o y , p g p , o

ur urea var . ca l cosa are fidel or n arl so to rai ri s. i f r Prairi rass s su h p p y , e y p e Ke f e e g e c as S hi a h rium s o a ri um Koeleria macrantha A ndro o on era rdii c z c y c p , Ledeb p g g ,

Sor trum n utans and P an i um ir a tum . are als hara risti of midwestem ghas , c v g L o c cte c rar p rres.

’ A comparison of our plant list with Ladd s ( 1997) for midwestem tallgrass prairies Thi s 1s i res i but an und r s im a shows a 6396 similarity rn species . mp s ve e e t te of the true similari in man e i es un in the ei f r Prairi s su h as r ty s ce y Sp c fo d K f e e , c Spo obolus

v n Mimo sa stri illosa T rre A . ra and N unc B eau . u h. ia j eus ( ) K t , g o y G y , eptun Iutea

Le a en rth en h . are l sel rela ed s u hern n en rs of m r n r h rn ( v wo ) B t , c o y t o t co g e o e o t e

s e i s. Furth r h n nl the hi h s rankin nser a i s i s are m ar d it p c e e , w e o y g e t g co v t ve pec e co p e , is clear that the Keiffer Prairies have basically the same species as the midwestem

tallgrass prairi es .

These results are encouraging and give us confidence that the Keiffer Prairi e r mnan are nl ai ni n h h i Tha the i r e ts not o y maint g t emselves but are hig qual ty . t Ke ffe Prairies may be more diverse (i n better condition fiori stically) today than they were in ’ the la 1930 s is not d m ns ra l r m hese s udi s but the da a do not x lude hi s te e o t b e f o t t e , t e c t

ssi ili hi h is an in ri uin one. In 2 050 s m n r risin ani s can s po b ty , w c t g g o e e te p g bot t te t thi s r si i n sin da do ha a ura fl ris i li s s n a l rare s i s p opo t o ce to y we ve cc te o t c t , ot b y of pec e , for all h m MacRoberts MacR of t e ( oberts l 996a) .

A CKNOWLEDGMENTS

Mar Ma field ura r of the LS U H r arium was ins rum n al in ai ni n k y , C to e b , t e t obt g ’ r n s n s and r rds and in r ar i n our h r ari um r Finan ial B ow otebook eco , fo w d g e b wo k . c su r w as r i in art hall n s - Shar A r m n s i h the ppo t p ov ded , p , by C e ge Co t e g ee e t w t isa hi a i al n F r s . P ar of the x ns s for h r arium ar hi al and l K tc e N t o o e t t e pe e e b , c v , fie d “ r r r i a ran S ur of Keiffer/Packton LA Prai ri s Pr wo k we e p ov ded by g t ( vey ( ) e , oject ’ Number 97 073 - 030) from National Fish and Wildlife Foundation s Wildli fe and Habitat Management Ini tiative and the Native Plant Conservation Ini tiative to the isa hi a i nal F r s and nn S a ildli i l is inn Ran r is ri K tc e N t o o e t Ly t cey , W fe B o og t , W ge D t ct,

i sa hi a i nal F r s . nn S a Gu s m a i M r Phil H a and K tc e N t o o e t Ly t cey , y Ne o , D v d oo e, y tt , R r E ans mm n on arli r rsi n hi a r obe t v co e ted e e ve o s of t s p pe . MacRoberts MacRoberts : Changes in Loui sian a prairi es 7 1

LITERA TURE CITED

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- 3 . S u ssi n in rasslands: i L D . E . A dams. 198 Th r ars of lli ns S . . Co , cce o g ty two ye

- a r V e etati o 1 . change in a central Oklahoma tallgr ss p airi e . g 90 - mm uni - Mur . 1 3 . rass d mina d i s. . 9 P s lm H . R. 9 . 87 141 De e , N dock G o te co t e p .

A . C . Echtem acht B iodi r H . Mar in S . G. c ve sit o t In: W . he t , Boy e , y f

outhea tern United S ta tes . hn il S ns r r . S s Jo W ey o , New Yo k , New Yo k i l 8 1 . The oundla er ati n r s s an s in an H ehn L . M . 19 r ur an o e , g y veget o of fo e t d b

- M . S h r an ma ri x . P . 4 54 In : R .L . ur ess D . ar e suburb t p 1 . B g p Fo est

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S rin r V rla r r . p ge e g , New Yo k , New Yo k

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Pa ar . F u l nd S . C . M tel The Ta l rass Restora tion H a book or P ra i ri es ck d g f , d W d Sa a nna s a n oo la nds . Island Pr ss l ali rnia. v , e , Cove o , C fo

MacRoberts B . R. M . a i H . M cRoberts . 1 6a . inn s ri rai ri r rt . , 99 W D t ct p e epo n u lish d r r i sa hi ati nal F r s Pi n ill oui si ana U p b e epo t , K tc e N o o e t , ev e , L

MacRo B . R . M . berts H . MacR berts . l 6 The flori stics f al u o 99 b . o are s , two c c o i ri s on the i sa - ra hi a i nal F r s oui i an a. Ph tolo i a 4 s 3 . p e K tc e N t o o e t, L y g MacR D T MacR rt . a oberts , . . , B .R. obe s, M H . M cRoberts . 1997 . A flori stic and l i al in r r a i n th m an d r x di i eco og c te p et t o of e Free an Custi s Red Rive e pe t on of 1 806 .

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MacRoberts M . H . R . ar ra f ar u B . R . Mac ul obert s 1995 a . as o al e s , V c flo two c c o i ri r mnan n ra ts o the Ki sa hi a i nal F r s ui si ana . Ph tolo ia p e e tc e N t o o e t , Lo y g 2 7

MacRoberts M . H . B . R . a e r h as ular lan M cRob rts. 19 5 b . , 9 Notewo t y v c p t ll i ns on the a - lo i a . i s hi a i nal F r s uisi ana . Ph to 3 13 co ect o K tc e N t o o e t, Lo y g Mar in L . . . M . Smi h . ur d i n f e na ural lan 19 1 . A s an s ri o th t , D L t 9 vey de c pt o t p t mmuni i f th s o e i sa hi a i nal F r s i nn and i sa hi i s ri s . co t e K tc e N t o o e t , W K tc e D t ct li sh r n u rt . ui a n U p b ed epo Lo siana Department of Wildli fe and Fi sheries . B to R i u u si ana . o ge , Lo 72 P H Y T O L O G I A February 1997 volume - 72

- Mas r L A . 1997. M ni rin i n a 2 7 3 1 . In : S a ar F . P . . te , o to g veget t o p 9 0 P ck d C . . Mutel The Tall rass Restora tion H andbook or P ra iri es Sa van na s and g f , ,

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Els i r r r . V ol. 8a. ev e , New Yo k , New Yo k

m r . P Mar i n G D e r . 1 M . N . M. il R . . . s 989 . i r al S mi h L. ar us t , , G o e , t , L te Ke ffe c c eo rest m l x : a r s ar h r or and r liminar man a prairie/fo co p e e e c ep t p e y gement plan . n u lish r rt uisi an a artm n of il li and Fi sh ri s a n U p b ed epo , Lo Dep e t W d fe e e , B to

R u ui siana . o ge, Lo 4 a l i i 1 . From Coas W lderness to Fru ted P la in . am hi n G.G. 99 ri ni . W t ey , t C b dge U v

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NO ME NCLATURA L NOTES ON NO RTH A ME RICA N GRA S S ES

l R s ar h en re A ri ul ur an d A ri - F d ana a Eastern Cereal and Oilseed e e c C t , g c t e g oo C d , a a n ari A A A KIA OC6 Ott w , O t o, C N D

2 ni R al Bri i sh lum ia Mus um PO . B ox 98 15 Stn. Pr i n i al Botany U t , oy t Co b e , , ov c rnmen V i ri a ri ish lum ia A ADA V 8W 9W2 Gove t, cto , B t Co b , C N

AB STRA CT

The source and nomenclatural type of the name A rctophila trichopoda - v inet rum Holm are discussed; a re lectotypificati on for Danthonia spica ta ar. p o

Piper i s proposed; the name Festuca trachyphylla Hack. ex Dru ce i s a nomen lectotypes are provided for five Festuca names; five new combinations

i n Festuca are proposed .

KEY R S : A rcto hila Danthonia s icata var . inetorum Festu a WO D p , p p , c , l e P a a ectoty p , o ce e

In the course of prepari ng flori stic treatments of vari ous North Ameri can grasses a num r of ax n mi n m n la ural and t ification issu s r n d. These are be t o o c , o e c t , yp e we e ote is u n al h H arium a r n ms i ed are r m d c ssed i p abeti cal order of tax a. erb c o y c t f o

H lm r n et a l. o g e ,

A rcto hila tr icho da o H lm a a a urali s 1902 . p p o , Ott w N t t

Thi s ax n i s n rall n r t hila ulva in t o ge e y co sidered as a synonym of A c op f (Tr . A n r Its s ur de sson . o ce of publication has been gi ven i n Index Kewensis (P rain 19 13) “ and Index to rass S e i es has il v s no . s . r n . G p c (C e N e 1962 ) as Repert . pec eg i veg 3 : In 1902 h r lm li al n a H u sh d A . tri cho oda i h , oweve , o p b e p o g w t diagnosis and clear holotype citati on (Holm He stated that it w as a new species “ " and ha he i n n u li sh a ull s ri i n and illus ra i n at an arl t t te ded to p b f de c pt o t t o e y date . 74 P H Y T O L O G I A February 1997 volume - 78

m r l s ri ti n was u lish d in the 1907 a r no ill us ra i n Although a o e comp ete de c p o p b e p pe , t t o was provided.

The h l i s CA N 3 6690 hi h w as m un on the sam sh as the o otype , w c o ted e eet

Gri seb . var . aristatum Poluni n CA N holotype of Colpodium fulvum (Trin . ) ( “

v Mscr . Fl ra of a i n A rcto hi la tricho oda s . no . the H u s n and bears the annot t o p p p , o d o ” ’ The h l and an i s US 947554 are r rre B ay Region in Holrn s scri pt . o oty pe otype ( ) efe d Man Tz l . HODOTYPE: s l an to x A rctodupontia scleroclada (Rupr. ) ve ev fie d [M sel] Island H u s n B a CA N Is : Mans ld Mans l Islan H uds n , d o y; otype fie [ e ] d , o

B a r i s r s Dr . R. B ell 61 S . 1884 US y , c ev ce of ock , , ept ;

ia s i a ta L. B eauv . var. rnetorum Pi r Er h a 7: 103 . 1 899 . Dantho n p c ( ) p pe , yt e

a aum 1974 s le ed a l WS 3075 and isolectot e W S Fi ndl y B ( ) e ct ectoty pe , , yp , “ ” “ ” 3079 for his ax n as on the ann a i ns C. V . Pi r and du l. , t t o b ed ot t o by pe type p ty pe , i n i h the r in ns s otolo e . Th n d ha the la l a a are u i e . respecti vely . ey ote t t be d t co te t w t p g , , han ul 1 the specimens were collected on 2 2 July 1890rather t 13 J y 890.

“ im n at US 19 135 82 ars the ann a i n an h ni a s i a a in t rum A spec e , , be ot t o , D t o p c t p e o ’

11 v T s im n i n Pi r s s ri . The ll i n num r l a i n and da . ar ype pec e pe c pt co ect o be , oc t o te on the label are all consistent with the protologue and the indicated habitat differs only “ ” “ l h l n ni r us ds mm n on the s cim n ra h r han dr n s ig t y; In Ope co fe o woo , co o pe e , t e t y Ope ”

ni r us ds i n ra l s il in the rotolo ue. A s indi a d in Art . of the co fe o woo , g ve o p g c te “ ” l a l ma be su rseded if it i s in s ri us n i I BN reuter et a . e C (G , ectotype y p e o co fl ct with the protologue and another element i s available whi ch i s not in confli ct . Also f R mm nda i n 9A .3 ad i s s ha the int n i ns the au h r o the nam e. . eco e t o v e t t e t o of t o e ( g , ” annota tions) are to be used for lectotypification only when not in serious conflict Th r are at l as thre s im ns a aila l hi h are m r with the protologue. e e e t e pec e v b e w c o e nsis n i h the rotolo ue han the l e s l d Findl a aum co te t w t p g t ectotyp e ecte by y B , although the label data are not always as complete as those on U S 19 1 35 82 : two at US (Findlay B aum 1974) and one at W S ( 1482 46) acqui red from the S uk sdorf The s e im n US 1 135 82 is s l c h r as the l Danthonia H erbarium . p c e 9 e e ted e e ectotype of “ " l it ann a i n Pi r as s i a ta var inetorum Pi r. n s ar but it p c . p pe Not o y doe be ot t o by pe type i s more clear ly consi stent with the protologue than the Specimen WS 3075 selected by Fi ndlay B aum

Fe r h Druc R x h . lu ri . Isl s 14 v s uca t a ch lla H a . ex r . E 19 ol. t yp y ck e , epo t Bot c C b B t e , ,

19 15 .

ilkins n S a 1988 ha i n r r e nam Festuca tra ch h lla W o t ce ( ) ve te p eted th e yp y H ack . ex ru as ali l u li sh and an arli r h m n m of Festu a tra ch h lla H a D ce v d y p b ed e e o o y c yp y ( ck . ) i ra h m . n K j na (A cta Bot . Bo e th e former referri g to a South A meri can tax on and the latter to a Eurasian taxon which has been widely di stri buted through the “ ” mm r i al s trad un r the nam H ar il i n n co e c eed e de e d Fescue . W k so S tace ( 1988) state ra ina 1930 w as the rs au h r r nis u var trach h l a K j ( ) fi t t o to ecog e s b . yp y la H ck . at the Darbyshire Pavli ck: Grass nomenclature 75

i l H r H a l ex ru e 19 15 had r i usl us the nam to Spec es leve . oweve , cke ( D c ) p ev o y ed e The one hara r ri l n descri be a totally different tax on from S outh America. c cte (p ck es o

i m F. dumet rum Philli i i on i s the glumes) used to di stin gu sh it fro o pp [s c] n L . ” su fficient to validate the name .

l a ru in 19 15 ul s m The use of Festuca trachyphy la H ck . by D ce wo d ee to have been “ ” r e a un dis ussin a S u h A m ri an a lapsus or orthographic erro . Th cco t c g o t e c species as “ newly adventi ve to Bri tain begins on page 2 9 with the title FES TUCA ” TRA YLLA H a l but n i nues on the n x a i h the ll i n CH YP H cke , co t e t p ge w t fo ow g , ’ quoted (but possibly tra nslated) by Druce from Hackel s letter:

Thi s is surely one of the numerous Chilian species imperfectly descri bed

hi lli i sic and mes er n ar F. dumetorum Phi l . not i nn . hi h by P pp [ ], co v y e , L (w c am F trach le is but it has not the minu ri l s on the I propose to n e . y p , te p ck e

E . HA E in lit . of ha s i s . ferti le glum e t t pec e ) CK L, ( )

the a Dru ce did not apply the nam e Festuca trachyphylla Hack . to dventi ve S outh

n su s u n u li ati ns but us d the nam F. trach le is American tax o in b eq e t p b c o , e e y p H a ard ru 19 19 ru e Thi s i s confu'med in H a ard ru ( yw D ce , D c yw D ce “

1 1 h n the i a i n of the name F. trach le is is i n as Re . . E . 2 9 ( 9 9) w e c t t o y p g ve p B C , “ ” 4 nam F trach h lla Ha el a ar on a 2 hil 1 1 . The . e s 9 F . 9 e yp y ck pp p ge , w e m ach le um t l on inn . hi h I r s to na e F. tr is . a d e orum Phi . n L (w c p opo e y p ppears on ’ a hat r ru s in n i ns in la r ars hi s non a c an e p ge 30. W eve D ce te t o te ye , c ept c of F . trach h lla Ha . ex ruc did not a ear in the sam u li a i n so ha I BN A rt yp y ck D e pp e p b c t o t t C .

cann ot apply .

Th r i s h r am i ui in the inter re a i n of the r n uns in the e e , oweve , b g ty p t t o p o o two ’

hras s ithin ar n h s s in ru s a un . The hi n s s im n s e i s or p e w p e t e e D ce cco t t g ( pec e , p c e “ ” “ ” nam to hi h the thr r n uns it and ha re r can in r r t d e) w c ee p o o , t t ) fe be te p e e “ ” il i ns a e r n u i n di fferent ways. W k on Stace (1988) h ve i nterpreted th p o o n whi ch in r i i n w am the first phrase as applyi ng to Festuca dumetorum Phil . p ov s o of a ne n e for an ill i imate h m n m but it ma als be nsid red r r the su e of eg t o o y ) , y o co e to efe to bj ct “ ” the s n n the s im n amin In the se nd hras it ma e te ce pec e ex ed by Hackel) . co p e y “ i n r r as a l in r u hr th e. F . be te p eted pp y g to either e Object of the p evio s p ase (i . ” ‘ ” tra ch le i or The r n u y p s ) to the subject of the sentence the specimen) . p o o n that ’ “ ” ma i n r r d n in on n s in r re ati n it as a l in i h r the y be te p eted, epe d g o e te p t o of , pp y g e t e to species represented by the specimen or to the Species represented by the name “ ” dumetor um Phi l. If one n i r n dum etorum Phi l the co s de s which as applyi g to F. “ descri ptive phrase but it has not the minute prickles on the fertile glume of that s i s i s not su i n l l ar n u h trach le is or the pec e ffic e t y c e e o g to determine whether F. y p ’ s cim n i s the n i i h the mi nu pe e e t ty w t te prickles. Phi lippi 5 description of F . “ dumeto i nn rum a a : 2 1 86 . a l rae v (L e 36 97. 5 st tes that the spikelets are g ab el omni no ” “ ” hirtae but s not s i all m n i n n ri l the l mm a s i ur , doe pec fic y e t o mi ute p ck es or e ve t t e .

A l h u h Festuca tra ch le is H a ex ru i s un in h Index Kewen is t o g y p ck . D ce fo d bot s Prai n 192 1 and Index to ra ss e ie h w ( ) G Sp c s (C ase Niles 1962 ) as a ne nam e for F . du metorum Phil . no m n i i s ma r n t a ch h lla H a . ex ru h , e t o de of F. yp y ck D ce Bot P ar di 195 3 and Ma th i 1982 n i r tra ch le is H a . ex ru o ( ) t e ( ) co s de ed F. y p ck D ce to be

as o n F . dum etor Phi um l but the nam F. trach h lla H a ex ru b ed , e yp y ck . D ce i s not i n lu rn h ir s n n m li i n S ain i c ded t e y o y y st gs. t Yves ( 192 7) d d not mention either name in hi s m n ra h on S u h A m ri an o og p o t e c fescues . 76 P H Y T O L O G I A February 1997 volume -78

ex ru i h r nsi r d a no men The name Festuca trachyphylla H ack . D ce s e e co de e to be nudum bec ause of ambiguity in the appli cation of the pronouns in the descri ptive ina an a r re a i n all s F. trach h lla H a . ra s s the phrase . This inte p t t o ow yp y ( ck ) K j to t d earliest available name at species rank for the Eurasian tax on.

Lectotypificati on of Festuca names.

r h A m ri an rass s B ea] 1896 i d h l s for In his book on No t e c g e , ( ) c te o oty pe two ’ ’ - l of S ri n r ams n S ri n r s h r ari um . Al s Festuca species names as bei ng i n F. L o c b e e b c b e h um x im ns P an i um w as d s r a fire in A u us 1894in erbari , e cept Spec e of c , e t oyed by g t

Gaz . H r sel l s for h s nam s r m Washi ngton (Bot . e e , we ect ectotype t e e e f o are now r s n at US and ul alm s r ainl ha n isotype materi al . These p e e t wo d o t ce t y ve bee

al al 1 896 a i v . The cim ns i ha n ann a d Pi r seen by Be (Be , p ge ) Spe e c ted ve bee ot te by pe as bein g type materi al .

lada H a . ex al ra ss. N. Amer ed. 2 Festuca dasyc ck Be , G , ‘

S n ral ah : 1875 C C. P a rr 93 . LEC I OTYPE: U ce t Ut , y

Gra ss. N. A m en . 2 Fe onesu V as var . on erta H a . ex al stuca j ey c f ck Be , , ed ,

OTYPE: US US A . ali rni a: S an 1 05 6 rm al 1896 . LEC F C fo No

School .

n ra h on Festuca in rth Am ri a Pi r 1906 i l In hi s mo og p No e c , pe ( ) effect ve y m r als r m the r i usl lectotypified many names in thi s genus. So e of these we e o f o p ev o y w l s for h s nam s d s r h r ari um S ri n r. H r si na ne e t oyed e b of c b e e e , we de g te ectotype t e e e “ ” ’ based on the materi al whi ch Piper indicated as duplicate to the materi al in S cribner s h ari um erb .

v H a . ex al ra ss . N. mer . 2 Festuca s a brella T rr ar . va se a na A c o ey y ck Be , G ed , '

l ra : V a P ass alt. 300 1 896 . LECI OTYP E: U S U S A . 9 Co o do et , [

1 886 . Va se . feet] , , G y

Fe bul ora S ri n . m Ma un t a na d. P l . 1 890. stuca su Ca . ifl c b co , C ‘ LE TYPE: AN A AD i h lum ia: an u r I lan C I O C A . ri s s C N B t Co b V co ve d , l s r am ra l 2 un a un 9 1887 J . M o . Go d t e , g ve , J e , c

Festuca v se a a a na x r 8 6 . H . e al G a ss . N. A m en . 2 1 9 y ck Be , , ed , E 4 . . T T PE: U S U S A . l ra : a Pass 1 8 8 G Va se L C O Y Co o do Vet , , y Darbyshire Pavlick: Grass nomenclature 77

Proposed new combinati ons in Festuca

i n u bu li ora e eev ar h F tu a u nus S u bu la ta e s S E .B . Alex D b s . stat. es c s bge ect o fl ( ) y , r B v I M: Festuca su nus Sub ult o ae E . . Alexee t m no v. A S e co b. B ONY bge fl ,

i n ra 1 80. Ras . en 9 is . V ssh . Novost. S t y t (L g d)

A lexeev ( 1980) considered the elongate and pubescent callus of Festuca subuliflora a r a monos e ific su u so disti nctive in th e genus th t he c e ted the p c bgen s for it. With the ” x i n of hi s har a r h r i a n s i m r h l i al and ana mi al a ur s e cept o t c cte , ot e d g o t c o p o og c to c fe t e are imi lar one or m r s i s assi n d su nus S ubula ta e. In addi i n no s to o e pec e g e to bge t o ,

i n si aria i n was d t d n F. ula ta difference in c NA restri ct o te v t o etec e betwee sub T ri m. and ubuli o ra ar shi r ar i 1992 in i a in a l s r la i nshi F. s fl by D by e W w ck ( ) d c t g c o e e t o p w u n ubula and t r between the type Species of the t o s bge era S ta e Subulflo a e.

I : Festuca r li n om . no v . A S M rub a . ub F estu a a m m obi a Pa s s . c v ck , B ONY L p

v M s . Obva. Is t. Prit . Otd. i l . a r n icola E.B . A lexee B ull. fi s . e , y o k py B o g

r i ola Prodin 8 00 . 1973 . 2 . non Festuca a en 3 . 198 c ( )

F estu a s eu do vi vi a r a Pa li P a li omb. et sta t. n o v. AS I N M: c p p ( v ck) v ck , c B O Y

Festuca rubra . subs . aucta rma seudovi vi a ra Pa li Ph tolo i a L p fo p p v ck , y g

1985 .

u bra . subs . m edi a n a Pa li Pa li mb ta t. F est a r u o . ct s n o v. c L p ( v ck) v ck , c M: estuca rubra u e var med A SI F . s bs . s unda Pr sl . ia na Pa li B ONY L p c e v ck ,

Phytologia 1985 .

F estu a u ver i i lla t E . B . ex eev a i l i oli a r D ar sh . s b t c a P rs. A l rm os b c ( e ) fo p f (Do e) y ,

omb. n v A M: Fe tuca bt a iehler rma i i olia r in o . SI s o us B los c B ONY fo p f Do e , McNeill r a ural 1 7 . i s an a i n 103 60. Do e , N t te C d e 5 97

LI TERATURE CITED

lex v A ee BB . 1980. Festu a a x A m ri r ali et , c L . S ubgenera et secti ones nov e e e ca bo e M v - i a . No ost i V x S s . . In R ssi an . . enin ra 5 3 . u e c t yssh Rast . (L g d) [ ] l W a J . 1 r G a sse . 896 . H n H l Co s o Nor h A m er i ca lum 2 . r Be , f t . Vo e e y o t , New

r r . Yo k , New Yo k ha A . . s il . . de to ra eci es . . K . H all s n s 1962 . In x ss S G C e , C D N e G p , Bo to , M assa hu c setts . ar hi r s S . S J . G ar A m an F a . 1 2 . Ph l n of r h ri es uc D by e , W wick . 99 y oge y No t e c t P ac a an d r la d i a ( o e e) e te genera usi ng chloroplast DNA restri ction s te vari tion . ana B t - . . o . 2 42 C d J 9 . 78 P H Y T 0 L O G I A February 1997 volume - 78

5 . Plan G. C . 19 1 n s . ani al ruc for 4. R r of the S i D e , t ote , etc , 19 1 epo t Bot c oc ety an lu the ri i - Exch ge C b of B t sh Isles 4: 1 48 .

ru G.C. 192 8 . B ri tish P la nt List onta inin the S en na to h tes P terido h tes D ce , C g p p y , p y and Cha ro h tes Found Either as Na ti es or rowin in a Wild S a te in B ri ta in p y v G g t ,

Ireland and the hannel Isles . 2 nd ed . T . un l Co . A r r a h r a , C B e e , b o t , G e t

4 m n la u al i li i 1 7 . The n r m a ns the ax n Fi ndla N . B .R. aum . 9 m y , L B o e c t p c t o of t o o y - 1 82 an ad. . . 5 . of Dan tho nia i n Can ada . C J Bot

Demouli n . t W hal n r V . H a W . . urde . G. . s rt h reuter . H M B P . M . G , , , C o e , , D L wk wo , i a eh i ls n P . C . S l P . Tr ane . McNeil r n n D . H . l. 1 4. Jo ge se , N co o , v , , J 99 t i a l men ur T A Interna tiona l Code of Bo an c No cla t e. ( okyo Code) . dopted by the n h rna i nal ani al n r ss hama A u us - S m r 1 fiftee t Inte t o Bot c Co g e , Yoko , g t epte be 993 .

S i n i s oni s in rm an . Koeltz c e t fic Book , K g te , Ge y i l r w id 1 1 . The Adven e F o a o T eeds e. T . B u M . G.C . ru . 9 9 ncle H a ard I . yw , D ce t v f

h . Co . A r r a , b o t r hila a a a urali - On the nus A cto Ru r. s 85 . H lm T . 1902 . o , ge p p Ott w N t t Index H er ba r ior um lm r n L.C . arn . 1990. . Par I: H lm r n P . K. NH . H o g e , , o g e , B ett t

- - 8 . R num V a il l 2 oz i x 1 693 . r The h r ari a of the rl . . e b wo d ed eg eget b e , New Yo k

ard. r nx r . Bot . G , B o , New Yo k l a - Fe t a P a a en hi . ana 37: 67. 82 . El n r s uc 1 Ma h i . 19 tt e , O ge e o ( o ce e) C e G y

Las s i s Festuca la Pa a ni a. R is a A r n i n Par i L . R. 195 3 . a od , e pec e de de t go ev t ge t de - 2 2 A gronomia 2 0: 177 9 .

6 . r h Am ri an s i s Festuca . n ri . U . S . Nat. H r Pi r C V . 190 . pe , No t e c pec e of Co t b e b 1 - -48 vii - i x 10 . . i vi 1 . pp , ,

Index Kewensis. S u lementum ua rtum . x r Prai n . 1 13 . at the , D 9 pp Q O fo d ,

lar nd n xf r r a ri ai n. C e o , O o d, G e t B t

dex Kewensis . u lementum uintum . x rd at Prain . 192 1 . In S the , D pp Q O fo ,

lar n n x r r a ri ai n. C e do , O fo d , G e t B t ' - - S ai n s A . 192 7. n ri u i n a 1 é ud des Festuca sub en . Etc Festuca t Yve , Co t b t o t e ( g ) de ' - an ll a 3 : 15 1 3 15 . 1 A méri que du S ud. C do e

il ins n MJ . C . A . S a . 1988 The ax n mi r la i nshi s and W k o , t ce t o o c e t o p i n tuca r i i d F lema nii ar f la r . typ ficatio o Fes b ev p T acey an B ast d (Poaceae) .

- Watsoni a 2 99 .

80 P H Y T O L O G I A February 1997 volume - 8 1

- in humid in ak r 1 2 . IX 1 76 . uare ss on re o s 2 640 rn . 9 0 S E . in mo t e p e fo e t, , , z,

- M l . reenwood E. Ha sa er G 2 96 A O G , g t ( )

- E FA LLE N S Sz la h . Es o z F rrari omb. n o v . DEIRE GYN ( c ) pejo L pe e , c

r ollens S z l ra m . Fl ri s . Geobot BAS IONYM: Oestlundo chis p ach F g o t .

T P S : ME I . ax a a: mi les r m i az r az on the 2 . 199 1 . fig . , 5 Y U X CO O c f o D O d

- r ad Cua imalo as 19 . V 111 . 1968 . P o lla rd S 1 2 6 SEL . o to j y , , G ( )

- F E IA KIA NA S z la h . Es o z rrari omb. n E H V o v . DEIRE GYN S ( c ) pejo L pe e , c

lundo rchis shevia kia na S z la h . Fra m . Fl ri s . Ge : Oest obot . BA SIONYM c , g o t

- ME I . hi a as: S an F li illa n ar 2 3 . 199 1 . T P S : fig . Y U X CO C p e pe , v ge e “ las asas S an ri s o al ML Hui c Eca 3 mil s n Ciudad C ( C t b ) , tepe tepec e W of tow , Ca rlson 1 600

ex ra m . Fl ri . Geobot. iranthes carlsona e . . illiams S z la h. F s Sp L O W c , g o t

ro. s n . 1993 . p y

- H IEDEELLA ON F USA ara Es o z F rrari omb . no v . S C C (G y) pejo L pe e , c

Mus. Leafl. . 8 SI N M: Dei re ne on usa ara . 1 C. 1980. BA O Y gy c f G y , Bot t

F kiella duran ens is A nt s . Schwei nf z la v unc g ( e C .) S ch . ar . “ ” a a m Geobot. 199 1 Funkiella on a ara S z l h. Fr . Fl ri s . . c fus (G y) c , g o t , Hi al : a n f Mez titlan a the h : ME I . o r n TYPUS X CO d go L goo , bove S o e betwee ° ' ° ' r s i n hi h rass 2 0 40 98 5 1 1600m 2 71 11 1933 J . Gon ale sub ock g g , N , W , , , z z lund 2 1 4 E. Oest 9

- S CH IEDEELLA RE N ULA TA . . illi ams Es o z F rrari omb. C (L O W ) pejo L pe e , c

lineata in . var . renula ta . . illiams . no v. AS I N M: S iranthes tri dl B O Y p L c L O W , Bot

- Leafl . 2 3 . 1 46 . renulata . . illi ams ara B ot . Mus . 7 9 Gula ria c (L O W ) G y , dee la tr i n u Leafl . . 2 8C. 1980. Schie l linea ta i dl . M s. t (L )

- v la ta . . illiams z la h . Fra m . Fl ri s . eob t um s B al . ar . renu S G o . B c (L O W ) c , g o t ME I Pu la: ass ld La M a . 2 f. 199 1 . T P S : . r s fig Y U X CO eb g fie , e , ° ' ° ' ax a 2 0 1 1 98 00 1 600 m 2 1 . II . 1 932 F. Froderstrom sub E . Nec , N , W , , , Oestlund 2 592 (AMES ")

S CH I EDEE LLA P A N D URA TA ara z - F Es o rr ari omb . n o v . (G y) pejo L pe e , c A SI Deire ne r M: ata ara . us . Lea fl . andu M . 2 1A . B ONY gy p G y , Bot t ‘

1980. Oestlundorchis andurata ara S z la h. Fra m . Fl ri s . p (G y) c , g o t ’

Geobot. . 2 . 1 1 P v . uran : n Guance r fig 99 TY US : ME XICO . D go betwee

° ' ° ' ° ' ° ’ and ua alu al 2 5 56 - 2 6 2 8 105 56 - 105 36 2 000- 2 400 m G d pe y C vo , N , W , , “ ale u E J . on estlund 0 1 0 AMES G z z s b . O 5 1 8 5 8 ( ") o z - F rr ari : M xi an m n s 8 1 Espejo S . L pe e e c o ocot

A CKNOWLEDGMENTS

h x r ss our in er a r i a i n th e ura rs the h r aria r is We wi s to e p e S c e pp ec t o to c to of e b ev ed. a and a i r ar ia ruz for h ir r i of hi s m anus ri T Also to Ramon Rib J v e G c C t e ev ew t c pt . his NA B IO r 16 work was partially supported by CO (p oject GO ) .

LITERATURE CI TED

- i ni i n i n u ri P . 1 82 . n r re s S iranthi nae r hi um al h 9 a a . B s B og , Ge e c defi t o bt be p (O c d ce e)

- 1 132 . A mer . J . Bot . - M ir an urn al h P . 1986 . A s n si s xi an S thinae. r ui a Méxi B s B og , y op of e c p O q de ( co

- City) 96 .

i i si n of the S iranthi nae . . u a 82 . A n r M s. Le fl . Gar a L . A . 19 y , ge et c ev o p Bot 42 5 . i schen u r nun 2 0. V rsu h in r S sternat der S iran thin ae S hl h r R. 19 . c ec te , e c e e y Ne o d g p - 454. B ot. Centralbl. Beih . he n ra chiedeella and Funki e S z lachetko . 199 1 a. s on t S lla r hi a a , D Note ge e (O c d ce e ,

- Fra . Fl ri . Geobot. 2 1 . Spiranthi nae) . gm o st

S z lachetko . 199 1b . nera and i s of the su ri S iranthi nae , D Ge Spec e bt be p le hi a ea . 1 . n ri uti n the r isi n of Ga otiella . Fra . (Orc d c e) Co t b o to ev o gm Flori st .

- Geobot. 12 .

S z lachetko . 199 10. Oestlundorchis r hida a S ir an thi nae a new r hi , D (O c ce e, p ) , o c d m - n a Fra m . Fl ri s . Geobot. 3 1 . ge us from Meso erica. g o t

S z lachetko . 199 1d. B urnsba lo hia a new m xi an nus of the su ri , D g , e c ge bt be -4 F lia t. Ph tax . 02 . S piranthi nae (Orchidaceae) . o Geobo yto

S z lachetko . l 992 a. n ra and s e i s of the su ri S iranthi nae , D Ge e p c e bt be p

hi a . 2 v r c a . A re ision chiedeella ra m . Fl ri s . Geobot. (O c d e e) of S . F g o t 4 2 0 . la S z chetk o . 1992 b . n ra and s i s the su ri S ira nthi nae , D Ge e pec e of bt be p hi a a 4 - r . P h so ne. Fra m . Fl . eo t. 43 6 . (O c d ce e) . y gy g ori st G bo S z lachetk i iran thi nae o . 1993a . en ra and i s the su r S , D G e Spec e of bt be p

- r hi da a . . P 7 Fra . Fl ri s . Geobot. 103 . (O c ce e) seudogoodyera . gm o t

S z lache ko . 1 thin t 993b . n ra an d i s of the su ri S iran ae , D Ge e Spec e bt be p hi a a r Fl ri s . Geobot . u l . . 6 . A r i si l a m . S (O c d ce e) ev on of Funkie la . Fr g o t pp

- 2 5 0. S z lac hetko . 1993c . n ra and i f the su ri S iranthin ae , D Ge e spec es o bt be p hi a a r . 8 . A rev i r i Fra . Fl ri s . Geobot. (O c d ce e) i s on of Oestlundo ch s . gm o t

1 2 9 .

S z lache ko . 1 4 an t I . Mis ll a 99 a. S u i s on the S iranthin ae r hi ac a . , D t d e p (O c d e e) ce e

- a Fl ri s . Fr m . Geobot 4 g o t . 3 8 . S z lachetk 4 anthi nae o . 199 b . n ra an i s of the su ri S ir , D Ge e d Spec e bt be p r hi a a 1 r - . 0. L o lossa . Fra m Fl ri s . Geobo t . 12 7. (O c d ce e) y g g . o t S z lach tko . e 1 4c na a I . 99 . S u i s on the S iranthi e r hi a . 10. I , D t d e p (O c d ce e) Co leorchis and W r Fra m . a scaea two genera of the Cyclopogon com plex . g Fl ri Geobot - s . 2 o t . 5 6 . - P hytologia (February 199 7) 85 .

REB UTTA L TO T ERRE LL’ S TAXONO MIC NOTES OF TURNER’ S TREATMENT OF TEXA N AND MEXICA N H ED YO TIS

L. Turn r B . e

ar m n of an ni rsi of T x as A us i n T x as 78713 U . S . A . Dep t e t Bot y , U ve ty e , t , e

ABSTR A C T

’ A vigorous rebuttal to Terrell s evaluation of my treatment of Texan and t u ha ula i nal r Mexican H edyotis i s presented. I is s ggested t t pop t o wo k will prove pivotal in judging between the merits of the two co ntrasting m n la ural s stems hi s r rm d un ias d l r rs usin no e c t y , t to be pe fo e by b e fie d wo ke g more Sophisticated techni ques than those employed by the contestants

concerned .

KEY R S : Ru i a a H ed otis H ouston ia Mexi co T x as T rrell WO D b ce e, y , , , e , e ,

“ ” Tenell ( 1996b) provided a critique of my tax onomic treatment of the H edyoti s ni ri ans F s er m l x T urn r 1995 a and h r l m n s of H ed otis g c (Lam ) o b g co p e ( e ) , yet ot e e e e t y

urri n in T xas and Mexico Turn r 1995b 1995 c l995d . Thi s a r an occ g e ( e , , ) fte in r du r d ns hi s ac an of H ed o tis H ous tonia and Oldenla ndia t o cto y efe e of cept ce y , , , hi h lassi a i n I did not ll r rri n ins a ll ha F s r w c c fic t o fo ow, p efe g te d to fo ow t t of o be g 1937 and Shinn rs if not h r r rs who i H ed otis in the r ad ( ) e ot e wo ke , v ew y b o

s ns r atin the s ral n ri s r at s as but infra eneric a ri s. The e e , t e g eve ge e c eg eg e g c tego e in formed reader will understand that either H edyotis or Hedyotis (S S ) i s equally a a l unl ss it can sh n ha the rm r i s a n l l h l i or erha s ccept b e, e be ow t t fo e p te t y po yp y et c p p

ara h letic hi h m n l d has not n d m nstra d. p p y , w c to y k ow e ge bee e o te

The only substantive comment made by Terrell in regard to my paper i s that he all e r c ed to th fore an error or lapse in my key to the varieties of H edyotis a ce osa A . ‘ ” ra in hi h l a v as i ula ta for the in nd d G y w c I inexp ic bly substi tuted the name ar. f c c te e var . a erosa . B ut hi s i s no bi al: an r a r ul ha d the la s and c t g de y e de co d ve detecte p e , ma all an l de ow ces according y .

What Terrell fails to comment upon adequately in hi s paper i s the cons iderable i n r r da i n i v v a d ot s acerosa ar . er sa and H a r n H e . . ol emoides te g t o betwee y ac o ar. p yp

A . r W a . H . i hi w mm n ( G y) Lew s i n west central Tex as . T s as co e ted upon and map in d ail me c Fi ur ut h r et by ( f. g e b t ese we e crudely remapped and treated by Terrell ’ Turner: Rebuttal to Terrell s revi ew of H edyotis 8 3

f a r noni nter radin s m atri su s ci s al h u he n d ha h r as i the tax we e g g y p c b pe e , t o gh ote t t t e e i ns rla as als surmi w a a i n in r I s d . S in T rr ll l as i ntergr d t o eg o of ove p , o e ce e e c aims to ha d n ld r er the ran of H . acerosa as I m s l ha he must n ve o e fie wo k ov ge , y e f ve , k ow i n int r rada i n the ula i ns ar som ha in s im that in the reg o of e g t o , pop t o v y , e v g pec ens

l m lin var . acerosa s m i h im ns m s l res m li n v ar most y rese b g , o e w t Spec e o t y e b g . It ma rs not if Tenell ish s call h s in r ra i axa polypremoides . tte w e to t e e two te g d ng t i : the la r a ri es can a h h us a mor ho eo ra hi cal ari mu subspec es tte c tego e c o e p g g p v ety , ch a in l i He mi h thi as a monotypic subgenus can house S g e Spec es. g t nk that the use of “ ” “ ” u s i s and ari are mu uall x lusi but I not r ad the terms s b pec e v ety t y e c ve , do e the i a l omen la ture in hi s ashi n nor h ul T In erna tiona l ode o B ota n N rr ll. t C f c c t f o , S o d e e

a s 1979 but see hi s c n rar i s as x r ss in 1996 as n d Tene ll st te ( , o t y v ew e p e ed ote “ below) that the differences between these two enti ties are on a hi gher level than the i n e o hi al se ara i n in M xi o and a usual ari . In a i th ra a n v ety dd t o , ge g p c p t o New e c dj ce t ” ar s i int rada i n in st rn T x as and n r h rn M xi arl m r . lands i s cle y ked , de p te e g t o we e e o t e e co I never deni ed that the two tax a mi ght not belong to different subspecies: I merely hea ted the two at the varietal level consi stent with the treafinent accorded most in r radi n infraS ecific a ri al uni s m s m d rn r rs r n ui s te g g p c tego c t by o t o e wo ke C o q t , 6 et a adi a 1 6 for a r as n numerous publicati ons; Turner 195 ; c of. K p 9 3 e o able review) . I reco niz ed ur mor ho eo ra hi cal el m n s und r the a ri of H ed oti s a erosa g fo p g g p e e t e f b c y c , a r a h Tha Tenell mi h not think h providi ng a key to these an d m ps fo e c . t g t t ese ula i nal uni s rt h r ni i n is fine i h me but he has not r d an pop t o t wo y of ecog t o w t , offe e y real data to di sprove their reality .

“ T rr ll n nds ha the e s cim n of H ed otis a erosa Wri ht 2 3 7 see e e co te t t ty p pe e y c , g ( ‘ ’ T rr ll 1996a was lle d i n Westem Texas El Pas M xi in 1 849 e e ) , co cte to o , New e co but he ails n t ha hi s nam n ds lectot ification as I learl in d out f to o e t t t e ee yp , c y po te , and h a Wri ht 2 3 7 was ll in ha is now inn or V al V r un i s t t g co ected w t K ey e de co t e , T x as as the P s Ri r uri n une 1 849 and T rrell re he ha e (e t of eco ve ) d g J of , e (we to ve ’ l e hi s u in ri h s u li shed eld n s nee d not as ri its e l ali as ook d t p W g t p b fi ote ) , c be typ oc ty “ “ s m h r e n s rn T x as and El Pas M xi the la r o ew e e b twee we te e o , New e co tte r i n a ra h r m ani n l ss l al eo ra hi call ea in . eg o t e e g e oc e , g g p y Sp k g

Furth r c mm n in u n the ra i of m var . otosina B .L . Turn r he e , o e t g po ve c ty y p e , contends that the pulvinate low plants from southernmost Coahuila and S an Luis “ P si Méxi ha i n l n a r lla u s are ar of a li n hat n r h ard has oto , co , v g e o g te co o t be , p t c e t o t w all r lan s i h r In s uth rn T x as a la s t e p t w t coarse leaves and longer i ntem odes. o e e [ p e h r he sh ul ha sai i n s rn or rans P e s h r are ll i ns ha e e , o d ve d we te t co t e e co ect o t t ” are s m ha ransi i nal i h ra h r fine l a es and small s a ur . I a the o ew t t t o , w t t e e v t t e t ke few Sheets he cited in defense of thi s statement to be depauperate or otherwi se atypical

lan s v ar . acerosa as urs in ula i ns r w h r h s r mar a l p t of , occ pop t o eve y e e , be t e e e k b y ari a l or mar a l in o ula tions in n r h n ra l r ari a l . In ru h h r are no v b e e k b y v b e t t , t e e p p o t ce t Méxi or s v otosina alle the co we tern Tex as which re semble the popula tions of ar. p c d to r in m a r nor is h r a lin o ula tions n h s in the r i n fo e y p pe , t e e c e of p p betwee t e e eg o n ha w co cerned . I ve traveled over thi s terrain many years no an d would have been ha fi nd su h u a r b t n n wa a var . otosina i s r l s s r d. In ppy to c , o e ob e ve f ct , p p ob b y bette m ar han var . o l r em oides hi h T rr ll a rds subs ecific s a us and I w as ked t p yp , w c e e cco p t t , “ ” sur ri s see ha T v o osina in its ow n p ed to t t errell did not elevate or positi on ar . p t su s i s as he di d v ar . ol remoides r ai nl the la t r sh s mu h r a r b pec e , p yp ; ce t y , t e ow c g e te “ ” linal i n r i r a a n r a r a r area h an s var. o osina as i s l ar r m c te g d t o ove b o de t doe p t , c e f o h of our is ri u i nal m a s f rm r m l x imilar mm n ul als bot d t b t o p o the fo e co p e . A s co e t co d o 84 P H Y T O L O G I A February 1997 volume - 85

an a an e m var. so hi la B . L. T urn r his in be made about his reluct t ccept c of y gyp p e , t be g

l remo ides . Tenell 1996b al s i l as s d r much better marked than var . po yp ( ) o gl b y p e ove

ha m r a l ir ums ri var. tama uli ana B .L . T urn r n ndin my somew t o e we k y c c c bed p e , co te g hil ll is la d and ss ss d a di f ri n al x and r lla it was not that, w e we o te po e e of f e g c y co o , t m m l a ll i n v wort hy of recogni tion because i so ewhat rese b ed co ect o of ar . - P s T xas Rollins hamber po lypremoides from Jeff Davis Co . in trans eco e ( C s 2 75 9 as if a single atypical element from the mile- high volcani c outcrops of the Davis Mountains mi ght miti gate the localiz ed vari ation found in the populational uni ts of the much lower calcareous outcrops of the Tamauli pan shrublands of northeastern

Mexico .

’ r di s ur i n me at l as has n T rr ll s n usi n res nta i n of the Mo e t b g , to e t , bee e e co f g p e t o

ific ari a i n un in H ed otis acerosa . Thus T rrell ri i nall infraspec v t o fo d y , e o g y " u s ies i n hi s ax n but in his 1996a re isi n he s a d ha The recogniz ed two s b pec t t o , v o t te t t ms a c n inuum h re are no a ar n dis n inui ies and n in H . acerosa s variatio [ ] ee o t ; t e pp e t co t t , ” r a l i s t ll: so wh r in n the su s I now recogniz e only obse v b e Spec e . Pray e y e ve t b pecies so as to casti gate my recogni ti on of these at the varietal level"

Terrell ( l 996b) also wax es in a grumbling fashi on about my recognition of '

W B . L. Turn H d ti s a bnen A . ra . H . e is var . mu ui ana r hi h i s e yo p ( G y) L w zq z e , w c l arl a mo ho eo ra hical ula i nal uni but mus we li his s a m n ha c e y rp g g p pop t o t, t be eve t te e t t t “ My presently limited sample lea ves me [T errell] reluctant to accept the exi sting m r h l i al da a as n lusi n rnin mu ui ana ndi n urth r c ll in o p o og c t co c ve co ce g z q z , pe g f e o ect g " it " In m ini n h re are su fi i n ll cti ns of the ax n LL TEX to of y op o , t e f c e t co e o t o ( , ) n the nam I ha r os d al h u h T rr ll mi h m an he r rs ll c ve ture e ve p op e , t o g e e g t e p efe to co e t e u a i thi s himself before accepting th p t t ve tax on.

It should also be noted that Terrell ( 1996b) sweeps under the fabric of H edyotis

ni ricans var . ni ricans m r s d ari i s H . n . var . austrotexana B .L. Turn r g g y p opo e v et e , e “

llac B . L . Turn r laimin the hara rs s h and H n . var . i ea aratin ese to . pap e , c g c cte ep g t be ” n h e i var oridana min r. B ut no m r mi r an th ari s H . n . . S andl. and H n . o , o e o t v et e fl t . l ta a l r n i a var u ina h di f d endernics Fl ri da hi h Tenell a ed. . p v , bot we k y f e e t te of o , w c ccept In had T rr ll not r niz ed the la t r mor ho eo ra hi cal uni s as rth deed , e e ecog t e two p g g p t wo y of n m n la ural s a us I rha s ul not ha r ided rm al ari al s a us o e c t t t , pe p wo d ve p ov fo v et t t to the T xas ula i nal s s ms . Thus Tenell set the minimal s andar s for two e pop t o y te , t d ari al ni ri ans in hi s 1 86 a r Si a v et recogni ti on within H . g c 9 p pe ( d

Fi nall it s ms r h n in ha T rrell ass s r m r s H ed ti y , ee wo t ot g t t e p e ove y p opo ed yo s

oolea na B .L. Turn r T urn r l 995 d laimin his but ar the ari a i n of p e ( e ) , c g t to be p t of v t o his nc f H mullerae F s r a i n r h c n ral Mex ic w o . s o . I as co ept o be g , Spec e of o t e t isa in ha he i ne n nl ooleana d ppo ted t t de g d eve to examine the o y specimen of H . p

n n me the h l T EX . The di f r n s w n m r s H . k ow to , o oty pe ( ) f e e ce bet ee y p opo ed ooleana and hi H mullerae are rtainl the n p s . ce y as great as differe ces between H oustonia butterw ickia e Terrell (a localiz ed taxon in close prox imi ty to H edyoti s oolea na and the i r a H ed ti r B ut he who r s a ax n li p ) w desp e d yo s nig icans . e ect t o kes to s and it m s l in lu . t by , y e f c ded

In the nal anal si s an two i ri n s s m a i rea m n s are li l s fi y , y d ffe g y te t c t t e t ke y to be te ted l r rs who ill a s h ir ula i nal ali i or DNA r rs by fie d wo ke , w tte t to t e pop t o v d ty , by wo ke usin r s ri i n i anal sis or s m su h the la t r r sum a l a h r g e t ct o S te y , o e c , t e p e b y g t e ed and

- - anal z i h u i as. I si ncerel li ha m lassi a i n of H ed otis vi s a vi s y ed w t o t b y be eve t t y c fic t o y , ’ Turner: Rebu ttal to Terrell s re view of H edyotis 8 5

ha of T rr ll ill me the s s im s rha s Tenell ls the t t e e , w et te t po ed; pe p fee same about hi s classificati on .

Fi ld r rs in th e r i n n rn d m s l i n lu d h ul ma a n rt e wo ke eg o co ce e , y e f c de , S o d ke co ce ed

r xamin and mm n u n o ulationa l uni s of H ed o tis . In u h effo t to e e co e t po p p t y deed , s c d x ra la i ns r m hese led m arti ul ar rea m n observations an e t po t o f o t to y p c t t e t . I do not doubt that Terrell has done considerable field work in Tex as and perhaps north central ex ic but hi s has not n es iall i us me in the ll i ns he ha M o, t bee pec y obv o to co ect o s ass m l and dis ri u d nor i s hi s i us r m the in rm a i n r s n d in i e b ed t b te , t obv o f o fo t o p e e te h s cri ti que of my own work .

Th r i s a rui sm in s s ma i an or sh ul : h r hi n s in ual e e t y te t c bot y , o d be ot e t g be g eq d x ri n the s s ma i s m s li l n s the t. e n an ( brai s e pe e ce) , y te t t o t ke y to k ow be t Specific and i nfraspecific boundari es of a given group i s that worker having the most field I are not to ud the m ri ex peri ence with the taxa concerned. c j ge e ts of the two an a in the r s n n r rs but I do find the m in h th t goni sts p e e t co t ove y , co pet g ypo eses Stimulati ng; hopefully some youn ger worker with more field ex peri ence and better sampli ng techni ques will ultimately resolve the systematic problems posed i n the r n a r p ese t p pe .

i F s r FR . 1937. S m Ru i a a s u h as rn P l n si a. as nal Pa rs o be g , o e b ce e of o t e te o y e Occ o pe i sh us - 2 B op M . 93 .

a adi a Z ] . 1963 . arie ies and su s i s a su s i n ards rea r K p , V t b pec e , gge t o tow g te u - niformity . Tax on 2 5 8 . 4 Shi nn rs L. H . 19 9 . Trans r of T xas i s H oustonia H ed o ti s . Fi l e , fe e Spec e of to y e d a ra r — 16 L bo to y 9 . T - rr ll E . E . l 996a . R i si n of H oustonia Ru i a a H ed otideae . S s . B ot . e e , ev o ( b ce e y ) y t

- Monographs 1 18 .

T rrell E .E . 1996b . Tax n mi n s on T x as and M xi an ci s of H ed otis e , o o c ote e e c Spe e y and H t n P i a - 4 ous o ia u i a a . h l (R b ce e) yto og 1 1 . T u rn r B .L . 195 6 . A t ax n mi ud f the nus H meno a u e , cy ot o o c St y o ge y p pp s m si a Rh - . od ra 0 (Co po t e) o 3 8 . Turn r B .L . 1995 a . Tax n mi r i f ed ti ni ricans Ru ia a and e , o o c ove v ew o H yo s g ( b ce e) l s l alli e — ax a. Ph t i a c o e y d t y olog 2 1 . Tu B . L rn r . 1995b . Tax n i r Ru i a a m l . e , o omy of the H edyot s a ce osa ( b ce e) co p ex Ph tolo ia - y g 88 . Tu L rn r B . Ph l a . 1995 c . Tax n mi s u f e ti a lmeri u ia a . to o i e , o o c t dy o H dyo s p (R b ce e) y g

- 92 . Tu rn r B . L . l 995d. H ed otis ooleana Ru i a a a new i s r m the a e , y p ( b ce e) , Spec e f o De d H rs M n i ns T a - - u a r ns P s T x as . Ph tolo ia 96 . o e o t , eco e y g - P hytologic (February 199 7) 93 .

M UGOS P ERMUM HOLZ. NEW O LOU S ANA W TH NOT TA LINU R , T I I I ES ON T ERETE -LEAVE D TA LINUM IN LO UIS IA NA

M . H . MacRoberts B .R MacRoberts .

4 ui i an a R s ar h 7 0 lum ia S hr r o s 71 104 U S A . Bog e e c , Co b , evepo t , L

H r arium Mus um of i S i n s ouisi ana S a ni rsi i n S hr r e b , e L fe c e ce , L t te U ve ty evepo t,

S hr r ui si an a 71 1 15 U . S . A . evepo t , Lo

A BSTRA C T

Ta linum ru os ermum H lz . P rtula a eae r i usl unr r d in g p o ( o c c ) , p ev o y epo te T lin ui iana has en un in x ri sand lands i n add Pari sh . a um Lo s , be fo d e c y C o

r v orum u . als urs in ui si ana but on sali n rai ri s and pa ifl N tt o occ Lo , e p e

sandstone outcrops.

KEY R S : Talin um ru os ermum Ta linum a rv orum P r ula a a WO D g p , p ifl , o t c ce e, Loui siana

INTRODUCTION/METHODS

Ta linum rugosperm um Holz a north-central and Great Plai ns Species recently

un in T xas ix n et a l. 1980 hran 1 3 S in hurst 1 96 T x as a ural fo d e (N o , ; Coc e 99 ; g 9 ; e N t Heri tage Program 1995 ; Tex as Organiz a tion for Endangered Species has not been report ed for Loui siana (MacRoberts 1989; Teague Wendt 1994; Louisiana Natural Heritage Program

In li ght of the frequency with which S i nghurst ( 1996) reported Ta linum ru os ermum in as T x as r x ami n h r arium ll i ns g p e t e , we ee ed e b co ect o r m oui iana H r arium ma ri al for r - l a Ta linum n f o L s . e b te te ete e ved i s ofte indeterminate: s ld m are rs ll d s s are asil l s a er the lan has n r ss d and e o flowe co ecte , eed e y o t ft p t bee p e e , r s are not al a s r s n . Fr sh or at l as ll r s r ma rial i s alm s a oot w y p e e t e , e t we p e e ved , te o t n ssi for a si i i nti a i n ar Th r r innin i n Ma 1997 ece ty po t ve de fic t o (W e e efo e, beg g y we exami ned flowering specimens in the field an d plants collected from ten loc ati ons in as T x as and s uisiana and main ain i la a e t e we t Lo t ed n the bor tory .

Our laboratory sam ple consisted of plants from five sandstone outcrops in a hi h s and S a in arish s uisiana and as r un T x as MacR N tc toc e b e p e , Lo , J pe Co ty , e ( oberts MacRoberts 1993 l 995 a Mar i a ix n ali n , ; ett N o two s e prairi es in Wi nn and

88 P H Y r o 1. 0 G I A February 1997 volume - 93

Raf. P aron chia dnumnondu T rre A . ra P h sa li s hetero h lla s P in us , y o y G y , y p y Nee , da P ediomelum h P Mill P inus tae o a eum N u . x rre echinata . , L yp g ( tt e To y A .

h . m ra R d . var . subula tum us I ri s P runus um bella ta Ell. uer us incana G y) y b (B ) G e , , Q c

lla ta an hus a roma ti a A it. utellar ia r h la n artr . . ste R Sc ca diO l E lm . A B , Q W g , c , p y ge .

ra Stilli n ia s l atica L. S t losanthes bt ora L. B . S . P Trades antia G y , g y v , y fl ( ) c honii ush Va cin ium a rboreum Marsh Vi tis aesti va lis Mi hx Vitis reverc B , c , c i rotundifolia Michx and Yucca lou sianensis Trel.

Ta linum rugospermum in east Tex as occurs in similar habitat with the same

a i s Si n hurst 1996 rs. associ ted spec e ( g , pe

x ami n d h r ari um ll i ns r - l a d ui sian a Ta lin um r m NLU We e e e b co ect o of te ete e ve Lo f o , For r as ns i n a s m s im ns r indetemrinate LS US and LS U . . , e o g ve bove , o e pec e we e

in mainl s d and ha i a hara ri s i s n lud ha T. a r vi orum urs Us g y ee b t t c cte t c , we co c e t t p fl occ in S a ine add and a hi ches ari sh s . Our s r a i ns on r sh ma ri al b , C o , N tc to p e ob e v t o f e te de ni el ut T. arvi o rum in add S a in inn and a hi h s ar i sh s. fi t y p p fl C o , b e , W , N tc toc e p e ri al for Red Ri r and V ern n ari sh s is ind rmin a but the ha i a Herbarium mate ve o p e ete te, b t t r Ex r im n su s h are T a r vi o um . u and plant siz e for both spec e s gge t t ey . p fl cept o earlier Caddo Pari sh collection (Ma cRoberts MacRoberts 2 75 7 [LS US ] origi nally misid n i d as T. a rvi orum n n the her ari um s im ns i s T e t fie p fl ) , o e of b pec e . r ugospermum .

The dis ri uti n of r e- l a Ta lin um i n uisiana is sh n i n Fi ur 2 t b o te et e ved Lo ow g e .

We found Ta lin um pa r viflorum in two habitats : Shallow soils on sandstone outcrops in S abine and Natchitoches pari shes and on sali ne prairies in Winn and ha des ri d the rm r san s n u r lan mmuni Caddo pari shes. We ve c be fo e d to e o tc op p t co ty in detail elsewhere (MacRoberts MacRoberts The saline prairies have yet to receive detailed attention (Smith Both habitats are very di fferent from the xeri c d lands i n hi h T ru os rmum urs acRo MacR be san e M be . y w c . g p occ ( rts o rts 1995b)

E n in the limi sam l xamin for his r n t ha a ar ve ted p e we e ed t p oject, we o ed w t ppe to

nsi s n m r h l i al i f r n s am n Ta linum ula i ns. For xam l be co te t o p o og c d f e e ce o g pop t o e p e , the T. a rvi orum ula i ns di r d r m a h h r in r iz s am n p fl pop t o ffe e f o e c ot e flowe S e , t e num r and s l l n th rela i s am n l n th: in s m ulati ns the i a be , ty e e g t ve to t e e g o e pop o st gm x n s nd the an h rs in h the i m a d th u h e te d beyo t e ; ot ers St g an an ers are the same height . S c local differences have been noted by Reinhard Ware who feel it would be im ra i al and und sira l r niz e su h l al ari ati ns ax n l p ct c e b e to ecog c oc v o t o omical y .

DISCUSSION

The literature on Ta linum and the comm uni ties in which it grows suggests that there is little awareness that Ta linum occurs in east Texas and west Louisiana (Carter Murdy 1985 ) or that sandstone outcrops flori stically comparable to those found i n T n n ss A r ansas and A la am a uarterrnan et a l. 1993 ur i n as T xas and e e ee , k , b (Q , ) occ e t e ui i an a T r - l a li Lo s . e ete e ved Ta num has been known from Lo uisi ana since at least the mid- nineteenth century when Riddell ( 1852 ) included it in hi s Ca ta logus Florae Ludovicia na n ral ui i e. Ce t Lo s ana and east Tex as sandstone outcrops are well known MacRoberts MacRoberts : Talinum in Louisiana 89

plant communi ties and have been the subjects of several floristi c surveys (Mari etta Nix on 1984; MacRoberts MacRoberts

Ta lin um pa rviflorum i s on the Loui siana Natural Heri tage Program ( 1995 ) rare t rare aus its ha i a sali n rai ri s sands n u r plant list . I is bec e b t t ( e p e , to e o tc ops) i s

e x ri d sand ha i at in hi h T. ru os ermum urs is s vanishing . Th e c eep b t w c g p occ al o rare Th s hre lan in Loui siana (MacRoberts MacRoberts 1995b) . e e t e p t co mmuni ties are on the Louisiana Natural Heritage imperiled list (S mi th 1988 ; Teague Wendt

Ta linum rugosperm um has been under considerati on for listing as a federally hran 1 3 z u ie endangered or threatened Species. Coc e ( 99 33) st d d its status and “ is ri uti n and und ha at leas in the n r h rn art of its is ri u i n it i s not in d t b o fo t t, t o t e p d t b t o , immin n dan r x in i n at the na i nal l l but it i s a nti all uln ra l e t ge of e t ct o t o eve , pote y v e b e s aus it curs in nl a s a r d l ali ies r m s of its al ran specie bec e oc o y few c tte e oc t ove o t tot ge , m s s a i ns and u i s r r s ri d ha i a w num rs at s . H e exi sts in lo be o t t t o , occ p e ve y e t cte b t t “ ” u li r ri a r s r a i n s a Thi s las in adds that few populations enjoy p b c o p v te p e e v t o t tus. t po t

a rvi orum and T. ru os erm um a r ha i a ha i underlines the problem: both T. p fl g p f vo b t t t t s

i r una l m s n n uisi ana T. a rv orum i i n r d ra dl . F s are be g dest oye p y o t te y , o t k ow Lo p ifl S te on the Ki satchi e Di stri ct of the Ki satchie National Forest where they are protected The one n n si in ui siana for (MacRoberts MacRoberts 1995 a) . k ow te Lo T - l S in i rugOSpen num is in an acti ve oil field and has a four whee er trail through it. ce t

nl rs a s uar m rs its xi s n i s re ari us . o y cove few q e ete , e te ce p c o

DOCUMENTATION

li uisiana: add Pari sh: MacRoberts Ta num rugosperm um : UNITED STA TES . Lo C o

Ma Robert 2 3 CM . c s 75 7 [LS US], 3338 [LS U] , 33 9 [ ]

T in uisiana: a d Parish: verb 1 8 a l um pa rviflorum: UNI TED STATES . Lo C d o O y 9 [NLU ]; Ma cRoberts Ma cRoberts 3354 Natchi toches Pari sh: B r own 00 LS 8 1 [ U]; MacRoberts Ma cRoberts 2 05 6 [SFRP] , 1 75 8 , 3332 LS U 1 72 6 3 1 B 1 [ ] , , 3 3 [LS US] , 1 75 9, 1 780 [VD ], 33 3 3333

IL I Thoma s 1 072 2 0 1 1 0300 NLU . Red Ri r Parish: ilmo re Sm ith [W L ]; , [ ] ve G S a in 3830 [NLU ] . b e Pari sh: A llen 1 2 8 1 6 [NLU ]; Ma cRoberts Ma cRoberts

335 9 LS U . V rn i nn Pari sh: [ ] e on Pari sh: Thoma s 1 05 079 [NLU] . W Ma cRoberts MacRobe rts 3353 [LS U] .

hil hi s a r is n rn i h r - lea ed Ta linum in ui si ana we did W e t p pe co ce ed w t te ete v Lo , ll s m i n rm a i n on T r hil our co ect o e fo t o exas Ta linum duri ng the course of the wo k . W e T x as r was not h r u h and did not xam in h r ari um ll i ns can e wo k t o o g , we e e e b co ect o , we r r Ta lin um a r vi orum r m n R ert a cR berts epo t p fl f o Newto Co . (Ma c ob s M o 3360 T EX and as r Co . Ma rietta and T ru s erm um r m [ ]) J pe ( 1 1 6 . go p f o a h Ma s . cRoberts Ma cRoberts 3334 TEX 3 335 WI S 333 6 N cogdoc e Co ( [ ] , [ ] , LS U S 33 3 7 W ILLI an d A n r n . Ma R berts Ma Roberts s . n . [ ], [ D de so Co ( c o c he N aco doche i s Co . 1 80 rs r r hi s i s is h r i x n et a l. 9 g S te w e e N o , ( ) fi t epo ted t Spec e Tr m o l ex as . me -93 90 P H Y T O L O G I A February 1997 volu

F i u r e 1 P i s t i l o f T r u o s e r mu m r i h t g . . g p ( g ) a n d T a r v i f l o r u m l e f t S c a l e i s 1 mm . p ( ) . . MacRoberts MacRoberts : Talin um in Lo ui sian a 9 1

u n f T a l i n u m r u o s e r mu m F i g u r e 2 . D i s t r i b t i o o g p

R a n d T a r v i f l o r u m P i n L o u i s i a n a . ( ) . p ( ) 92 P H Y T O L O G I A February 1997 volume — 93

A CKNOWLEDGMENTS

d re S . hran S ar ar R rt i r R . al Th mas as n Theo o Coc e , tew t W e obe K ge , D e o , J o

Si n hurst and Mar Ma field aid d in varrous as e s of hi s r . a i M r g , k y e p ct t wo k D v d oo e

the inn Pari sh salin rairi on Ma 5 1997. The H l ma R ad sit in took us to W e p e y , e p te o e i on anu r 15 1 w fi rs sh n us Nelw n McInn s a 995 . Part of Caddo Pari sh as t ow to by y J y , thi s work was conducted while we were engaged in vari ous volunteer and cost- Share agreements with the Kisatchi e National Forest and the National Forests and

lan s in T x as . S ar ar R r i r and Th r hran Grass d e tew t W e , obe t K ge , eodo e Coc e li r r i n hi s a r commented on an ear e ve s o of t p pe .

linum a rvi r W ur . 85 . S s ma i of Ta o um u . H . M 19 s ar r M. E . B . . C te , dy y te t c p fl N tt

- re olium Pursh P rtula a a . Rh d ra 87: 12 1 1 0. and the ori gin of T. te tif ( o c ce e) o o 3 linum r u e hr an T S . 1993 . S a us and dis ri u i n Ta os rm um H lz . Coc e , t t t b t o of g p o

- a ural Areas urnal 41 . (Portulacaceae) . N t Jo W W a in W . Ma un . . Kil ri . H . G. Mart J . nr 1980. Ed ards J . P P . w , , go , p t ck, C e y

il S ur ve o addo P a rish Lo uisia na . S il ns r a i n S r i U . S . So y f C , o Co e v t o e v ce , ri ul ur Department of A g c t e . d r wn Illustrated Fla m l n H . A 1 68 . The New B ritton an B o o the G easo , . 9 f

i e ta te and . V l 2 . Ha n r Pu li shi n Northeas tern Un t d S s Canada o . f e b g Co New

r r . Yo k, New Yo k

oui siana atural H ri a Pr ram . 1995 . Rar lan lis . n u li sh d r r L N e t ge og e p t t U p b e epo t ,

ouisi ana ar m n il li and Fi sh ri s a n R u ouisi ana. L Dep t e t of W d fe e e , B to o ge , L

MacRoberts D .T . 1989 . A um n d hec li s and a las of the as ular ra , doc e te c k t t v c flo of

Mus um if S i nc s . . oui siana S a ni rsi uisiana. ull. 9 Lo B e of L e c e e , No L t te U ve ty ,

Shr r uisi ana . evepo t, Lo

M . H . MacRoberts. 1 a. rth as ular lan MacRoberts B . R. 995 , Notewo y v c p t

- ll i ns on the Kisa hi a i nal F r s ouisi an a. Ph tolo i a 3 13 . co ect o tc e N t o o e t, L y g

M . H . MacRoberts . l 5b . Fl ri s i s x ri an hi ll MacRoberts B . R . 99 s s in , o t c of e c d

- i i ana . Ph l ia 2 northwestern Lou s yto og 79: 1 3 13 1 .

MacR R ac . 1 V as ular r an obert s M .H . B . . M Roberts 993 . a s ds n ut r , c flo of to e o c op mmuni i n rn uisi ana i h n rar es i s n s o and n r h s i s . co t we te Lo , w t ote e otewo t y pec e Ph l i a -4 yto og 80.

- Mari a . L i x n . 4 V a i n n a n rairi li u K . E . S . 198 . o mm ni in ett , N o eget t o ope , p e ke co ty

- as rn x . x i 2 e te Te as Te as Journal of Sc ence 3 .

i x n E . S K L . ari c . ra ch r . M a M. M Crar 1980 B el t um ere tum and N o , ett , y , y y c linum ru os erm um new s i s x s n h en l r Ta T as and n o Sc o o i ion wr i htii . g p , pec e to e ote g i - S da 35 6 .

uart errn an E . M . P . B urbanck D .I . S hur . 1993 . R u r mmuni i s: Q , , , e ock o tc op co t e im - W i n s n S an s n and rani . P . 35 85 . In : . H . Mar S .G. L e to e , d to e , G te p t , Boyce ,

A . C . Echtem acht i di ver it h u hea rn hn B o s y of t e So t ste Un ited Sta tes . Jo il S ns r r . W ey o , New Yo k , New Yo k Ri ll J .L . 1 85 2 . Catalo us Fl ra u i ana rl i . ans Med. ur . urnal dde , g o e L dov c New O e S g Jo

- 754. MacRoberts MacRoberts: Ta lin um in Louisian a 93

R i nhar R. T . S . ar . 1 98 9 . A a ati n su s rat in r u r lan s : e d , W e d pt o to b t e ock o tc op p t la - hl an Ta lin um P r u ac a . Gaz . 1 z44 4 . i nterior hig ds ( o t c e e) B ot. 50 9 5 3 i n n S i n hurst J R . 199 6 . The s a us n an r lan s as T x as: Hi s ri al g , t t of e e d ge ed p t of e t e to c ,

h l i al n s. M . S . i l i al and h s s S h n F. A us in S a eco og c , p yto og c ote t e , tep e t t te

ni rsi a h s T x as . U ve ty , N cogdoc e , e l mmuni i s f ui a i M . 1 88 . The na ura o sian . n u l sh r S mi h L . 9 r . t , t co t e Lo U p b ed epo t ram a n R u ui si ana a ural H ri a Pr u o isian a . Lo N t e t ge og , B to o ge , L 4 a and i ar i T a u . T . n . 199 . ss r sh s LA : na ural ar as e g e , J We dt C ddo Bo e p e , t e

li h d r . The a ur ns r an a n R u u n u s r isiana . survey . U p b e epo t N t e Co e v cy , B to o ge , Lo al lan lish a Pr ram . 1995 . S i lis . n u Tex as Natural Heri t ge og pec p t t U p b ed report .

T x as a ural H ri a Pr ram A us i n T x as . e N t e t ge og , t , e

T x as r aniz a i n for En an r S i s . 1993 . En an r hr a n and e O g t o d ge ed pec e d ge ed , t e te ed

a ch lis s of T x as lan s . Pu li a i n 9 . A us i n T x as . w t t e p t b c t o t , e w Ta linum P r ula a a r m the ar lad s of mi 1 67. A ne l ar S . 9 W e , ( o t c ce e) f o ced g e dd e -4 Rh ra 75 . Tennessee . odo r 7 — P hytolo gia (Februa y 199 ) 102 .

NEW S P ECIES OF CA LA TH EA (MA RA NTA CEA E) FRO M EA S T ERN B RAZI L

1 2 H elen Kennedy Joao Marcelo

1H r ari um an artm n ni rsi ri i sh lum ia e b , Bot y Dep e t, U ve ty of B t Co b

V an u r B . C . V 6T 2 B 1 A A A co ve , C N D

2 H r ari S a de ani a Si st mai a ardirn ani do Ri o de aneir o e b o, ec o Bot c e t c , J Bot co J - Rua ardi m an i CEP 2 2 460 Ri o an ir R. RA S I J Bot co , 000 de J e o , J B L

AB STRA C T

Ri o an ir and . a n n a e Ca la thea fa ti ma e from northern Edo . de J e o C and

r a i ela e r n in ulti a i n are d s ri d as new . All hr s ci s C. g z , g ow c v t o , e c be t ee pe e

in s i n B revisca us . Cala thea a tima e i s r la d . ebrina are C . ect o p f e te to C z ; C .

l s l r la . l u a e and . albertii and r ielae to annae is most c o e y e ted to C o is C ; C. g az

a emula . C.

KEY R S : Ca la thea Maran a a raz il ax n m h rti ul ur WO D , t ce e , B , t o o y , o c t e

MA E H . nn. Mar l e n ur CA LA TH EA FA TI e s . o v. Fi l . T PE: Ke c o, p c ( g e ) Y a i Marx arra de uara a M RA IL . ul i a d t Si R r url i uni Rio B Z C t v te t o obe to B e , B G t b , c .

- an ir Edo . Ri o an ir raz il a ssi n num r MA R 101 r m de J e o , de J e o , B , cce o be , f o hi z m s ll d Fatima omes de r G Sou a r m M rr do ca . 40 km o e co ecte by z f o o o Coco ,

n r h the i am s Muni . do am s Edo . Rio de an ir raz il o t of c ty of C po , c C po , J e o , B ,

r ul i a i n 1 1 an . M. de F me e d in J 199 1 H . Kenned Go s d ou a flowe e c t v t o , y . S z 4700 H T PE: RB Is s: K S itio RB M UB C . ( OLO Y ; otype , , )

- Plan a m al a . F li a asili a 1 3 caulina 0 a a vel a a t t o b , , ov t obov t , usa cum acumi ne asi a u a vel usa su ra viridi a la ra s us costarn obt , b c t obt , p g b , ec allid viri dia minu m n sa subtus ur ur a la ra ul inus ars p e , te to e to , p p e , g b ; p v Sp e minu tomentosus ad 10 cm l n us i lus iri is interdum ru - ti nctus te o g , pet o v d fo , mi nu tomentosus 30- 5 cm l n us a ina minu m n sa 35 - 65 cm te 7 o g ; v g te to e to ,

l n a . S i a 1 sub lobosa vel ovoida ad x cm edunculo mi nu o g p c , g , p te m n s 5 0- 68 cm l n l a a bracteae ad 40 S irali ter di s ositae to e to o o go e ev t ; , p p , cori aceae r ss ovatae vel la ovatae a i mar s n i us vel , dep e e te , p ce ce ce t , obt o rotundato virides ad resse tomentosae ari a rum ad 10 vel ul ra , , p , p flo t , prophyllum et bracteae secondari ae dense mi nute tomentosis apicibus m arcescentibus bracteolis 2 membranaceis ari um al um la rum s ala , , ; ov b , g b ; ep m m rana a lli i a rotundata ad a icem i la a ad 1 1 mm l n a e b ce , e pt c , , p v o ce , o g ;

96 P H Y T O L O G I A February 1997 volume - 102

lea es numer us s irall arra n ed ra s and the re u n r en i n of sheen of the v , o p y g b ct , f q e t et t o unpollinated flowers in the inflorescence which turn black upon drying . Ca lathea fatima e i s a very stri king foli age plant because of the contrast of the pale midrib regi on r l r It l arl has hor i ul ural with the dark green above and the pu p e unde leaf. c e y t c t potential as a landscape plant in the wet tropics and as a house plant in the temperate The c r lla and s ami n d s urn la u n dr in or hen reser ed i n 70% regi on. o o t o e t b ck po y g w p v '

S same rea i n i s s n in Ca lathea ca tata Ruiz Pav . i n l . hi h ETOH . IT1i ct o ee pi ( ) L d w c likewise Shows a color change m the style after trippin g .

n r a Faima m s Calathea fatimae is named in ho o of S ra . Mari de t Go e de S ouz a aniz a i n d um n a i n and u h ri n the li in ll i ns whose org t o , oc e t t o , vo c e g of v g co ect o , all Maran a eae and su r i si n the h r ari um for Funda a a i nal Pro i y t c , pe v o of e b c o N c o Memoria Slti o Roberto Burle Marx has been invaluable to both botani sts and late R r url Marx re u st d ha his s i s hi h she had horticulturists . The obe to B e q e e t t t pec e , w c

in her h n r and als mmissi n d a ain i n of it Mr. Mar collected, be named o o o co o e p t g by k

ai n in is at Si i REM. Fothergill . The p t g t o

Mar el e . no v. i ur A A NA E H . nn . s F T PE . CA LA TH E N Ke c o , p c ( g e Y

ti vated at Si i R r url Marx arra de uara i a Edo . Ri o de an ir O rl t o obe to B e , B G t b , J e o ,

- n r MA R 1 r m raz il Edo . unkn n 1 1 J an . 1 raz i l a essi n um 99 1 H . B , cc o be , f o B , ow , , Ga mes de Sou a 4696 H T PE: RB Is t : Kenn e M. de F. s dy z ( OLO Y ; o ype K ,

M . RB ,Siti o RB ,UB C)

an a cauli na m al a r u n er demum rami ficans . F li a asali a Pl t t , f eq e t o b 0- 1 caulina 1 -4 h r a ea a a interdum elli i a usa cum acumi ne , ( e b c , ov t , pt c , obt , data vel r un a a ad 5 3 x 2 3 cm su ra ni ida sa ura viri di a base rotun t c t , , p t t te , la ra subtus ur ur a la ra ul i nus olivaceus vel ur ure -ti nctus g b , p p e , g b ; p v p p o , minu tomentosus ad cm l n us i lus atroviridi s interdum ru te , o g ; pet o fo tinctus mi nu e tomentosus ad 48 cm l n us a ina minu m n sa in folii s t o g ; v g te to e to ,

- - caulinis 10 2 1 cm l n a . S i a 1 3 elli soideae vel subc lindricae o g p c e , p y cm lon ae edunculo - 15 cm l n bracteae 10-30 S ira liter dis ositae g , p o go; p p herbaceae reniform ae vel transverse ellipticae apice rotundato cum acumi ne vel us recurvato viri des cm lon ae ari a florum ad 8 bracteolis 1 obt o , g ; p , ; ovari um album glabrum; sepala anguste ellipti ca vel anguste obovata rotundata; r lla et s ami n i a cremea u us corollae ad 2 4 mm l n us l i an us co o t od , t b o g ; ob g te elli tici a u i vel obtusi starrri nodium x t ri us la lli i um mar ina um p c t ; e e te e pt c e g t , n u al n - stami odi m c losum oblo go obovatum .

- - aul s n h r cm hi h . Fl ri n h s ari n 0 4 5 l a s 0 C e ce t e b g owe g S oot be g ( ( ) e ve , 1 basal and (0 l (- 2 ) cauline on the mai n stern; axillary shoots formed in the ax il of the l a su ndin the in r s n ar 2 - l a s and an n n e f bte g flo e ce ce be 3 e ve i floresce ce . Up to 3 axillar infl c a u i orescen es m r d in hi s m an n r . a a h lls s hi n d y y be p od ce t e C t p y t ff, t cke e asall narr l a a x 4x r und a i ula ur l mi nu l m n s l . b y , ow y ov te, pe o ed , p c te, p p e , te y to e to e ( ) ea la h r a us a asi nall lli i a x us i h a um n a u L f b de e b ceo , ov te , occ o y e pt c , pe obt e w t c e to c te in small r l a s as r un run a r h r l a t nua 18 - 5 3 x 10 e e ve , b e o ded to t c te , ve y S o t y t e te , ( (

- - 13 2 3 cm . ea la a hin r n la r us mi ri li r n L f b de bove S y deep g ee , g b o , d b o ve g ee , la r us l r l af sur a s mi - hi n ur l la r us mi ri li - r en asall g b o ; owe e f ce e S y p p e , g b o , d b o ve g e b y , ur l a i all s ars l mi nu l a r s x p p e p c y , p e y te y pp e sed tomentose ( l 4 m agni fication) . Pul inus lli i 1n r ss- s i n li - reen or in i h ur l ll n v e pt c c o ect o deep o ve g t ged w t p p e , ye ow gree us at un i n i h i l mi nu l a r ss m n s l4x n arl n u n j t j ct o w t pet o e , te y pp e ed to e to e ( ) , e y co fl e t Kennedy Marcelo: New Ca la thea from eastern Braz il 97

i th i l ul i nus sli h l lar r i n di am r cm l n P i l ar w pet o e , p v g t y ge ete , o g . et o e d k ish r n minu l a r ss m n - r en r n s 48 cm l n . ea g e to b ow g ee , te y pp e ed to e to e , o g L f h a h not auri ul a dar r n r ni sh mi nu l a r ss d m n s sh a h S e t c te, k g ee to b ow , te y pp e e to e to e , e t - m l n h r - of aulin su ndi n l a 2 1 c s 1 5 5 8 cm l n . S m dar c e , bte g e f o g; ot e o g te k n in d ur lish mi nut l a r ss m n s 14x l not s n intem e gree t ge p p , e y pp e ed to e to e ( ) , fe t ee ; od n su n in l af and n x l r se n l a 14- 71 cm intem ode n betwee bte d g e e t owe ( co d) e f , betwee

- nd and hi r l a s h n r s n 30 5 1 cm l n . seco t d e ve , w e p e e t, o g

In florescences 1 - 3 1 rmi nal on m ain h a i i nal inflorescences rmi na , te S oot, dd t o te ting the 1 - 2 ax illar sh s rar l the in r s n rn on a s ara non- l f h y oot , e y flo e ce ce bo e ep te ea y S oot la l h a h lli s i u lindri x - su nd d a ss s 3 5 cm . bte e by b de e S e t , e p o d to bcy c , P dun l r n i h r al n one si mi nu l a r ss d- m n s at as e c e g ee w t g oove o g de , te y pp e e to e to e b e ,

- - ars l so a su la r us in a i al hal 15 cm l n . ra s 10 30 Sp e y bove to bg b o p c f, o g B ct , s irall arran d h r a us l r n s r ni rm ran s rs lli i a x r un p y ge e b ceo , owe o e e fo to t ve e e pt c , pe o ded i h a um n u r ra s lli i a x us the a i al mar in r ur x w t c e , ppe b ct e pt c , pe obt e , p c g ec ved ,

cm a h su ndin u 8 r airs. u r sur a of ra shi n r n ; e c bte g p to flowe p O te f ce b ct y g ee , asal minu l m n s 14x ma ni a i n ra uall ars r a and b te y to e to e ( g fic t o ) , g d y Sp e bove

la r us n ar a x inn r sur a hin li h r r n la r us. i ari na r h ll g b o e pe , e f ce S y g te g ee , g b o B c te p op y m m ran us a a x r und d ranslu n Chartr us r n mi nu l a r ss e b o , ov te , pe o e , t ce t e e g ee , te y pp e ed m n s on sid s n r and arina la r us x cm to e to e e , ce te c g b o , , n S ndar ra m m ran us a a x r cm id arina ari a. und w e , c to c eco y b ct e b o , ov te , pe o ed , ranslu n Char r us minu l a r ss d m n s at sid s and a x x t ce t t e e , te y pp e e to e to e e pe , l r ai m ial ari na la r cm . ra 1 er r us cm l n B cteo e p flowe p , ed , c te , g b o , o g .

Fl rs n h d r m in r s n if not lli na d. S als h r a e us owe Ope , S e f o flo e ce ce po te ep e b c o n ral l mar ins h ali n narr l lli i narr l a a x r und hi ce t y , g y e, ow y e pt c to ow y obov te , pe o ed , w te

- - at as a i al Char r us la r us 14 16 x 4 5 mm . r lla and s amin s b e , p c t e e , g b o , Co o t ode r am -c l r d la r us r lla 32 - 36 mm l n u 2 2 - 2 4 mm l n r lla l s c e o o e , g b o ; co o o g , t be o g; co o obe 1 1 - 12 x 4- su ual narr l a narr l lli i a u to us ca . 6 mm . beq , ow y ov te to ow y e pt c , c te obt e ,

- - u r s ami no r a l lli i mar ina 12 14 x 9 1 1 mm . all s s amin O te t de b o d y e pt c , e g te , C o e t ode

- l n a a i al 3 mm al i the r s all s ca . 12 x 6 mm . O rcullate ob o g obov te , p c pet o d, e t c o e , ami i a x 4 a al i a s n hi at t c . 6 mm . S am n i h l ral nda t ode w te p, t e w t te pet o d ppe ge to n i n S l and s i ma r am mm wide exte d g to basal of anther; anther 2 mm long . ty e t g c e

l r ar sm h hi la r u s. a sul un n n . co o ed . Ov y oot , w te, g b o C p e k ow

a la the al n s i n B revisca us and i s s l s l r la C a anna e so belo gs to C. ect o p mo t c o e y e ted

lo ui a e a n . an d i a n . i u i . W . l n a lbert i I. . ail R ffill s o P rs to C s G g ep C. B ey ( y C g a z ete e

non t liet ei E . Mort en . The a I i s somewhat less closely related to C. z bove species all have wider than long spirally arran ged green bracts; herbaceous green sepals; cream - whi te corolla and staminodes with an apically petaloid callose stami node and l a s ur l n a a anna e i in i sh r m . lo uisae and . e ve p p e be e th . Calathe i s d st gu ed f o C C albertii the ar r n un attem ed l a s the all r s a ur m vs. 111 by d k g ee , p , e ve ; t e t t e hi gh) ; the development of axi llary Shoots bearing additi onal inflorescences i n the ax il f h o t e l a su n i al r n or hi ra s. e f bte d ng the i nflorescence; the deep green vs . p e g ee w te b ct The l m n deve op e t of axillary shoots (and addi tional inflorescences) i s shared with C .

- - liet ei . The un atte v 3 7 r a il z p m ed leaves and more numerous bracts ( 10 30 s . ) e d y i i n i s u sh it r m . Iie d t g f o C tz ei .

Ca la thea anna e is nam in h n r i anis A nna an i a the musi al ll a u ed o o of p t C d d , c co e g e of the la R l M a a he h n r r ur arx t his r u s h s so . H e te obe to B e , eq e t t t be o o ed 98 P H Y T O L O G I A February 1997 volume

Fi . l . Ca la thea mi n ute A . H . bi g a t . B . Infl r / o escence.

C . Flower on u er leaf surf ace. Measurem ent in cm pp . Kennedy Marcelo: New Ca la thea from eastern Braz il 99

Infl o rescence . Fi A . H a bit . B . 2 . a la thea a n na e . g . C

w r m ent i n cm . Fl l f rface . Measu e C . o er o n u pper ea su — 102 F ruar 1997 lum 100 P H Y T O L O G I A eb y vo e

Infl orescence with it . B . A . H ab r iela e. i 3 . a la thea a F g. C g z

w wi h mm ruler on u er leaf surface. w . Flo er t flo ers . C pp

Measurem ent i n cm . Kennedy Marcelo: New Ca la thea from eastern Braz il 101

in anna e Mr . Mar F h r ill hi h commi ssioned a watercolor paint g of C. by k ot e g w c i s at

S iti o REM .

. nn. Mar l e . n o v Fi ur RA ZIE LA E H s . T PE: CA LA TH EA G Ke ce o , p c ( g e Y l a t S i i R rt ur l Marx arra uara i a Muni I . u i a . Rio BRA Z L C t v ted t o obe o B e , B de G t b , c ir raz il r m rhi z m s ll in raz il E an i r Edo . Ri o an do . de J e o , de J e o , B , f o o e co ected B , d i a i n 1 J an. 1 1 H . Kenne M. de F m un n n r d in ul 9 99 . o es k ow , flowe e c t v t o , y G M iti RB M de ou a 4699 H T PE: RB Is s: K O S o UB C . S z ( OLO Y ; otype , , , )

- 4 u a i sa ad 0 cm al a . F lia as alia 3 in u urcul Planta c esp to 9 t o b , q oq e s o ri n i ali 2 - 3 i n u u surculo ax illari a a breviter a umi na a vel usa p c p , q oq e , ov t , c t obt a mi no as rotundata vel usa r issim an us a a ad 42 x 2 2 cum cu , b e obt , b ev e g t t , cm su ra smara di na S lendentia la ra subtus nitentia i ri i - ri s a ur ur , p g p g b , v d g e p p eo i vel n ul sa la ra ul i nus s ars mi nu tomentosus cm p cta eb o , g b ; p v p e te longus; peti olus obscure brunneo- vi nosus mi nute tomentosus ad 3 1 cm

- - l n u a ina s ur brunneo vi nosus mi nut m n sa . S i a 1 3 o g s; v g ob c e e to e to p c e , elli soideae vel lindri a 4- 1 1 x cm edunculo veluti no 1 1 - 18 cm p cy c , p l n bracteae 5 0- 12 0 S iraliter dis o sitae herbaceae r ss obovatae o go; , p p , , dep e e , obtusae cum acumine a i recurvato viri des interdum ur ur - ictae p ce , p p eo p , velutinae ad x cm se ala mem rana a a a vel lli i a , ; p b ce , obov t e pt c , usa la ra ad 13 mm l n a corollae aureae labrae u us 38 -45 mm obt , g b , o g ; , g , t b l n us lobis elli ticis ad 1 3 mm l n is s ami n ia aur a s amin dium o g ; p o g ; t od e , t o x rius a um mar ina um ad 13 x 10mm s ami n ium ad a icem vix e te obov t , e g t , , t od p

etaloideum ad 10 x 5 mm ari um la ru m . p , ov g b

R sula h r m hi h h s d ns l lus r d mai n sh ari n 3 -4 o te e b , g , S oot e e y c te e , oot be g basal leaves with ax illary Shoots beari ng 2 - 3 leaves developing in the ax ils of th e upper

- l . a la h r a us 2 3 l a s . a a h lls narr l a a i ulat ur o e ve C t p y , ow y ov te , p c e , p p e Le f b de e b ce , a a x us i h a um n h r l a uminat ase use r un ov te, pe obt e w t c e to S o t y c e , b obt to o ded , a l sh a umi na 2 -42 x 1 - 2 2 la a s ar lin dar r en ru rt 3 3 cm . eaf b pt y o c te , L b de bove p k g k g e i h mi ri al to hi ti sh r n la r us l a sur a l s mi - shi n li h r w t d b p e w g ee , g b o ; e f f ce be ow e y g t g ey r n irr ularl s r a and in i h ur l la r us mi ri in ur l g ee , eg y t e ked t ged w t p p e , g b o , d b t ged p p e ard as a i all ll - r n la r ul inus r adl lli ti i n r ss us. P tow b e, p c y ye ow g ee , g b o v b o y e p c c o s i n sam diam r as i l dar mar n asi nall i h a li h r n lin ect o , e ete pet o e, k oo , occ o y w t g t g ee e al n r n s ars l mi nu l a r ss d m n s m r s ars l so al n a o g f o t , p e y te y pp e e to e to e , o e p e y o g b ck ,

- cm l n . P i l ar ur l r n ars l minu l a r ss m n s o g et o e d k p p e b ow , Sp e y te y pp e ed to e to e s x - l - r n mi nu l n at 14 6 3 1 cm l n . a h a h not auri ula ar ur ( ee ) , o g Le f S e t c te , d k p p e b ow , te y a r ss m n s hai rs rn on a mi nu ushi n of lls hai rs l n r and m r pp e ed to e to e , bo e te c o ce , o ge o e d nse ard as i si l the na 18 -40cm l n e tow b e , v b e to ked eye, o g

Inflorescences 1 - 3 er sh the rs one rmi nal on the main sh a i i nal ( ) p oot , fi t te oot , dd t o n s rminal on la ral sh s im ri a lli s i li n ri 4- 7 - l l x o e te te oot , b c te , e p o d to cy d c , ( )

cm . P un l r n s r a or tin i h ur l a i all or ur l hrou h u ed c e g ee t e ked ged w t p p e p c y p p e t g o t , ns l l t m - 50- 12 0 n s leS S ns l so n ar as 1 1 18 cm l n . ra s de e y ve ve y to e to e , de e y e b e , o g B ct s irall arran r ss a a x us i h a um n u r mar in p y ged , dep e ed obov te , pe obt e w t c e , ppe g and a x r ur x cm the rs and s n ra s the pe ec ved , , fi t eco d b ct i s cm i a h ra su n in u 2 or m r r airs. u r w de t , w de; e c b ct bte d g p to o e flowe p O te sur a of ra s li h r n l rm s ra s ma i n i h ur l l f ce b ct g t g ee , owe o t b ct y be t ged w t p p e , ve vety m n s inn r u i ari na r s r a shi n al r n la r us . B h ll to e to e; e f ce y , p e g ee , g b o c te p op y 102 P 11 Y T o L 0 G I A February 1997 volume - 102

m m rana e us a e lli i a x r und d asi nall r us ranslu n al e b c o , ov t to e pt c , pe o e , occ o y et e , t ce t p e r n a i all l rl ss l la r us at as u r hal s arsel il s al n g ee p c y , co o e be ow , g b o b e , ppe f p y p o e o g 4- m cm i na a c i ar ari n . S n i s x l . 2 . l ar s de , ( , w de , c to c eco d y ra m m ranac us a a x r un al ranslu en r n a i all la r us b ct e b eo , ov te , pe o ded , p e t c t g ee p c y , g b o

at as u r hal s ars l il s x cm . ra l s 1 er r b e , ppe f p e y p o e , B cteo e p flowe ai r medial m m rane us han n led lin ar ran s ar n la r us p , , e b o , c e , e , t p e t, g b o ,

cm long .

m ran us a lli i us rans ar n la r us 1 1 - 13 x 4 Sepals me b o , obov te to e pt c , obt e , t p e t, g b o ,

- lla and s ami n d s ri h l n ll la r us. r lla u 38 45 5 mm . Coro t o e b g t go de ye ow , g b o Co o t be

- l n l s su ual lli i a u 1 1 1 3 x mm . u r mm o g; obe beq , e pt c , c te to O te

- - s ami n at mar i na 10 13 x 8 10 mm . all s s ami n a x t ode obov e , e g te, C o e t ode , pe - i reflexed 8 - 10 x 4- un uall 2 l d nl the a i al mm al 5 mm . eq y obe , o y p c pet o d , ,

- - ulla stami n d 5 6 x 3 4mm . S l and s i ma ll . ar sm h r am Cuc te o e ty e t g ye ow Ov y oot , c e

l r la r us ca. 3 x 2 mm . a sul un n n . co o ed , g b o , C p e k ow

tru r aemula Ca la thea graz iela e is most Simi lar in i nflorescence and floral S ctu e to C. hari n the s irall arran r n us ra s i h r ur a i al mar in Kom . s , g p y ged , g ee , obt e b ct w t ec ved p c g ; d a al i all s s ami n yellow corolla lobes and staminodes; an pically pet o d c o e t ode . Calathea

ra ielae i s r adil i s in uish d r m . a emula its lain r n un attemed g z e y d t g e f o C by p g ee , p , l a v a t rned i h a li h r ll i sh 2 -4cm i r n and al n the midri e ves s. p t e w t g te ye ow w de g ee b o g b;

the r l i n d la r us vs. li h r en l t m n s l r l a sur a and pu p e t ge , g b o g t g e , ve ve y to e to e owe e f f ce

l h r n vs. r en i l and l a sh a h . The rs in r the deep purp is b ow g e pet o e e f e t flowe C. g adela e are ll x rted i h at l as 1 cm of r lla u id n nd the ra s h r as we e e w t e t co o t be ev e t beyo b ct , w e e

la e u i s n ain d i hi n the ra s. al hea ra iela e li in . aemu th C at . C t be co t e w t b ct g z , ke C

a tima e is an a ra i ard n rnam n al . The s ri i n n ras of the shi n lim f , tt ct ve g e o e t t k g co t t y e green i nflorescences and yellow flowers against the background of the dark maroon peti oles combined with the di stinctive whitish mi dri b of the blade above and the purple r d n i t r St eaki ng an mottli g below g ves i g eat potenti al for horticulture .

raz iela ar s n a n l Thi s species i s named i n honor of Dra . G B o o i ck ow edgment of her long devotion to the study of Braz ili an plants and her numerous contri butions to our n l h m hr u h her a hi n and num r us u li a i n k ow edge of t e t o g te c g e o p b c t o s.

A CKNOWLEDGMENT S

S cial ra i u s e a R pe g t t de goe to th l te S r . oberto B urle Marx for the opportuni ty to r i h his x nsi Maran a a ll ti n and for the man i s ussi ns a u wo k w t e te ve t ce e co ec o , y d c o bo t h m r the ar . h t e ove ye s T anks to Dra . Dorothy Dunn de Araujo for her continui ng assis an h at RB and la r GUA and for hari n he a a i n t ce bot te S g r ccommod t o s; and to Dra . Graz iela ar s for r i in a ili i s in the h r arium at RB r l r B o o p ov d g f c t e e b fo fie d wo k in 197 1 .

han S ra . Faima m s for her h l at the Si i and for r s r i n um n i n We t k t Go e e p t o p e e v g , doc e t g and u h ri n the su r li in ll i n h r Elain H m hr Dr . u and Ms. Een vo c e g pe b v g co ect o t e e , e p ey Ten for h ir h l i h the illus ra i ns and Mr t e e p w t t t o s . Margaret B utschler for her

n ri u i n ar a har s . han the ura rs of GU A MO RB co t b t o tow d p ge c ge We t k c to , , NY , , and US for the l an or use of h ir ma ri al . ish han UBC , o t e te We w to t k Mr . B ruce H olst

and Dr . Fr an rs for h ir r i s and h l ul mm n s and Dr an ed G de t e ev ew e pf co e t . G ders for

su r H . nn ppo t to Ke edy .

104 P H Y T O L O G I A February 1997 volum e - 104

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R .L. H ar m an . 1996 . R lassi a i n of rth M . A . Lane , t ec fic t o No

Ha lo a us m si a : Astereae m l : Ra a cks on ia en . nov . p p pp (Co po t e ) co p eted yj g A mer.

- 370. J . Bot . na i n n ria L. 1 8 . m i s i Ericame m si a : te s m G. 9 9 As reae Ne o , New co b t o (Co po t e ) . 6 Phytologia 10 .

h . R M r an . i . L . . S u D . B B . S m s n 1 90. X l tha s m G. 9 o mia Ne o , , Y , o g , p o y terea e a n nus r la Eu ha mia Si a - A s ra a : A s . 1 16 . ( te ce e ) , ew ge e ted to t d - P hytolo gia (February 1997) 106 .

A new commun ion m MA CHA ERA NTH ERA CA RN OS A (A S T E RA CE A E : A S T E R E A E )

s m Guy L . Ne o

f r En ir nm n al S udi s S am H us n S a ni rsi Tex as Regional Institute o v o e t t e , o to t te U ve ty ,

H un s ill T x as 77341 U . S .A . t v e, e

A BSTRA CT

sa m ri s s all a ri ari i s ll in the Machaeranthera com o co p e two op t c v et e , fo ow g i al arie i s r m Ari z na and S n ra S S und r . The study of cott be g typ c v ty f o o o o ,

The ari r m ali rni a and a a i s re niz d as M . Mexi co . v ety f o C fo Nev d cog e

b no . m om . v i ntrica ta A . ra s a var . ca m os ( G y) Ne o , c

: Macha eranthera A stereae A s rac a n m n la ur KEY WORDS , , te e e, o e c t e

ra la The generic sta tus and infraspecific tax onomy of As ter intrica tus (A . G y) B ke ra r n r s udi d S un r who has Leuc osyris co m osa [A . G y] G ee e) we e t e by dbe g

r i ai ls m r h l a h and n m n la ur . H e n lud d ha the p ov ded det of o p o ogy , geogr p y , o e c t e co c e t t Single Species should be maintained as the monotypic genus Leucosyr is Greene but later decided that a positi on in the genus Haza rdia Greene would be more appropri ate . The transfer of A ster in trica tus to Macha eranthera NeeS (Nesom 1989) reflec ted yet an other point of view .

S undberg recogniz ed tw o allopatri c elements wi thi n the species: (a) a system of l - v i s of s u h as rn A riz na i h an ar h a lan s ar . ca m osus in ur un ge e ded p t ( ) fo co t e o t e te o , w t a a n l ali i n S n ra Mexi co l s th e in rna i nal r r and a s s m dj ce t oc ty o o , , c o e to te t o bo de ; (b) y te small- h a lan s var r ta h un i s s u h rn ali rni a and ur of e ded p t ( . in t ica ) in eig t co t e of o t e C fo fo l r m a s u h rn a a . He n ha var . omosa is n n n of o t e Nev d oted (p . 197) t t c k ow o y f o few “ ” l i aus unsu c ss ul a m s co lect ons and may be threatened with ex tin ction . Bec e of c e f tte pt re l a lan s at r i us ll i n si s he ma ha sus ha the nti to oc te p t p ev o co ect o te , y ve pected t t e ty alr a is i n e dy ex t ct or nearly so .

In the rans r Ma ha eran thera I rr to S un r for hi s h i of n ra t fe to c , defe ed dbe g c o ce ge e for the ari al S in h n h r r n uis 1 4 main ain the m ina i n . 99 v et co b t o ce t e , oweve , C o q t ( ) t ed i s i hi n Leucos ris and ma the n essar ari al om in a i n i n h a nu s Spec e w t y de ec y v et c b t o t t ge , ’ reflecti ng hi s agreement with S undberg s assessment of i nfraspeci fic vari ati on and hi s ini ial h i t c o ce of genera . 106 P H Y T O L O G I A February 1997 volum e - 106

A ra m Machaeranthera com osa ( . G y) Neso has not been included i n recent

l ular s udies of the ri Astereae an et . a l. 1996 or the u ri mo ec t t be (L e, ) s bt be Machaerantheri nae (Morgan Simpson 1990) to provide a test of the competing

h l al h h ses of n ri la em n . The s i s has h r n morp o ogi c ypot e ge e c p c e t pec e , oweve , bee accepted within Machaeranthera by Hartman ( 1990) and in the treatm ent of that genus ’ rni a il r n In Cron uist s r a m n of the i s 19 4 in Cali fo by Ke B ow q t e t e t Spec e ( 9 , “

2 60 as Leucos ris he mad the ll in s r a i n . For the as n ur p . , y ) , e fo ow g ob e v t o p t ce t y as usuall n in lu in A ster hi h it ars no r l Leucosyris h y bee c ded , to w c be ve y c ose i l t i s alli d Ha lo a us s inulo sus hi h has r similar relati onshi p . Poss b y i e to p p pp p , w c ve y

H a lo a us s inulosus Pursh . was rea Turn r achenes and pappus . p p pp p ( ) DC t ted by e n n m Macha eranthera inna ti da Shi nn rs sensu la to H artman ( 1976) as a sy o y of p fi e .

Following preliminary agreement that the species i s correctly placed in ’ M ha eranthera ha eranthera i h its rea s similari m m rs of ac s . A n da Mac , w t g te t ty to e be ect

artman s m 1989 Hartman 1990 s m et a l. its rm al R.L . H (Ne o ; ; Ne o , fo taxonomy is completed here with the vari etal combination .

m var R ra s . intrica ta A . r MA CH A E RA N TH E RA CA N OS A (A . G y) Ne o ( G ay) '

M: B i elo ia intnca ta A . ra P s m omb. no v. AS I N ro . A mer Ne o , c B O Y g v G y , c . l ad. Ar s 1882 . Aster intr ica tus A . ra a . ash . A ad. A c t ( G y) B ke , J W c S ci .

1 37 Leucos ris omosa A . ra s m var . mm 9 . y c ( G y) Ne o (A . Gray)

4. m . r n . Inter . Fl 199 C o q ,

V 4 Inte un ta in lora . m r n uis A . l The 199 . rm o F u Fi : A s ral s. r C o q t, o e ve te e New Yo k

ani al ar n r nx r . Bot c G de , B o , New Yo k a f ac r H ar m n R.L . 1990. A ns us o M hae anthera As ra a : te t , co pect ( te ce e A s reae) . Ph ol ia - yt og 465 .

- K. r r il D .J . 1 & G. n . 993 . Machae anthera . . 3 n : P 08 3 10 I . Hi man Ke , B ow p , J ck ’ Manua l the Flora J e son s o o a li ornia . ni . ali rn i a P r ss p f f C f U v C fo e ,

r l ali rnia. Be ke ey , C fo A D an M . . .R . M r an . S uh B . B i S m s n R. K. ans n L e , , o g , Y , . p o , J e . 1996 . R la i nshi s of r h Am ri an n ra of A stereae as d on hl r las DNA e t o p No t e c ge e , b e c o op t

r s ri i n si a a . In : Hi n D . 1 . N . H . J . B eent e om ositae: e t ct o te d t d , j C p i - tema 49 P ntem tl. S s t s . I m 4 77. r . . n . 199 . y c , pp oc Co p Co f , Kew , M r an . . B B m n Machaeran hera . . S i s . 1990. A s ma i s u of o g , D L p o sy te t c t dy t As era a l ( t ce e) and re ated groups using res tri cti on site an alysi s of chloroplast DNA .

- S s . y t Bot . 5 3 1 . m s C . L . 1989 . A ster intrica tus A s ra a : A stereae rans rr Ne o , ( te ce e ) t fe ed to Ma ha eran h r - Ph tolo ia 44 . c t e a . y g 0 m L s G. . L. A . V ro ik R. L i n i f er o b . H artman . 1990. The o Ast Ne o , , , de t ty ble ha r h - llus A . Gr A ster a o a e e: A s ra a S . p p y y ( te ce e) . yst B ot. 642 . S un r S D . 1986 . The s s ma i s of A ster su . tri lium m i dbe g , y te t c bg Oxy po (Co pos tae) and hi ri ll alli s i s a s. PhD . iss. ni . T xas A us i n T x as to c y ed pec e d , U v e , t , e . T rn L u r B . R . L . m a . 1 6 . Inf H ar n . 7 e , t 9 raspecific categori es of Macha eranthera inna ti da m a W htia - si . ri 3 1 p fi (Co po t e) g 5 . - P hytologic (Februa ry 1997) 1 13 .

S YNO P S IS O F S TE P HA NODORIA (A S TE RA CEA E : A S TEREA E)

s m Guy L . Ne o

x as R i nal Ins i u for En ir nm n a l S u i s S am H us n S a ni rsi Te eg o t t te v o e t t d e , o to t te U ve ty ,

H un s ill T x as 77341 U . S . A . t v e , e

A BSTRA CT

A full descri pti on and ci tation of Specimens are provided for the monotypic t ria te ha nodoria mentella i s ri Mexican genus S ephanodo . S p to rest cted to a n rth as rn S an uis P si h r it i s rar and in dan r small area of o e te L oto , w e e e ge of “ ” n e i s i s art the Xanthoce ha lum r u of the su ri ex tincti o . Th Spec e p of p g o p bt be Machaerantheri nae but its h l n i si i n i hi n the Xan tho ce ha lum , p y oge et c po t o w t p m as i l r la i n hi group i s not clearly resolved. So e pects of poss b e e t o s ps among u other genera of thi s group also are disc ssed.

KEY R S : Ste hanodo ri a As ra eae Mexico ax n m WO D p , te c , , t o o y

e r n was rs s ri d Th Single species tha t comprises Stephan odo ria E . G ee e fi t de c be by It was rmall x lud Robinson ( 1892 ) as a species of Xanthocepha lum Willd. fo y e c ed

r m the la r r n who r a the new nus for it. Af r ha it w as f o tte by G ee e c e ted ge te t t, not discussed or i ncluded i n publi shed studies evaluati ng relationships among Astereae until the recent di sserta tion by Sub ( 1989) and its publication by S uh S impson ’ who substantiated Greene s rationale for placing Stephanodo ria as a m n i nus l s l r la e t anthoce ha lu o oty p c ge c o e y e t d o X p m .

te me S hanodoria to ntella B .L . re n i n d in h r ari a nl a p ( . Rob ) G e e s represe te e b by o y s im ns in ludi the ma ri al ll Pri n l and a mu h m r few pec e , c ng type te co ected by g e few c o e r n ll ti ns m n ar aus te hanodo r ia is rl n n ece t co ec o fro e the type locali ty . Bec e S p poo y k ow and n an r i n its n i nu d x is n a ull s ri i n and i a i n of im ns e d ge ed co t e e te ce, f de c pt o c t t o Spec e are r i p ov ded .

S TEP HAN ODORIA T MEN L . R ins n E . r n O TELLA (B . ob o ) G ee e

Ste ha nodoria t me lla B v o nte E r n en . m . no h a .L . Rob . Er p ( ) . G ee e , g co b . yt e

1 895 . A SI M: Xan th l oce ha um tomentellum B .L . Rob . Pr . Am r. B ONY p , oc e

A a . A rt s 1 T PE: ME I . an ui P si : alkalin c d 892 . Y X CO S L s oto e 108 P H Y T O L O G I A February 1997 volume - 1 13

W an ar l 1 1 ul m ad s H a i nda de A n s ura ca . 7 mi SS of S J 1 89 1 e ow , c e go t [ B to o] , , rin le 3 761 H TYPE: GH Is s: CC . P g ( OLO ; oty pe

P r nni al h r s r m d r s r du in h r as al s s m e e e b f o woo y oot , p o c g S o t b off et , ran hi n ri mari l in the in resc n s ms l a s and h llari es i i a tall, b c g p y flo e ce; te , e ve , p y St p t te

- ar d minu el hirt ll us ilosulous not lu i n us. a s hi and glandul an t y e o p , g t o Le ve t ck l sh h n resh the asal n s num r us rsis n the la s a noti ceab y fle y w e f , b o e e o , pe te t , b de obov te l n - lli i n ir m s l 15 - 30cm l n 12 -40mm i as all a enuat to a to ob o g e pt c , e t e , o t y o g , w de , b y tt e i l 3 - 12 cm l n and the l n th the l a auli n l a s r du in siz pet o e o g e g of e f; c e e ve e ced e ,

il u ards. H ads in an en l s r m s ssil sh becoming sess e pw e Op , oo e co y b , e e to ort pedicellate in corymboid clusters of 2 -6 ; pedi cels mm long; involucres ur ina mm l n 3 - 5 mm i ; h ll ri s r n l ra uat in 5 -7 s ri s t b te , o g , w de p y a e St o g y g d ed e e i h the u erm s as l n as the i nn r lan la - a m s l w t o t o t o g e , ob ceo te obov te , o t y in ura d the a x r ni sh s r a in to reflexed th e la ral mar ins i h a hi n d te , pe g ee , p e d g , te g w t t , i al mar i ns n minu l s i i a - landular r a l s sli l hyaline rim, ap c g ofte te y t p t te g ; ecept c e ght y

- - n x i h low al lar ri s. Ra rs ca. 2 0 30 ld n ll istilla co ve , w t veo dge y flowe , go e ye ow , p te and r il i h r llas mm l n ars l and mi nut l s i i a - lan ular fe t e, w t co o o g , Sp e y e y t p t te g d on the u and li ul s mm l n mm i . is rs rtil t be , g e o g , w de D c flowe fe e , ell minutel s i i a - landular on the u e and l r lim r x e for the y ow , y t p t te g t b owe b , e ect e c pt uter hi h l an u ard a r l n a i n the u i h o r llas mm l n o , w c e o tw fte e o g t o of t be , w t c o o g , in ludin the mm l n u l n a in at ma uri and l a in the r lla c g o g t be , e o g t g t ty e ev t g co o duri n an hesi s a ru l iden d in the lim i h the deltate l s mm g t , b pt y w e to b , w t obe l n re an h rs ins r ed ca . hal a d n the u i h the u usuall o g , e ct; t e e t fw y ow t be, w t t be y becomi ng noticeably different in color and tex ture at that point; style branches with - A h n s m s l lan la - deltate tri angular collecting appendages. c e e o t y ob ceo te oblong to l n - lli i s r n l a ned 2 - id i h 2 thi ri s or s m im s i h a ob o g e pt c , t o g y fl tte , S ed w t ck b o et e w t narr hird sid and hird ri b mm l n mm id s ra - l red ow t e t , o g , w e , t w co o , shi n la r us or r ars l sh r -s ri se nl at the a x a us a h ali n y , g b o ve y Sp e y o t t go o y pe ; p pp y e , la rate or er se- mar i n r na mm hi h or s m i m s 1 - id d and ce o g ed co o g , o et e S e

- somewhat awn like .

= Fl ri n r m un b r. hr m s m num r 2 n 6II La ne 2 92 0 owe g f o J e to Octo e C o o o e be , ( , r rt d an Li 1993 Nesom Wells 668 1 r r h r r m a un epo e by L e ; , epo ted e e f o co t - ll obtained from field co ected buds) .

” The n ri nam i s d ri d r m the r l m n s s han s r n ge e c e e ve f o G eek e e e t tep o c ow , “ ” referring to the coroniforrn pappus) and dori a (an early name for the goldenrod Solida go) .

i i nal ll i A ns xamin : ME I . S an uis P si : M i . del dd t o co ect o e ed X CO L oto p o Cd . Maiz : km of the Es u la Primeri a in Las T a las 2 7 J ul 1979 Lane 2 5 35 N c e b , , TEX Las T a las Lane 2 92 0 not s n mi Las Ta las RR ( ) ; b , (COLO , ee ) ; N of b ( r ssin i n n 14S e 1 m Wells 6681 u h r for hr m s m 988 Neso J . c o g tow ) , p , vo c e c o o o e un and hl r las DNA s u i s is ri u A RI CA S F GH M MEXU co t c o op t t d e (d t b ted to Z , , , , , , MO M TEX S i i n PA T R U M . rri s n ar the railr a s a at rri s , NY , Z, , , ) ; p o Ce to , e o d t t o Ce to , J un 1 8 1 Leidi La e 9 , g n s . n . (TEX) .

Stephanodo ria is endemic to a small area of northeastern S an Luis Potosi near the ns rri Las T a las and an s S art l . The l a i n ran s r m a u tow of Ce to , b , B o o e ev t o ge f o bo t 95 0 to 1050 m rs a min d r m s r ra hi m a . The lan s r in low ete ( dete e f o topog p c p) p t g ow , ri i all rass s in ar as of us- al al i n pe od c y wet , g y pocket e gypseo k e meadows with scattered Nesom : S yn opsis of S tepha nodo ria 109

m s ui e un i r and su ul n - l a d shru s and h r s in lu i n L i u m e q t , j pe , cc e t e ve b e b c d g yc ,

Ma tenus Fla veria Sa m ol s S esu i um and Sua eda . Prin l s ri the y , . u , v , g e de c bed “ disti nctive area around Las Tablas in hi s di ary (Davi s 1936) as meadows miles in x n r i h d rass i n rru l s of uni r r s and un e te t, cove ed w t eep g , te pted by be t j pe fo e t bo ded by - lim - r Th s m ad s a ar to r s gray desert hills of half bare e ock . e e e ow ppe e t on a A u h sod and la r of la s il a in hi n ss subterranean lake . to g ye b ck o few feet t ck e covers ”

o r bolus ai roides rn T rr . i ne of the min mud and water. Sp o ( To ) o s o do ant grass ec1e Sp s.

The gypseous soil i n thi s area i s deri ved from the ex posed gypsum and gyp seous “ " limestone of the surroundi ng hill s and mountai ns (the gray desert hills descri bed by The an i n na ur of thi s ha i a i s a s d the urr n of h r Pri ngle) . c e t t e b t t tte te to by occ e ce ot e gypSOphilic endemics restri cted to the same immedi ate area or nearly so: Oldenla ndia P luchea mex icana d r s m Vi uiera otosina l L . Rob . a rin lei B . p g , (Go f ey) Ne o , g p B ke ,

ul B rande ee and eissole is sua edi olia B . L . h r a multi a is G Rob . P ina op ppus c g , p f Bot Stepha nodo ria and Geissolepis are monotypic and are Strongly divergent in la i morphology compared to their closest re t ves.

The water table around Las Tablas and An gostura (the type locali ty) apparently has been lowered by artifi cially channeled drai nage and the area is being modi fied in other a it has n r u h in as ura and in 1988 the n inu d n w ys. Much of bee b o g t to p t ge co t e existe ce of S tepha nodo ria was estim ated to be seri ously endan gered .

The Stephanodo ria population sampled by Nesom Well s in 1988 immedi ately north of Las Tablas consisted of about 50 to 75 indi viduals scattered along low areas ar th e The ula i n of Ste hanodoria sam l d in 198 1 an eidi ne road. pop t o p p e by L e L g i hi n the n erri s ul not be r l at d in 1988 the re s n au h r w t tow of C to co d e oc e by p e t t o , apparently because the ar ea in the vicinity of the railroad sta ti on is almost entirely n n u der culti vatio .

Phylogenetic position of Stepha nodo ria

In tran s rri n Xa ntho ce ha lum tomentellum to Ste han odoria re n 1 895 . fe g p p , G e e ( , p 12 ) noted that it is

a lan not far r m d i n na ur r m olida o ri ida but i h a coroni forrn p t e ove t e f o S g g , w t ra h r han a illar a us hi h on a un a m r anal of its t e t c p y p pp , [w c ] cco t of e e ogy a T i al us has n u lish d un r the r n nus . p pp , bee p b e de w o g ge yp c Xan thoce ha lum has no a us at all but a ur i ann ulus ins a and the p p pp , t g d te d; i nus sh ul limi h s s i s a ar n l . B ut n i h r the ha ge o d be ted to t o e pec e , pp e t y e t e b t , the in r s n the i n lu r nor n the m r ss a h n s of hi s lan flo e ce ce , vo c e eve co p e ed c e e t p t ” ar e at agreement with Xa ntho cepha lum or an y of its near allies .

The habit and overall appearance of Stepha nodo ria are i ndeed similar to Oli oneuron olida r i ll s r r n but h s two S o idum L . S ma as g ( g ) g ( ) , ob e ved by G ee e , t e e species are relati vely distan tly separated wi thi n the Astereae (Nesom 1994) an d their ’ similari i s are n n a a l su rt R i ns n s ri inal t e co vergent . Rece t d t (be ow) ppo ob o o g 1 10 P H Y T O L O G I A February 1997 volume - 1 13

’ h sis hat Ste hanodoria and Xanthoce ha lum are l sel relate but r n s hypot e t p p c o y d, G ee e exclusion of Stephanodoria fr om Xan thocephalum has not been form ally challenged te by any subsequent students of the A s reae.

Morphologi cal and molecular evidence that Stephanodo ria i s most closely related to in lu i n Xanthoce ha lum i s summari z ed an et l lus er n ra a . a small c t of ge e , c d g p , by L e , “ ” This is the Xan thoce ha lum r u es m Xa ntho ce ha lum p g o p (N o p , u Ra ks onia H ar man an ri ndelia ill ha nodoria Isocoma ac . Step , N tt yj t L e , G W d “ ” i the rindelia r u in F (including P rio nopsis and Ol vaea Benth . G g o p igure 9 et E id n for m m rshi the Xanthoce ha lum r u i hi n the of Lane, v e ce e be p of p g o p w t l d ned rth Am ri an Machaeranthem s lad was h n i n the strong y efi , No e c Nee c e S ow f S ub 1 8 and M r an summ ari z d s m l ular da a o 9 9 et a l. mo ec t ( ) o g e by Ne o ,

and rr ra d in ane et a l. 1996 in a r a r n x . The co obo te L , ( ) b o de co te t Xanthocepha lum group was formally placed withi n the subtri be Machaerantherinae by Nesom

ar hi n ir lan la l a s Ste hanodoria are r mar a l The l ge , t ck , e t e , ob ceo te e ve of p e k b y

m s i s P rrocoma H . P . rocea A a similar to those in so e pec e of y ook c [ . Gr y]

re n m r so han an i n the Xa n thoce ha lum r u . P rrocoma i s a m m r G e e) , o e t to y p g o p y e be of the Machaerantheri nae but consistently placed outside of the Xanthocepha lum group (S ub 1989; Morgan 1990; Morgan S impson The strongly flattened and elongate achenes of Stepha nodoria also are more similar to those of Pyrrocoma than to ha lum r u Ma es 1 76 s ula ed ha a l s r la i any of the Xanthocep g o p . y ( 9 ) po t t t t c o e e t onship mi h xi n P rrocoma and Ra a cksonia ased on th ir mu ual r du g t e st betwee y yj , b e t p o ction of

flavones. r ad r iden e h er indi a s ha the similari n P rr ocoma B o e ev c , owev , c te t t ty betwee y and Stepha nodori a (as well as Rayjacks onia ) has resulted from evolutionary parallelism (but still reflective of close common ancestry) .

Stephanodori a and Xan thocepha lum are positioned essentially as Si ster tax a by m l ular a a an et a l. 1996 an Har man but her is li tl in h ir o ec d t (L e, ; L e t t e t e t e n n i The are la ed as s r tax a nl morphology leadi g to the same co clus on. two p c si te in o y the re s as s l l on m r h l an H artm an 1996 Fi ur s 1 two of five t e b ed o e y o p o ogy (L e , g e Plan ts of both genera produce a coroni forrn pappus and sparsely pilose and s i i a - lan ular ra and dis r lla an H ar man s r d a h n s h t p t te g d y c co o tubes. L e t co e c e e of bot la r us ut in m r ati ns - as b s h ha e minu e in hairs n ar the rui a x . g b o , y ob e v o bot v t tw e f t pe The lack or slowness of coiling of the ray corollas also might be taken as a Similarity

- n the n ra . In the DNA as anal ses an Hart m an the betwee two ge e b ed y of L e , two genera as a pair are placed in vari ous positi ons relative to others of the Xanth e oc pha lum group .

Other tax a of the Xa nthocepha lum group

Gri ndelia P riono sis and li va ea , p , O

The m n i P rionO sis has n r a as a i s of rindelia s m S uh o otyp c p bee t e ted Spec e G (Ne o , , S im s n The m l ular a a of an et a l. 1996 su rt hi s si i ni n p o o ec d t L e , ( ) ppo t po t o g , but Lane H artm an ( 1996) mai ntai ned these two as disti nct genera bec ause of two m r h l i al i r n i s s s the a us . In P riono s is the o p o og c d ffe e ce (be de p pp ) (p p ,

1 12 P H Y T O L O G I A February 1997 volume - 1 13

Rayjack sonia

Har man 1976 1990 ri inall re niz ed the hr ies the H a lo a an/ t ( , ) o g y cog t ee Spec of p p pp “ ” Machaerantheran phyllocephalus group as an undescri bed genus closely related to es ri ti n the new nus and new com ina i ns Xan thocepha lum . D c p o of ge b t o were provided as Rayjacksonia H artman Lane The three species of Rayj acksonia r n in sti ur s inul se li ar ee h s li ar h ads and ra are cohe e t ve t e , p o fo t t , o t y e , y flowers with min n li ul s but h a aren l la a set m r h l i al a ures ha pro e t g e , t ey pp t y ck of o p o og c fe t t t could Tw s i n hi h n be interpreted as Specializ ed . o pec es have mo omorp c ac e es and phyllari es

n l ndura e at the as hi l R. h lloce ha la . H artman an stro g y i t b e , w e p y p (DC ) L e produces di morphic achenes (apparently developed in parallel with those of Gri ndelia and l n n- n ari Oli vaea ) and comp etely herbaceous ( o i durate) phyll es.

Xanthocepha lum hum ile

One of the most interesti ng problems remai ni ng in the Xan thocepha lum group i s

humile n h . the r s f the the nature of the relationship of X. Be t to e t o genus and the ’ S uh s m l ular da a h his s i s m r l sel r generic group . o ec t S ow t pec e o e c o y elated to

ma lun ora T rr . ra re n han to h r s i s Xan thoce Isoco p fl ( o G y) G e e t ot e pec e of pha lum .

humi le and I. veneta un h ha n Natural hybri ds between X. K t ve bee collected from se ral l ali ties in en ral Mexico Hartman ane The anal sis an ve oc c t ( L y by L e, et al. in ntras sh s . hum ile asal to a lad ha in lud s the r s co t , ow X b c e t t c e e t of

as te ri a . h u hum ile i s set Xanthocepha lum as well S pha nodo A lt o gh X. apart from the r s f anthoce ha lum its l s s m r h l i al imilari i h h s s i e t o X p , c o e t o p o og c S ty w t t o e pec es seems a aren It ill in r s in see if ur h r s udies r id id n for pp t. w be te e t g to f t e t p ov e ev e ce thi s.

a i s i e and W r r H .B . 1936 . L o k o us uernse P rin le. Fr Press Pri n i n D v , f f Cy G y g ee t g l n n V rm n Co . ur i , B gto , e o t .

- C.D . .H . aman . 1 6 on D. 9 3 . The nus liva ea m si a De J g, J Be ge O (Co po t e A stereae) .

- B ri ttoni a 92 .

- - r n E L. 1 895 . s r a i ns on the m si a . V III . E h a G ee e, Ob e v t o Co po t e ryt e 15 . Har man R . M r L. 1976 . A ns us of acha e an hera m si a : te t , co pect t (Co po t e A s reae) and a i ma i ud the i n s s r D . se Ble ha odon . Ph . di ss rt a i n ni f b o y te t c St y of ct o p e t o , U v . o T x as A us in T x as . e , t , e

1990. A ns us of Ma haeranthera A st rac a : e co pect c ( e e e A st reae) . Ph tolo i a -4 y g 65 . H man R. L ar M .A . an . 1 . 99 1 . A na ural in r n ri h i e t , L e t te ge e c ybrid n th x 6 group he - t A stereae A s ra a . Si a of ( te ce e) d 32 9 .

an M. A . i . 1 hr . L 993 . m s m num r r L e , J C o o o e be eports i n Compositae with m has is on ri A stereae the s u h s rn ni S a an d M e p t be of o t we te U ted t tes exico . Sida

- 546 .

an M . A . R. L . H ar m an . l i L e , t 1996 . Rec ass fication of Nort h A meri can H a lo a us m si a : Astereae m l : R a cks onia en . n v a o . Am p p pp (Co po t e ) co p eted yj g er.

- J . Bot . 370. Nesom : S ynopsis of Stephanodoria 1 1 3

M r an . S uh A D . R. B . B . S im s n R an M . . K. ans n . . L e , o g , Y , p o , J e 1 996 R la i nshi s of r h Am ri an n ra of Astereae as d on hl r l NA e t o p No t e c ge e , b e c o op as t D

In : Hi n D . J r s ri i n si da a . .N . H . J . Beent e i a e e t ct o te t d , j Co mpos t : 4 - m ti . 9 77. Pr . Inte tl. o m . n . S stema s 1994. y c , pp oc C p Co f , Kew ,

- - li vaea n h . Fla m N al . o G iciano 645 . M v au h R. 1984 O o ni rsi c g , Be t v U ve ty

hi an Press A nn A r r Mi hi an . of Mi c g , bo , c g mi an d c s 6 . A ax n hemos tematic u f a s R A . 197 s o the nu M ye , cytot o o c y t dy ge s

P rrocoma As ra eae: A stereae . Ph . D . i sser a i n ni . of T x as A us i n y ( te c ) d t t o , U v e , t ,

Tex as . d Machaeranthera M r an DR . 1990. A s st ma i s u As rac a and r la o g , y e t c t y of ( te e e) e ted groups usi ng restri ction analysi s of chloroplast DNA and a tax onomi c revision of

Ma ha eran hera s i n P sila ctis . Ph. D . i ss r a i n ni rsi of T x as A us i n c t ect o d e t t o , U ve ty e , t ,

Tex as.

2 . i B . Sim s n. 199 A s s ema s ud Macha er M r an D . R. B . anthera o g , p o y t t c t y of and r la d ro u s usin res ri i n i anal sis of hl r las (A steraceae) e te g p g t ct o S te y c o op t DNA .

- 5 3 1 . S yst . Bot . i in the s ma i x an and M xi an rindel 0. S u s s s T ia s m G. L . 199 Ne o , t d e y te t c of e e c G

- tolo ia 3 2 . (A steraceae: A stereae) . Phy g 3 a P 4 S u ri al lassi i n i n the Astereae A st ra a . h tolo ia 199 . bt b c fic t o ( e ce e) y g

- 15 1 .

R. r an B .B . S im s n . . M . t s m G.L . S uh D 1990 X lo ham ia Ne o , Y , o g , p o y ew n r la d utha m i a - A s ra eae: A stereae a n us e E ia . S d 1 16 . ( te c ) , ge e t to

S uh B . B . S im s n. 1992 . P ri ono si s As era a : Astereae s m G. L . Ne o , Y , p o p ( t ce e )

- Ph olo i a 346 . uni ted with Gri ndelia . yt g

ri i new an ll M xi c . . R ins n B .L . 1 892 . s ns l s d in ob o , De c pt o of p t co ecte e o by C G

d 1 8 1 i h n t s u n a her i s. Pr A m r . Pri n l in 1890 an 9 . g e , w t o e po few ot Spec e oc e

- a 185 . A c d . A rts

S uh 1989 . Ph l n i s udi s of rth Am ri an Astereae A st ra a ase , Y . y oge et c t e No e c ( e ce e) b d

on hl r las DNA . Ph .D . di ss r a i n ni . of T x as A us in T x as . c o op t e t t o , U v e , t , e

S uh B .B . S im s n . 1990. Ph l n i anal si s hl r las DNA in rth , Y . p o y oge et c y of c o op t No

ter . A meri can Gutierrezia and related genera (Asteraceae : A s eae) S y st . Bot .

- 670. - l 2 8 P hytolo gia (February 1997) .

N A THE ECO LOGY OF A GRIMONIA I CIS A TORREY 81 . GRAY (ROS A CEA E) IN THE WES T GULF COA S TA L P LA I N

MacRoberts M .H . MacRoberts B . R.

i i a R sear h 740 lum ia Shr rt o u s na 71 104 U . S . A . Bog e c , Co b , evepo , L

arium Mus um of i S i n s uisiana S at ni rsi i n Shr rt Herb , e L fe c e ce , Lo t e U ve ty evepo ,

S hr r ui siana 71 1 15 U . S .A . evepo t , Lo

A BSTRA CT

l A rimonia incisa in is d roovebur in eas T x as i s The eco ogy of g , c e g , t e

a undan in n in r s . It is r a l n described. It can be b t ope p e fo e t p ob b y ot widespread today because of widespread habitat alterati on resulti ng from fire

suppression during the twenti eth century .

RDS : A rimonia incisa T rr A . ra T x as R sa eae KEY WO g o ey G y , e , o c , A ngeli na National Forest

INTRODUC TION

Little is known about the biology and ecology of Agrimonia inci sa (Kral 1983 ;

R ins Hardi n 1987 rz ll 1990 rac 1993 Sin hurst 1996 . li n rs . obb ; O e ; G e ; g ; G K e , pe

mm . n il r n l its no n di s ri u i n was the l r as al lain r m S u h co ) U t ece t y , k w t b t o owe co t p f o o t

ar lina Fl rida s ar d Mi ssissi i Ra r et a l. 1968 rz ell 1990 C o to o , we tw to pp ( dfo d , ; O ;

Si n hurst 1996 lin rs. mm . In 1989 a di s un ula i n was r r e g ; K e , pe co ) j ct pop t o epo t d r m A n lina ati nal F r s as r un T x as a u 500 km s of the f o ge N o o e t , J pe Co ty , e , bo t we t closest known Mi ssissippi Site (Mahler

In 1993 and 1994 S inghurst ( 1996) reported A g rimonia incisa for eleven Sites in

as r ina - a in inni n in 1 95 ha An l n and S un i s . 9 J pe , ge , Newto , b e co t e Beg g , we ve ndu in isa ar un the ri inal n n l a i ns on the co cted extensive surveys for A . c o d o g k ow oc t o An lina a i nal F r ha i n i a is ri u i n ar a a r x ima l 1 1 km ge N t o o est . We ve de t fied d t b t o e pp o te y x 4 km straddling the Jasper- A ngelina county li ne south of the S am Rayburn

° ’ ° ' R s r ir n rin n 4 i hi n his area the s i s i s now e e vo ce te g o 3 1 04 N 9 1 1 W . W t t pec e n n r m i hi s ar a r 50 i s man hi h ha d ns ula i ns. u s k ow f o ove S te , y of w c ve e e pop t o O t de t e , ha ar h s for it i n li l ha i a on the A n li na a i nal F r s e. . n r h of we ve e c ed ke y b t t ge N t o o e t ( g , o t MacRoberts MacRoberts: Ec ology ofA grimonia incisa 1 15

th e S am Ra urn R s r ir and s iall al n the area to the s in lu in yb e e vo e pec y o g we t , c d g in S ri n s and lan Islan il rn ss hi h has similar 5 0113 but ha Boyk p g Up d d W de e , w c ) ve failed to locate it.

i the a i nal F r s ha un A rimonia inc i sa at hr l a i ns in Outs de N t o o e t, we ve fo d g t ee oc t o rn n un al n S a Hi h a R2 5 5 a u 32 34 and 5 1 km as of northe Newto Co ty o g t te g w y , bo t , , e t a i nal F r s i s and S i n hurst 1996 has r rt d it for the A ngeli na N t o o e t S te , g ( ) epo e one as the S am Ra urn dam in as r un locati on about 6 km e t of yb J pe Co ty .

S inghurst ( 1996) reports it for two Sites on the S abine National Forest: one in s u h rn S a in un and one in n r h rn n un . A ls re n h r ari um o t e b e Co ty o t e Newto Co ty o , ce t e b urn u im ns r m A nd rs n un T x as 19 km n r h searches have t ed p Spec e f o e o Co ty , e , o t west Thi s si i s a r xi mat l 185 km n r h s f of Palestine near S and Lake . te pp o e y o t we t o the A h th A n n un A ngelina Nati onal Forest populations. searc of e derso Co ty area on July 2 4 1996 ail d di s r A rimonia incisa and r n rm hi s im ortan ran , , f e to cove g eco fi t p t ge e i s is not lis d as ar of the ra n ex tension of the Species. Th Spec e te p t flo of E geling

a m n A r a us n r h of S and a e T l ai r et a l. Wildlife Man ge e t e j t o t L k ( e f ,

GENERA L DES CRIPTION A ND BIOLOGY

A rimonia incisa is one of s n r h Am ri an s i s in the nus li n g eve No t e c pec e ge (K e , hni al ri i ns of it and its n n rs can und pers. comm ) . Tec c desc pt o co ge e be fo in several sources (Kral 1983 ‘ Robbi ns Hardin 1987; Orz ell 1990; Singhurst Four s i s A rimonia are n n for T x as hre of hi h ur in the south eastem pec e of g k ow e , t e w c occ W r W allr . and A . rostellata allr . rrell ar the s a e: A . in isa A . mi ro a a p t of t t c , c c p , , (Co

hns n 1970 hns n 1990 ix n ll 1993 li n rs . mm . A rimon ia Jo to ; Jo to ; N o Ke ; K e , pe co ) g in isa i s the m s di s i n i : its mid- s m l a es i h nin h r ars l i n i s c o t t ct ve te e v w t e S o t , co e y c ed ma r l a s mid- s m rminal l a leSS h an 3 cm l n usuall i h i h de jo e flet ( te te e flet t o g , y w t e g t ep i n isi ns or ni n h i n i it imm i at l Al h u h it i s not r u n l c o e teet ) de t fy ed e y . t o g f eq e t y illus ra the li n ra in s i n Ri t 1967: 192 h the l af a rn r l . t ted , e d w g cket ( ) S ow e p tte pe fect y

A ri n ia i isa g mo nc is an easy plant to moni tor because it can be located year roun d . It l ms r m ul to m r an i a al B ut r s ms of the b oo f o J y Septe be d d es b ck i n the f l. befo e te die a a w a ar ne sal r s a ars r m a bud a a n the old s m . E n ye b ck , b o ette ppe f o dj ce t to te ve the all dr s ms and i h r l a s old lan s rsi s l n i n the ll i n t , y te w t e ed e ve of p t pe t o g to fo ow g ar ye .

PLA NT A SSOCIA TIONS

In as T x as A rimonia in isa a rs s ars - an i i n u lands and e t e , g c f vo p e c op ed p e p m ra l n in - oak ass i a i ns and h r a u la r i h r ode te y ope p e oc t o a ric he b ceo s ye w t few sh ub s . “ ” H arcombe et a l. 1993 r r imi lar ha i a in th e B i Thi as u lan , ( ) efe to S b t t g cket dry p d ” forests and savan nas and provide a good descripti on (see also Mari etta 1979 ; W ard 1984; B ri dges Orz ell 1 16 P H Y T O L O G I A February 1997 volume - 12 8

lan ia i ns sur e d ten rman n 2 p t assoc t o , we v ye pe e t 10m sites on three asi n Ma 19 ul 2 8 and er 3 1996 and re or d all i n i a l s e i s occ o s ( y , J y , Octob , ) c de de t fi b e p c e a i ns on the An lina a i nal F r s A n li na and as r present . The plot loc t o ge N t o o e t ( ge J pe counties) were chosen for their abundance ofAgrimonia incisa indi viduals and to gi ve a t r r the F ar a u i d the lan a representative geographi c sc t e ove orest e occ p e by p ts.

Species occurri ng in the sample plots are listed below according to relati ve

urr n d fin d as r s n in num r of l s. m n la ur frequency of occ e ce , e e p e e ce be p ot No e c t e mainly follows Kartesz

Occurred in ten plots: A ndmpogon ternar ias Mi chx Centrosema vi rgin ia n um

hemus Mi hx . Dichantheli um acicula re x r r P i r . n h . roton a ant s . e (L.) Be t , C gy c , (De v o )

i ramin olia Mi hx . u . P in us alustris Mill . ul lar P it o s s P . Go d C k , y p g if ( c ) N tt , p , L un z and hi a h rium s o a ri um Mi r ica ns . S hx . ash. Tox icodendron ad ( ) K t e , c z c y c p ( c ) N

l tida r r in nin l s: Ambrosia artemisi o ia L . A ris u u as ens P i r . Occurred e p ot if , p p c o ,

lochia reticula ta ac . Berla ndiera umi la n i h l s: A risto Mi hx . Occurred i e g t p ot J q , p ( c )

mi um em er vi rens L. St. Hil. H elia n thus mollis Lam . Pas alum Nutt Gelse s p ( ) , , p

hx Rhus co a llina L. Ruellia humilis u Sassa ras a lbidum u setaceum Mic p , N tt f (N tt .)

Mi hx . ees Sti llin ia s lva ti a ard n ex L. and Tra ia urti olia N , g y c G e , g cif c

urr d i n se n l s: As imina a rvr ora Mi hx . unal a llicar a Occ e ve p ot p fl ( c ) D , C p

Mi hx . H e otis n i ricans m L. G mn0 o on ambi uus d La F s r , y p g g ( c ) y g ( ) o be g ,

nia A . Smal P aspalum plica tulum Michx and Vem o texana ( Gray) l.

curred in six l s: Eu ho rbia orollata L. Galactia re ula ris L . Oc p ot p c , g ( )

Gna ha lium obtusi oli um L . Rh n hosia lat olia u . ex T rr A . ra p f , y c if N tt o ey G y ,

he t ora . B . . P and Vitis a esti va lis Rudbe kia hi rta L. St losa nt s b L S Mi hx . c , y fl ( ) c

urr d in l s: roton micha ux n G.L. s r Era rostis s e ta bilis Occ e five p ot C Web te , g p c - Pursh S u . landula ria anadensis L . u . Di ita ria co na ta J . A . S hul s ( ) te d , G c ( ) N tt , g g ( c te )

Pil r uer us incana B artr uercus ma rila ndica Muenchh . Rub us S . Schrankia ge , Q c Q , p , h r st icina S mall S an l. and S orobolus un ceus B eauv . un h . y ( ) t d , p j ( ) K t

urr i n ur l s: A ster a tens A 1t Ba tisia leuco ha ea u . eanothus Occ ed fo p ot p p p N tt , C me a ida L . Eri er n tri o uhl x l . rican us ornus or o s sus M . e il Erio onum L C fl , g g W d , g lon i li ian themum a r lin Mi hx H meno a o um u . H el o ianum al . us g f N tt , c (W t ) c y p pp a rtemi ia e olius . er i h er i an s H um oides L. r z Ilex omitor ia A it. Le hea f DC , yp c yp c ( ) C t , v , c ' mucr na W alt. Mi hx . da . e i u o a Raf. Lia ms ele ans P inus tae L P t rid um a ilinum t , g ( ) c , , q

L . uhn S utella ria elli tica Muhl. S t lodon a rneus M i c Moldenk e and ( ) K , c p , y c ( ed ) , Trades an tia re h c verc onii B ush.

urr i n hr l : l hi r m C . F s r a r a s A o a d u mondu raham R. aIba Occ ed t ee p ot p (G ) o te , C y

- L. l . u . ex E l h d u ex a n u l . r r so sis ilosa u . n i os ol s t s Mu l A . S m all ( ) N tt , C y p p N tt , C c ( e g ) , Echina cea a ll ida u . u . el b ia uberula Mi hx . Ph sa li m ll Lo s o is u . p (N tt ) N tt , p c , y N tt , ' uer W an us s ella ta enh . Rh n hos ora lobula n s ha man S mall R ra i Q c t g , y c p g (C p ) , . g y un h Te hrosia i r in i a na Pe and ts Tra ia sma llii Shi nn rs . K t , p v g , g e MacRoberts MacRoberts : Ecology of Agrimon ia incisa 1 17

l ristida la no sa Muhl . e urred in s: A x Ell. As le ias am lex ica ul Occ two p ot , c p p is r ilon di a ri a tum i ceus n . C o t u . Raf. Sm . A s er ser rotalaria sa i a lis L . , t Ve t , p v c (N tt ) C g tt , H eri um entia no ide . B . S . P e r a ea L s L. l hr ina he b Lob ia a endi culata . Eryt c , yp c g ( ) pp A

r raci ua L . Li thos erm m a rolin iens e al Li uida mba st u . ex m l . DC . , q y fl , p c (W t G e )

lea c nancho ides En lm . ds P enstemon australis u Mate s bs . MacM . , y ( ge ) Woo , p P nn ll nn t P h sa lis hetero h lla s P inus e hina ta Mill la xi rus P . flo ( e e ) Be et , y p y Nee , c , v b ula t i m lum h o a eum u t . ex T rr . A . ra R . ar. su um u h P ed o e yp g (N t o G y) ydb (B s ) J . ho t ma rim s S leri a ilia ta Mi hx . and Tri s e seta ceum H u . G e , c c c , c o tt

urred in one l : A er rubrum . A a linis ul hella P nn ll As ter Occ p ot c L g p c e e ,

i a rex om lana ta T rr . H . on a anade is inea rii ol us ns L . r n . l f L , C c p o ook C yz c ( ) C o q t m ul Di ita ria i W Dichan theli um la x orum La v llo sa al . P rs . ailla rdia fl ( ) Go d , g ( t ) e , G

W alt. H R H elia n thus an usti oli us H ieraceum r n vii aesti va lis o o L . ( ) ock ” g f L g , v as i ra r r m i a l e L . ri t P s lutea L . P e sea bo bon ia No thos ordu b o L . S r n c v ( ) B t , fl , ( ) p e g ,

mum rocumbens L . Sa lvia a rea Mi hx . ex Lam S olida ra P ol re o odo A it . yp p , zu c , g ,

Te hrosia onobr cho ides u . Trichostema di hotomum L . Ulm us alata Mi hx p y N tt , c , c d Vitis rot t m a rboreum Marsh Vac in i um stamineum L. an und olia Mi hx V i iu . acc n , c , f c

i i The num r of s i s In the study plots we ident fied 1 1 1 Spec es. be pec e per plot ran d r m 37 5 3 m an 46 S D As ra a d mi na d ll d ge f o to ( e , te ce e o te fo owe by Poaceae Fabaceae and Euphorbi aceae

The plant frequencies li sted above are fairly typical of West Gulf Coastal Plai n u lan l a i n sa ann a sub- mmuni i n hi h A rimonia p d long e f p e v . The co ty w c g incis a occurs most resembles upland longleaf pine savanna subtype 1 (B ridges Orz ell 1989;

H arcombe et a l. but h r are s m im r an i r nc s. The lan s li s d for , t e e o e po t t d ffe e e p t te su e 1 in rid s rz ll 1989 and H arcombe et al . 1993 l arl inha i m re btyp B ge O e ( ) , ( ) c e y b t o

x ri i s han h s in hi h A . inci sa usuall urs. For i ns anc su lan s e c S te t t o e w c y occ t e , btype p t r i t t i d A u eola ria e t na a u t . P nn B ulbos lis cilia t olia El Fem . Da lea s . an p c (N t ) e , y f ( , pp , ' i i all i f x ri Scutella na ca rd o h lla En lm . 81 A . ra hara ri st s o p y ge G y , c cte c y Spec e e c ha i a s are a s n r m our sam l l s h s s i s a r san dhills MacRoberts b t t , b e t f o p e p ot ; t e e pec e f vo ( MacRoberts

Our observati ons suggest that A grimonia incisa favors a sli ghtly more mesic ha i a h an d s ri d ri s rz ll 1989 and H arcombe et a l. 1993 for b t t t e c be by B dge O e ( ) , ( ) su t e 1 an d ha the u lan l n l a in c mm uni a r d A . incisa alls b yp , t t p d o g e f p e o ty f vo e by f ’ sli h l the l Orz ell g t y to eft (see their Tables 1 and 3) of the subty pe 1 communi ty . s “ 1 1 W - t ri an 9902 3 s ri i n of A . inci sa as urri n on ll dr ain bu not x s ( ) de c pt o occ g e ed e c , dy soils agrees more closely with our observations (see also Orz ell

The herbaceous associ ates for A g rimonia in cisa i n the East Gulf Coastal Plain are much the same as those described for the West Gulf Coas tal Plain (Kral ex cept ha h in i a a li h l t t t ey d c te s g t y more x eri c habitat i n the east .

R r s al a s ass i a A r imonia in isa i h l n l a i n hi h m a s the epo t w y oc te g c w t o g e f p e , w c ke A nderson County loc ation particularly i nteresting Since it i s outside the range of l n l a i n and rha l r s the i stri u i n of in al h r. n r una o g e f p e pe p d b t o p e toget e U fo t te y , we we e una l r l a the A n rs n un ula i n but the n ral mm uni in b e to e oc te de o Co ty pop t o , ge e co ty which it was found i s oak - san dylands with many of the species char acteri stic of u lan l n l a in sa anna The n un si s are i n l l ll and slash in p d o g e f p e v . Newto Co ty te ob o y p e lan a i ns ha r r a l n p t t o t t we e p ob b y longleaf pi e prior to thi s century . 1 1 8 P H Y T O L O G I A February 1997 volume - 12 8

LIGHT CONDI TIONS

In order to gather some quantitative informati on on th e light condi ti ons favored by

r i nia in isa a h r a a on an and shru r. Ag mo c , we g t e ed d t c opy b cove

l s u l s i n A n lina n and as r un i s stim a d In twe ve t dy p ot ge , Newto , J pe co t e we e te s ran r m 2 0% 55 % and a ra d A rimonia incisa canopy cover. Thi ged f o to ve ge g

the da but i s not ir l x s m s the da . A t the gets direct sun part Of y , d ect y e po ed o t Of y am im it a rs ri h i ndir li h . s e t e , f vo b g t ect g t

in savanna/shrub x amin d the di s ri u i n Of in i i ual Along a p e edge , we e e t b t o d v d

la i n er. S hru s r a llica r a americana Li uidamba r plants in re t o to cov b we e C p , q st rac ua Ilex omitoria P ersea borbonia A simina a rvi ora om us o rida L . y ifl , v , , p fl , C fl , mi n the m er i idi n it i n hr arts: h co a llinum . xa and R us p We e ed two et edge , d v g to t ee p rmal % an s m dire sun but sha m s Of the im Open (no 35 c opy) , edge ( o e ct , ded o t t e) , n hi s d l a ed 7 lan s: 50 lan Shade (always shaded) . A lo g t e ge we oc t 5 p t p ts were

e n 2 2 lan s r at the d and 3 lan s r in sha . in th Ope , p t we e e ge, p t we e de Of the hr lan s ha und in the shad a ear d i n r ndi i n t ee p t t t we fo e , two pp e to be poo co t o leggy and chlorotic .

sin a s n li h m r m asur li h ndi i ns Of A rimonia incisa U g We to g t ete , we e ed g t co t o g % an und r n rmal ndi i ns at no n . In one s ud l i h 35 the s n e o co t o o t y p ot w t c opy , We to m as was in dire sunli h in ha it was 16/ 17 ha i s in had the li h e ure ct g t; S de , ; t t , S e g t i s about one quart er to one thi rd that measured in di rect sunlight .

These findings confirm our general i mpression that Agrimonia incisa prefers Open

lands and isa ars as li h l ls r as a i in ha al h r . wood d ppe g t eve dec e e , vo d g deep S de toget e

We have never encountered Agri mo nia incisa in closed canopy forest nor in any area with a dense mi dstory or Shrub layer; even a dense cover of P teridium aq ui linum a ars be inh s i a l it. On the h r han ha n r l a it in n ppe to o p t b e to ot e d , we ve eve oc ted Ope ar a a e s with no Sh de .

SOILS

In the s u ar a A rimonia incisa urs on san s on un ula in to hill n l t dy e , g occ d d t g y , ge t y l in u lan s ha are ll rain m ra l rm a l and hi h ui l dr S op g p d t t we d ed , ode te y pe e b e , w c q ck y y uri n r u h d g d o g t .

un it on hre s il s: LTC l e tne - T hran ass ia i n We fo d t e o type ( ) y e oc t o , (DUB) u - in ass i a i n and LeC Letn e l am san s hi h are lassi as Do cette Boyk oc t o , ( ) y o y d , w c c fied l am sili us therrni c A r nic i l A ar n l L e Pl nthic Grossarenic Pa eudults . TC o y , ceo , ( , ) pp e t y , and Le C are the m or sa r itsch et a i . simi lar Dolez el et a l . 1988 Ne e ve y ( , ; , Th s s ils are un in n r h rn as r and n un i s s u h rn S a in e e o fo d o t e J pe Newto co t e , o t e b e

12 0 P H Y T O L O G I A February 1997 volume - 12 8

Ta l 2 . um er lan s in ten l s uri n u i b e N b Of p t p ot d g two s ccess ve years .

Fr m our un uantified sur s r a i ns sus d ha l o q vey Ob e v t o , we pecte t t p an t distri bution was lum w - - c ped. This as confirm ed by the hi gh density to frequency ratio found for all ranse s . In the km rans the rans s m i n l t ct t ect ( five t ect co b ed) , we ocated 88 places Th wi th plants . ese occupied 493 meters Of the total and averaged m (range The a 1 to 37 m) . verage number of plants per clump was (range 1

In km Of ranse m asur not nl the ar a u i d lan t ct, we e ed o y e occ p e by p ts, but al so the i s an n lum s. In the km rans c the a ra d t ce betwee c p t e t , ve ge di stan ce between lum sit s was 55 m ran 3 - 170m and in the km rans the a ra c p e ( ge ) t ect, ve ge distance n lum si s was 3 1 m ran 4- 13 m betwee c p te ( ge 5 ) .

A t only four places Of the 88 in the five transects where A g ri monia incisa w as und was h r nl fo t e e o y one plant .

Put sim l if one lan i s un the r a ili i s hi h a p y , p t fo d, p ob b ty g th t more (Often many m r ill un i hi n a m rs o e) w be fo d w t few ete of it.

FLOWERING

In T x as A rimonia in isa r and - e , g c flowe s produces fruit from mid July through mid S m r Thi s a epte be . p ttern appears to chara cteriz e other populations i n the southeast (Robbi ns H ardin 1987; Grace MacRoberts MacRoberts : Ecology on rimonia in cisa 12 1

The in r s n i s a s i li ra m s m imes ran h ari n num r flo e ce ce p ke ke ce e , o et b c ed , be g e ous al rn a l arran d rs. Fl rs n in su ssi n u the ra em at su h te te y ge flowe owe ope cce o p c e , c

- in r al s ha nl a are n h r . Frui i s to sha i h s ral r s of te v t t o y few Ope toget e t p ped , w t eve ow ri stl s s r a in r m the mi dl and readil a a h s as s rs- b and a h r s b e p e d g f o d e , y tt c e to p e y d e e to air h la s and l hin for a hni al d s ri i n see ral 1983 R i ns h , S oe ce , c ot g ( tec c e c pt o K ; obb H ardi n 1987; Orz ell

TO ath r in rma i n on A rimo nta 1nc1sa ri n a rn and olli na rs on 7 g e fo t o g flowe g p tte p to , , 8 and 18 A u us 1996 m a n i nu us s r a i ns and ri di h s on the , g t , we de co t o Ob e v t o pe o c c eck in the ld an d a ur d s llina rs . The A status Of flowers i n fie c pt e ect po to . in cisa population used for h s s r a i ns w as a u hal a hr u h ri n . In hi s ulati n t e e Ob e v t o bo t fw y t o g flowe g t pop o , i lan s and rs and m ni r d h m hr u h u e we marked Spec fic p t flowe o to e t e t o g o t th day .

We al so made continuous Observations on two greenhouse plants from 15 A ugust in the r nh us s r a i ns r alm s i nti i to 7 S eptember. S ce g ee o e Ob e v t o we e o t de cal w th those n the l but in l a lar r num r Of rs and r mad a t r we made i fie d, vo ve ge be flowe , we e e f e we had rmi n d the asi utlin of the ri n a t rn ill d s ri h s dete e b c o e flowe g p t e , we w e c be t e e

Observati ons in deta il after giving the basic outli ne Of flowering in the field .

For thi s descri pti on we use local time (CDT) : sunri se on 7 and 8 A ugust w as u about am and sunset was about pm . On 30 A gust (the middle Of the i n sunri s was am and suns w r nh us s r a s as m . The g ee o e ob e v t o ) , e et p r nh us s r a i ns r mad in Shr r uisiana 160km n r h of h r g ee o e ob e v t o we e e evepo t , Lo , o t w e e The l s r a i ns r in the plants grow naturally . fie d Ob e v t o we e made northern Jas per un on l ar sunn a s a sh r la e aft rn n hun r sh rs Co ty c e y d y ; few o t , t e oo t de owe occurred on i n the nh us s r a i ns h r was one ai n 7 A u us . ur ree r r as g t D g g o e ob e v t o , t e e wet , y , ove c t

all da s it was hot and humi d. peri od . On y

a n Each The floweri ng beh vior of A grimo ia incisa i s relati vely Simple . flower lasts for two days and Opens twice: the petals Open the firs t time at about pm n th en close at about pm for the night; the flower opens for the second time the nex t day at a u n n and the als in dr in at a u m . ns u n l on an bo t oo , pet beg opp g bo t p Co eq e t y , y given day there are two classes Of flowers: those that ori gi nally Opened the previous da and new rs th e da S ms ha i ndi iduals h lass s at an i n y, flowe Of y . te ve v Of bot c e y g ve im the l r rs n ral l in l the n r on the ra m inc uds n t e , O de flowe ge e y be g be ow ewe ce e S e b Ope su s i l u the a cce s ve y p r ceme .

Floweri ng: Field Observations

Fl r are n owe s begin Ope ning j ust after noon . These flowers th ose that first Ope ed the r Th p evious day . ey tak e about 30 mi nutes to Open and stay Open un til about m at hi h im h in r i n h ir als hi h are all r d m . p w c t e t ey beg d opp g t e pet , w c d oppe by p the 30 rs mar e at m on A u u s 7 all r n on 8 A u us Of flowe we k d p g t , eope ed g t n m and betwee p pm .

S li h l lat r at a u m new rs in nin and n inu g t y e ( bo t p ) , flowe beg Ope g co t e nin un il a u m i h m s ni n uri n the rs h urs m Ope g t bo t p , w t o t Ope g d g fi t two o p

and m . Fl rs n a i l a Th p ) owe Ope r p d y : e ch takes less than half an hour to Open . ey rem ai n Open unti l between pm and pm and remai n closed until mi d- day the n x da at hi h im h ll e a t The an h r e t y , w c t e t ey fo ow th p t ern described above . t e s Of 12 2 P H Y T O L O G I A February 1997 volume - 128

flowers-Of- the-da do not ull in r rse un il la in the a rn n i nni n a u y f y t o t te fte oo , beg g bo t

m. hen h se rs n the n x da heir an h rs r main i ntrorsed. p W t e flowe Ope e t y, t t e e

TO a h r inf rma i n on the rc n a Of s ms ri n s l d g t e o t o pe e t ge te flowe g , we e ecte five widely separated areas where Agrimonia incisa occurs in typical habitat on the al d rans A ngeli na National Forest . We w ke tra nsects counti ng all plants within the t ect Ta l 3 and notin g whether it had flowered or not (had a raceme with fr ui t or not) . b e summari z es these data

i rimon a incisa s ms i h ra m and ru i . Table 3 . Ag te w t ce e f t

2 1 lan s in the rans s 36% had r d and had rui and Of 9 p t five t ect , flowe e f t 64% The rin lan s r alm s in ari a l lar r lacked racemes . flowe g p t we e o t v b y ge than those that did no ha did ma t flower. Those t t not flower y be plants Of the year germinated from last ’

ears seeds or r m u rs. In the firs r i n s as n A rimonia r du y f o t be t g ow g e o , g Spp . p o ce

- nl a h r few l a d lan . a s rar l ha m r han hr l a o y S o t , e ve p t Le ve e y ve o e t t ee e flets . In the s nd ar h d l l a s i al the s i s and r li n rs eco ye , t ey eve op e ve typ c Of pec e flowe (K e , pe mm co . )

Flowering : Greenhouse

Floweri ng sequence and duration of greenhouse plants was the same as for field

rs. ma s r ati ns n lan flowe We de Ob e v o o two p ts.

Plan 1 rs l m on 15 A u u s and had its nal r on 7 S m r t fi t b oo ed g t fi flowe epte be . It had ra m s and 64 r Plan 2 two ce e flowe s. t first bloomed on 30 A ugust and w as followed nl until 4 S m r - a u the rs uart r its l mi n o y epte be bo t fi t q e Of b oo g period . It had si x rs urin hi flowe d g t s peri od .

On l ar a s rs ha had firs n the r i us da re- c e d y , flowe t t t Ope ed p ev o y opened on average at am ran am m sam l siz 67 and an ( ge to p , p e e ) beg droppi ng their MacRoberts MacRoberts: Ecology Of A g rimon ia incisa 12 3

als at m ran m m sam l siz 5 7 rs . 5 5 pet p ( ge p to p , p e e flowe ) Of he al im n was h urs ran - h ur rs t 3 5 s . flowe , tot t e Ope o ( ge o )

On r as da s rs n d at m ran am m ove c t y , flowe Ope e p ( ge to p , sam ple siz e 15 ) and began dropping their petals at pm (ran ge pm to m sarn le siz Tim n on r as a s a ra h urs ran p , p e e Ope ove c t d y ve ged o ( ge h urs sam l siz Thus not nl is imi n la but the ura i n o , p e e , o y t g de yed d t o of

Openi ng i s ex tended on overcast days.

n l ar a s flowers- of- the- da n d on a ra at m ran m O c e d y , y Ope e ve ge p ( ge p sam l siz 66 rs and l s at m ran to pm . p e e flowe ) c o ed p ( ge to

m sam l siz e 66 rs . On r as a s h n at m ran p , p e flowe ) ove c t d y , t ey ope ed p ( ge m to m sam l iz 2 9 and l s at m ran m to p p , p e S e ) c o ed p ( ge p m sam l siz e 85 rs the al im n w as h urs p , p e Of flowe , tot t e Ope o - r w no i r n e n l ar r l u (range 6 10 hours) . The e as d ffe e ce betw e c e o c o dy days: on cloudy days the timi ng of Openi ng and closing i s simply delayed about two hours .

- - - r the r i u da are il i i Flowers of th e da y and flowe s Of p ev o s y eas y d st ngui shed . Th the da are ri h ll and i n i h als an d s am ns at ri h ose Of y b g t ye ow w de Ope , w t pet t e g t r il a r l h an l s the ar and not in rs un l aft rn n o ar nin . A n rs g e to ov y , do t o e t te e oo e y eve g t e P r i us -da rs are dull ll i h s am n are ri h ll . als s ull b g t ye ow et Of p ev o y flowe ye ow , w t t e f y a A n h are lam n s i s d or ar h an h rs ai ns th e s i m a. rs i ntrorsed w r n . , fi e t t te c ed , t e g t t g t e b ow ll da a r n l in P llin ati n ill usua ur the rs i s rs . mm . o o w y occ fi t y flowe Ope (K e , pe co ) A rimonia ies are sel - m a i l if h not u r ss h ill s l - llinat g Spec f co p t b e; t ey do o tc o , t ey w e f po e

h n the an h rs in r rs li n rs . mm . w e t e t o e (K e , pe co )

On the hirt n lan s mar d Each stern has 1 to 9 flowers Open at a time . t ee p t we ke in the ld and ll d on 7 and 8 A u us und 48 rs on the rs da and 5 1 fie fo owe g t, we fo flowe fi t y on the se n a ra i n ran 1 - 9 S D rs er lan er da hal of co d , ve g g ( ge , flowe p p t p y , f hi ld and w Thi s means ha on a ra a h lan has a u w ch were O half ne . t t ve ge e c p t bo t two n w e flowers per day .

The greenhouse plan ts were Slightly different bec ause the mai n plant had two

- ra m s. Thi s lan had a u ran 1 9 SD rs a h da hal ce e p t bo t ( ge , flowe e c y , f Of hi h r new an w c we e d half Old.

POLLINATORS

a u r lli na rs r sm all s the su amil H alicti nae H m n ra C pt ed po to we e bee Of bf y ( y e opte , A i H ali i a a all r a l the sam i s. po de , ct d e) , p ob b y e Spec e

On sunn a s th e s r a i en a u m and m . y d y , bee we e ct ve betwe bo t p p We w sa n n af r m m s a i i i n imm dia l a r the rs n . o e te p , o t ct v ty be g e te y fte flowe Ope ed Th i si h flow - - - did not ma ey v ted bot ers of the day and those Of the previous day . We ke l s r a i ns on r as and rai n a s SO not n how ins i si s mi h fie d Ob e v t o ove c t y d y , do k ow ect v t g t ha i h r on s asi ns. No i ns r s r on the r nh us lan s ve d ffe ed t o e occ o ects we e ob e ved g ee o e p t , al h u h the lan s r not n l t o g p t we e e c osed . 124 P H Y T O L O G I A February 1997 volume - 12 8

FRUI T

rimonia incisa li all A rimonia s e i s r du s lar ar ed rui d si n Ag , ke g p c e , p o ce ge , b b f t e g ed - ran i s rsal al h u h man rha s m st r a l sim l r n ar the for long ge d pe ( t o g y , pe p o , p ob b y p y d op e l n m lin rs . can add no in rm a i n on s i s rsa pare t ste [K e, pe We fo t o eed d pe , x sa ha in la summ r and all our an s and h la s r n r d e cept to y t t te e f , p t S oe ce we e Ofte cove e

h nia rui a es i n the i ness of heir ar s. If s d r wit Agrimo f t, tt t g to effect ve t b b we toope ove an A nia a ch the ma ur ru i asil au h i n our hair . grimo p t , t e f t e y c g t

ROOTS

and m u r Agrimonia incisa has fusiforrn thi ckened tuberous roots rhiz o es. The t be s measure about cm long and mm wide (n To determ i ne if these might play any part in reproduction and in the clumped nature Of A inc isa di ri u i n ll d s ral lan s cut the u rs and lan d ten in a st b t o , we co ecte eve p t , Off t be , p te The u rs r ll c d n 2 pot with soils taken from the collection Site . t be we e co e te o 0May and

lant d on 2 1 Ma 1996 . On 16 ul 1996 ur s r ut d 2 2 ul five had p e y J y , fo p o e ; by J y , maini n did n t s r u The r u s sur i an sprouted . The re g five o p o t . five Sp o t v ved d produced a typical leafy rosette.

hal Of all rth Ameri an A rimonia s i s ha u A bout f No c g pec e ve t berous roots. In

i be W allr and A . a di i n A . in isa A . m ro a r a A . u s ens . rostella ta a d t o to c , c c p , p c , h ve such r s lin rs . mm . Sinc li l is n n a u the c l Of an Of h oot (K e , pe co ) e tt e k ow bo t e o ogy y t ese i s it i s hard ll ha the un i n Of u rs mi h x su s ha Spec e , to te w t f ct o t be g t be, e cept to gge t t t all Of these species might occur in fire-dependent or droughty areas where food reserves l nal re r du i n mi h m or the l ern a i i rt an . r ainl a A . in isa t t ve of c o p o ct o g t be po t Ce t y , c i s su h a s i s and r m ha li tl n a u A . micro a r a it too c pec e , f o w t t e we k ow bo t c p , might experience frequent fire or Short periods of drought

All di scussions of Agrimonia incisa management include statements about fire aus the lan i s ass ia d i h the l n l a s s m hi h i s l arl bec e p t oc te w t o g e f y te , w c c e y pyrogenic

Harcombe et a l. 1993 Pla et a l. ( , ; tt ,

H r li l i s n n a u the c s Of fire or its s as nali i n nsi oweve , tt e k ow bo t effe t ( e o ty , te ty , or timi ng) on the herbs and shrubs of any plant community in the southeastern United S a s R ins M rs 1 2 ra 1 99 993 S r n et a l. t te ( obb ye ; G ce ; t e g , Certainly nothi ng has been publi shed on Agrimonia incisa and fire (S i nghurst The consensus appears to be that these upland pi ne systems burned regularly (every 1 to 3 years) and ha h urn n the r i n s as n t t t ey b ed i g ow g e o . MacRoberts MacRoberts: Ecology Of A gri mo nia incisa 1 2 5

hil not x ensi ha mad s m s rva i ns r l an u W e e t ve , we ve e o e Ob e t o e ev t to thi s i ss e . In 2 Of our 5 . 5 m s ud l s num r 7 la F ruar 1997 s and 8 Ta l 2 urn . te eb y , two t y p ot ( be , b e ) b ed he um r n 4 ar h 1997 un t n O rimo n ia in isa r s s in h . The O 2 M c , we co ted be n c o ette bot num rs r 46 and 18 r s i l . l arl i n hi s small sam l fire did not af be we e , e pect ve y C e y , t p e fect 2 i hin hi s sam urn ar a was als one Of our 10m l s see s i n numbers . W t t e b e o p ot ( ect o “ an hil had not un A rimonia s ms in hi s l Pl t W e we co ted g te t p ot , we u i n h ir num rs s em d to ha en una f the kn ew their distri b t o ; t e be e e ve be f ected by fire .

We have not seen the effec ts of spri ng or summer fire on Agrimon ia incisa and cannot say how plants might be affected by a growing season burn . B ut it i s not imi lar a t rn ha s r d for his arl urn ul un unlikely that a s p t e to t t ob e ve t e y b wo d be fo d .

DISCUSSION

ar It i s on all sta rar lis s h r it urs and is Ag r imon ia inci sa i s r e . te e t w e e occ “

S . F r s S r i . It is d s ri d a l al considered threatened by the U . o e t e v ce e c be s very oc and ” rarely encountered throughout its range (Orz ell 1990; see al so Kral

u in the ulati ns i n the A n lina a i nal F r s it is har ima in ha S t dy g pop o ge N t o o e t, d to g e t t hi e i s i s rar ha i s un il it i s r aliz ed ha its ha ita en l n l a in lan i s t s Sp c e e; t t , t e t t b t , Op o g e f p e d , The i s m s i s wh the s i s a very rare communi ty i ndeed. b gge t y tery y pec e in the West ul as al Plain i s ra h r n ai n d r a hi all on the A n lina a i nal F r s G f Co t t e co t e geog p c y ge N t o o e t , with apparently simi lar habita t existi ng along the Catahoula formation eastw ard into i x Loui si ana and farther westward n Te as .

d t R r du i l A rimo nia in isa a ars to s a l in the s u ula i ns. I ep o ct ve y , g c ppe be t b e t y pop t o r u an rui i u l a a l n - ran i rsal h n a shru a r p od ces m y f ts Obv o s y d pted to o g ge d spe . W e b l ye r s u it i s a a l sur i in al n the r s d ed of r adsi s the ed s of g ow p, c p b e Of v v g o g fo e te ge o de , ge w rl h - f- wa a e d o e ine ri s o and i lin s. M s r ai nl it i s not i s but p g t y , p pe e o t ce t y , w e y Spec e , aus an hr ni a i i it n nl sur i s in r u i a h r its n s are bec e Of t opoge c ct v ty , Ofte o y v ve ef g w e e eed met: ar as ha mimi the n a ann a n i i n n r fire e t t c Ope s v co d t o s o ce c eated by .

On the A n li na a i nal F r s the s i s i s no m ans n n su h ge N t o o e t , pec e by e co fi ed to c r u ia: i u a n ef g t occ rs in Open savanna longleaf pinelands. Where we h ve fou d it in n un it i s al s a a r m the r a s in n slash and l l ll in Newto Co ty , o w y f o o d Ope ob o y p e lan a i n p t t o s.

MA NA GEMENT

The populations of A grimon ia in cisa on the National Forests and Grasslands in Tex as are the only populations on public lan d in Tex as or in the West Gu lf Coastal Plai n hi h ma s h m r a il a i al r , w c ke t e e d y ccessible and potent ly secu e . 12 6 P H Y T O L O G I A February 1997 volume - 12 8

The major consideration for Agrimonia incisa management i s how to maintai n the integri ty of the canopy and the herbaceous layer to promote proper light and Shade

li l ri m n al A . l l . ns an shru la r and h a r u h are r eve s A de e c opy , b ye , e vy o g ve y ke y det e t to l main ain O n n and main nan e . Tradi i nall fire r a incisa developme t te c t o y , p ob b y t ed pe f fi re ha a illi n od i n aders and redu in r un li er. In the a s n o bit t, k g wo y v c g g o d tt b e ce , n the an sir d l ls these condi tions probably can be maintai ned by thinni g c opy to de e eve , - h h in hru s and m in the rass la r. Thi s r dur mi bus hogg g S b , ow g g ye p oce e g t be x r lar ra s but mi h r ll for small ula i ns . e pensive fo ge t ct , g t wo k we pop t o

n x n hi n a u the one thi n fire as it in ra s i h B ut we know e t to ot g bo t g , te ct w t Ag rimonia incisa about whi ch we Should be i nformed best in order to man age this s hus un il the in era i n of s as nal fire and A rimo nia i s s u i li tl pecies. T , t t ct o e o g t d ed, t e a a m n definiti ve can be said bout man ge e t .

ds h r a a m n a rs l arl x ssi r un i s ur an and A s regar ot e m n ge e t f cto , c e y e ce ve g o d d t b ce herbicides should be avoided (see Kral 1983 ; Robbi ns H ardin 1987; Orz ell 1990; a Singhurst 1996 for further comments on man gement) .

A CKNOWLEDGMENTS

R er E . E ans E l is a i nal F r s s and rasslan s i n T x as and ob t v , co og t, N t o o e t G d e , Suz ann alk r ani s S a in a i nal F r s r ins ru m n al in mak in his e W e , Bot t, b e N t o o e t , we e t e t g t l r was und r a n as s - S har A r m n n study possib e . The wo k e t ke a Co t e g ee e t betwee the au h rs and the a i nal F r s s and rasslands in T xas . nn h R r s n t o N t o o e t G e Ke et obe t o , R er ral n i li n i nda a s n S n h rinda S am ob t K , Ge ev eve K e , L W t o , teve Ly c , Do cott , J e m r e nar all ai in ari n i A s and S l e us a s. lin har o o o , t ve o d ded v o w y Ge ev eve K e s ed her

ia R rt E . E ans and en Observations Of Agrimon incisa with us. obe v G evieve Kline r li i n the eviewed an ear er vers o of manuscri pt .

A rimonia in isa has n r r d r m Ala ama Fl ri a r i a Mi ssi ssi i g c bee epo te f o b , o d , Geo g , pp , S u h ar lina and r h ar lina T xas a ural H a o t C o , No t C o ( e N t erit ge Program A ar n l the r h ar lina r r i s an un ri d si r or am A m r s pp e t y , No t C o epo t ve fie te ep t (J e o o o ,

rs. mm li n rs. mm . It has en r rt d als for An lina as r pe co ; K e, pe co ) be epo e o ge , J pe , n and S a in un i s in T x as S in hurst i n Newto , b e co t e e ( g N x o S niffen collected

- A . incisa in A n rs n un n five ars a o but mi si n i de o Co ty twe ty ye g de t fied it as A . a r vi r l d in ha a p flo a . We ooke w t ppeared to be suitable habitat (A nderson County i s u si the in l i n the l a i n in i a d on h ir h r arium la l but r o t de p e be t) oc t o d c te t e e b be , we e unable

r l a it. l are u h rs for T x un i ha to e oc te Be ow vo c e e as co t es. We ve not seen a voucher for S a in un al h u h Si n hurst 1996 r rts it r m ha un b e Co ty t o g g ( ) epo f o t t co ty .

A n rs n C0 ° Nix on S ni en 3 4 A An “ de o ff 8 0 [ STC] . gelina C0 Ma cRoberts MacRoberts 3007 3 1 1 3 A STC 3006 LS US 3 1 1 5 DE K . a , [ ], [ ], [ ] J sper Co . z M Ma R be acRoberts c o rt s 2 886 2 896, 2 898 2 900 2 91 0 2 92 9 A STC , , , , [ ] , 2 899

128 P H Y T O L O G I A February 1997 volume - 12 8

Sin hurst . 1996 . The s a us nin ndan r lan s of as xas : Hi s ri al g , J t t of e e ge ed p t e t Te to c , l i al and h l i ta e al n s. MS h si s. us in S co og c , p yto og c ote t e S tephen F. A t te

ni rs i a h s T x as . U ve ty , N cogdoc e , e Gli tz enstei n W J f s as n S r n D . R. J . S . . . Pla t . 1993 . E al ua i n the s o t e g, , , t v t g effect e o of burn in longleaf pine forest: A cri ti cal literature review and some results for an temi s u ongoing long t dy . Proceedings of the Tall Timbers Fire Ecology

- n r n . 2 63 Co fe e ce , No V il T l air R. C. .H . R se O .H . . n.d. a i n the Gus En eli n dlif e f , , J o , Veteto eget t o of g g W e

Mana m n A r a. T xas Par s il li ar m n A us in T x as . ge e t e e k W d fe Dep t e t, t , e

. a i n f MS h si . ard J R . 1984 d o the dr u lan s in as T x as . s W , Woo y veget t o y p d e t e t e i n S a ni r i a S e h n F. A us s h s T x as . t p e t t te U ve ty , N cogdoc e , e P hy tologia (February 199 7)

TWO NEW S UP RAGE NERIC NA MES IN MA GNO LI O P HYTA

am R J es L . eveal

- . a t r al M an n r n H r ari um H J . P s n H l ni rsi of ar l ll Norto B ow e b , t e o , U ve ty y d , Co ege

- 42 . . P ar Mar lan 2 07 5 8 15 U S . A k , y d

A B STRA C T

The r r na Two new suprageneric nam es are proposed. S upe o der A cor a e i s n her in lud nl the nus A orus A ra a a m m r of the defi ed e to c e o y ge c ( co ce e) , e be am M usa les has n i n use for s m i m ut S ubclass A ridae. The n e bee o e t e b never

It i s alida d h r . vali dly publi shed . v te e e

KEY RDS : Ma n li h ta Ari dae A coranae Musal s n m n la ur WO g o op y , , , e , o e c t e

Su r rd r A coranae R al su erord. no v. as d on the a i n d s ri i n Tri pe o e eve , p , b e L t e c pt o of be

ardh A hor. B ot. : 1 33 . 1 un A coreae C . A g ( p 9 J

The ax n i s d n d in lud nl a sin l nus A orus . A ra eae t o efi e to c e o y g e ge , c L ( co c ,

A corales) and included in A ri dae Takht .

r r Musa l s R al ord. nov ali a a ull and ir r r n to the a i n O de e eve , v d ted by f d ect efe e ce L t 4A u 1789 as d s ri i n Musa a uss. Gen . P l 6 1 . e c pt o of ce e J ( g ,

The a name has e n in use for s m im and r i usl R al 1993 bove b e o e t e, p ev o y eve ( ) attri buted it to B urnett Thi s proved to be an error on my part as the B urnett nam was r s at a mi s lac ran . A r in l the nam i s alida d h r so it e p opo ed p ed k cco d g y , e v te e e ma us d h s who ish m l the nam y be e by t o e w to e p oy e .

urn line ain . 1 835 . Out s o B o an . n n r a ri . B ett , G f t y Lo do , G e t B t

R al J .L . 1993 . A r lim inar li s alidl u li sh au m a i all i d eve , p e y t of v y p b ed to t c y typ fie i - r nal nam f T ax n 844. o d es o vascular plants . o - Phytologia ( February 1997) 133 .

VAN CLEVEA (ASTERA CEAE: A STEREAE) : n=9 CONFIRMED

s m Guy L. Ne o

r En ir nm n al S udi s S am H us n S a ni rs it Tex as Regional Institute fo v o e t t e , o to t te U ve y ,

H un s ill T x as 77341 U . S . A . t v e , e

AB STRA CT

A reexaminati on of Vanclevea stylosa shows its chromosome number to be = a re m n i h the ri i nal un for the i s and it ma n 9 , in g e e t w t o g co t Spec e , y yet be = “ ” A confirm ed as a member of the x 9 Chrysotha nmu s/P etrado ria group . recently reported n=6 for Va nclevea w as based on a mi sidenti fied collection of

Isocoma .

R S : Vanclevea A s ra a l s s ma i s KEY WO D , te ce e, cyto ogy , y te t c

The recent chromosome coun t of 2 n= 12 for Vanclevea stylosa (Eastwood) Greene = Li 1 3 n radi d an earli r re rt of 2 n 18 S olbri et a l. reported by Lane ( 99 ) co t cte e po by g, A phylogenetic study of North Ameri can Astereae based on c NA r s ri i n sit mu a i ns an ar a l. 1996 ad d n d n ha the new r r w as e t ct o e t t o (L e , ) de co fi e ce t t epo t correct by placing Vanclevea Greene (a monotypic genus) where it would be ex pected with a base chromosome number of x=6 (withi n the subtri be Machaerantherinae sensu Nesom

In a subtri bal classification of the A stereae (Nesom I di d not accept the proposed revised placement of Vanclevea and called for a recount of the chr omosome num r M i i un s 2 n=9H r rt h re r m n l ll be . e ot c co t of epo ed e f o ew y co ected materi al

n rm the ri inal un as rr c for ha s i s. Fur h r an x amin a i n co fi o g co t co e t t t pec e t e , e t o of the u h r for the un 2 u= l 2 sh s the ll i n a s i s of Isocoma vo c e co t of ow co ect o to be pec e Nutt . ra h l ea t er than Vanc ev .

Fl r u s ula i n ll i ns al n i h l a ma ri al for owe b d (pop t o co ect o ) , o g w t e f te molecular anal sis r ll e and s n me r m hr ula i ns Va ncleve y , we e co ect d e t to f o t ee pop t o of a i n north n r n im th la i um ura r al A riz a T o A ss . H r ar at r h ce t o by o y Z , t e b C to No t ern A riz ona U n Pr ss u h rs for of the ula i n iversity . e ed vo c e two pop t o s were i ncluded with these mat ri als the u h rs ha n si at the h r ari a n l e ; vo c e ve bee depo ted e b oted be ow . The leaf materi al has been sent for molecular analysi s to David Morgan (Western Washi ngton ni WWB wh s r rsi h r . o i in i h ail d U ve ty , e b ) , p oceed g w t det e studies of th e Machaerantheri nae and ri h rall r la ax a the ster a pe p e y e ted t of A e e. Nesom : Va nclevea chromosomes 13 1

ITE STA TES . A riz na: ni n un . l n an n R r a i n A r a UN D o Coco o Co ty G e C yo ec e t o e , n US 89 si h a uar rs of the NP S GCRA i n Pa san un s on the si of o Oppo te e dq te ge , d d e de d 4 1 u us . mmo n H a 1 1 2 76 i h T . . la he w l ca. 000 2 A 1997 H D. N t h y ; e ev ft , g t , w t Zo

KA N U WW B . ( , )

n f a STA TES . A ri z na: ni un . US 89 ca . 9 mi S o P mil UNITED o Coco o Co ty ge , e

3 on the a a R s r a i n san un s on the E si the hw l ca . 5400 5 8 N v jo e e v t o , d d e de of y ; e ev ft ,

u 1997 H a mmond 1 1 2 74 i h T .N . la 2 1 A g , w t Zo

UNTTED STA TES . A ri z na: nin un . H w 98 3 mi E of int rs i n o Coco o Co ty y , e ect o i h Hw 89 n ar r lan sand ar a on S si the r a l 4000 2 1 w t y , e powe p t, y e de of o d; e ev ft ,

A u 1 7 T M o la i h u u h r . g 99 , Z (w t o t vo c e )

Meiotic counts of 2 nz 9H were obtained from flowers in three heads of each cl v a n ui a l un s r m all hr ula i population of Van e e . U eq voc co t f o t ee pop t ons were i r h and a has i i observed in configur at ons at p op ase I met p e I . Me os s apparentl y i s synchronous or nearly so am ong the flowers of a single head and pollen formati on The ri nal oun for v apparently occurs over a relatively short peri od. o gi c t Vancle ea “ ”

S olbri et a l. 1964 was al ula ed r m s ma i un s An derson 1 976 the ( g, ) c c t f o o t c co t ,

u h r r m nin un A ri z na. vo c e f o Coco o Co ty , o

Va nclevea of Lane Li

The voucher for the count of 2 n= 12 by Lane Li ( 1993) was made from field ll ed ma rial c ll t d r m an oun ah mi E ana 13 J ul co ect te o ec e f o K e C ty , Ut of K b , 1989 La ne 32 47 It i s a ll i n of Isocoma r usb i r n its i n i , co ect o y G ee e , de t ty ral ri l ran h s are m un confirmed by vegetati ve and capitular features. Seve ste e b c e o ted - a v li n ar- tri an ar in on the sheet; the leaves are narrowly elliptic oblanceol te ( s. e gul

- - Vanclevea sin l in vs . dis in l arall l in d and l sel as ndin vs . ) , g e ve ed ( t ct y p e ve e ) , oo y ce g (

s i s r adin or ar ua . The h ads hi h are not nn d an the s ms t ffly p e g c te) e , w c co ecte to y of te , ’ are old and straw -colored (presumably from the previous year s floweri ng) but n a h - v l n - un ula and s li r m s and arranged i s ort pedicellate cluster ( s. o g ped c te o ta y to cy o e di stinctly pedicellate in Vanclevea ) and the involucral bracts are apically rounded or lun and r v n l an ular a the an t n shar l ur b t e ect ( s. arrow y tri g t apex d of e p y rec ved) . None ’ of the old las ar mai r A t l as un u s ( t ye s) florets re n in the involuc es. e t yo g b d r suma l r r s n in the ula i n h r al h u h n n are i n on the p e b y we e p e e t pop t o , oweve , t o g o e ev de t h ran Fl ri n an h is . rus i usuall ins in A u b c es of the vouchers. owe g ( t es ) in 1 by y beg gust

(Nesom 199 1 a) .

The diploid count by Lane Li for Isocoma rusbyi corroborates several others for ha i s hi h = a Th ir h ra h of s r r rt as 2 n 6II see s m 199 1 . t t pec e , w c we e epo ed ( Ne o ) e p otog p 12 mi totic chromosomes (published as Fi gure 1B) apparently w as made from a tapetal ll aus the au h im li ha all un in h ir r r r ma r m r ce , bec e t ors p ed t t co ts t e epo t we e de f o flowe b u ds .

The m l ular la d Vanclevea l s s anal si s an et a l. 1996 o ec y by L e , ( ) p ce c o e t to Lessin ia h i = r u am . X lorhi a r n and H a lo a us ass. i h n the x 6 g C , y z G ee e , p p pp C w t g o p but a ar r m Isocoma The sam l anal z for h ir NA s u i s i s r m the p t f o . p e y ed t e c t d e f o ula i n u h re n 4 DNA s u i s on the su ri a i M r an pop t o vo c e d by La e 3 2 7. t d e bt be by D v d o g 1 32 P H Y T O L O G I A February 1997 volume 130- 133

i n in u lear ri s mal ITS ari a i n rs . mm . sh the sam n i si (us g n c bo o v t o , pe co ) ow e e t ty po t o ed wi thin the branches of Isocoma based on hi s sample (the same sample of Lane 2 47 us d an et 3 e by L e ,

Vanclevea phylogeny

4 asin flori sticians su h as lsh et a l. 1993 and r n uis 199 ha Great B c We , ( ) C o q t ( ) ve maintained Vanclevea as a monotypic genus without comment regardi ng simil ar or

lated s ci s. A nd rs n r h r had r i usl possibly re pe e e o Webe g oweve , p ev o y concluded that Vanclevea is most similar to species of H esperodo ria Greene at that time regarded as a monotypic genus .

My interpretation has been essenti ally in agreement with A nders on Weberg : Vanclevea i s most closely similar and presumably closely related to species hr t (particularly the two of H esperodoria ) of a broadly interpreted C yso ha mn us Nutt . = s m 1 1b all i h a as hr m s m num r of x 9 . hr sotha mn us (Ne o 99 , w t b e c o o o e be C y in Hes rodo ria and P etradoria r n S tenotus u . and Vanclevea r ( cluding pe G ee e) , N tt , we e “ " placed together as the Chrysotha mn us group of the subtri be Solidagi ni nae (Nesom Confirmation of n=9 as the chromosome number for Vanc levea indicates th at a molecular reevaluation of its phylogenetic positi on based on the newly collected materi al will be interesting .

r a a One other species also belongs with the Ch y sotha mnus group . H plop ppus

w h l ne t ls. microcepha lus Cronqui st as too asti y transferred to To s us A . Ne (Nesom

M r an 1 90 but was rr l laced in the anal sis an et a l. A m re o g 9 ) co ect y p y of L e, o appmpriate nomenclatural combinati on has not yet been supplied .

Vanclevea should fit (as a prediction) in the exact position where the analysis of

an et al . 1996 la es Oreochr sum a rr i A . ra R d . i hin h ir L e, ( ) p c y p y ( G y) y b w t t e “ ” P etradoria group (their Figure My subtri bal classification (Nesom 1994) places Oreochr sum in the S olida ini nae but i h an s r a i n its rela i is l y g , w t ob e v t o of t ve o ati on hr ha mnu al s i among putative North A merican relatives. C ysot s o i s posit oned in my “ ' ” arran m n am n n ra the P etradona r u Fi ur 3 an e l t a . ra h r ge e t o g ge e of g o p ( g e of L e , ) t e

han am n h s Fi ur 6 of an et a l. t o g t o e of g e L e ,

The m n i nus Columbia doria s m Columbia do ria ha llu A . o otyp c ge Ne o , ( Gray) ’ s m was d s ri as a m m r the hr sotha mn us/ P etr Ne o , e c bed e be of C y adona group (Nesom l 99 1b in the su ri S olida ini nae. a r it was r m r m hi ) bt be g L te , e oved f o t s group and placed as a basal element of the subtribe (Nesom Columbiadoria w as i na r n l mi d r m the a c un in of the S lida ininae 1 4 o 99 . dve te t y o tte f o c o t g g ( , p

A CKNOWLEDGMENT S

I han Tina A rs an d Tim la rth rn Ariz na U ni rsi Fla s a t k ye Zo (No e o ve ty , g t ff) for their work in arranging and m aki ng the collection of bud and leaf material and vouchers of Van clevea and the s a of A for all i n me x amin the ri i al u t ff K NU ow g to e e c t c vo cher. Nesom : Vanclevea chromosomes 1 33

LITE RATURE CITED

4 A n rs n L . C . P . S . r . 197 . The ana m an de o , Webe g to y d tax onomy of Van clevea 34: - A s ra a . r a asin N at . 15 1 1 ( te ce e) G e t B 60. 4 - r n uis A . 199 . In term oun ta in Flo ra . lum 496: A s r al s. The C o q t , Vo e te e New r ani al ard n r nx r Yo k Bot c G e , B o , New Yo k .

M A . . Li . 19 3 . hr m an . 9 s m num r r r s in L e , J C o o o e be epo t Compositae with m hasis on ri Astereae of the s u h s rn ni S a s and e p t be o t we te U ted t te Mex ico . S ida

- 546 .

M . A . D . R . M r an . S uh an B B . i n . S m s 8c R. K. an s n . L e , , o g , Y , p o , J e 1996 . R la i nshi s of r h A m ri an n ra of A stereae as on hl e t o p No t e c ge e , b ed c oroplast DNA i a i nd r s ri i n s d a . In : H D . J . . B eent e N H .J . m e t ct o te t , j Co posita e:

- stema ti s . 4 77. Pr . Intem tl S 9 . m . n . 4 1 . y c , pp oc Co p Co f , Kew , 99

s m G. L . 199 1 a. Tax n m of Isocoma A st ra a : A stereae Ph tol Ne o , o o y ( e ce e ) . y ogia - 4 1 1 . Redefmiti n 199 1b . o of H esperodo ria (A steraceae: Astereae) and the s r a i n of Columbiadoria a new m n i nus r eg eg t o , o otyp c ge f om western North

- Ph tolo i a 2 1 . A merica. y g 5 Tax n mi n 1993 . o o c i frastructure of Solida g o and Oligoneu ron As ra a : A stereae and mm n s on h ir h l n i i i t ( te ce e ) co e t t e p y oge et c pos t on . Phy ologia

- 44. S u al l 1994. btri b c assification i n the Astereae (Asteraceae) Phytologia - 2 74.

m G.L . D . R s M r an . Ne o , . o g 1990. Reinstatement of Tonestus (Asteraceae:

A stereae . Ph tolo ia 68 : 174 18 ) y g 0.

S olbri O .T . L . . A n W C rs n D . K h s . P . H . Ra n . Ruden r o bu . 1 4 g, , de o , y , ve , L g 96 . hr m s m nu - m rs i n m si a . V . A ter ae r C o o o e be Co po t e s e H . Ame . J . B ot. 5 19 .

lsh S .L . N . D . A d S . ri h L. C . H i i ns We , , twoo , Good c , gg 1993 . A Utah l r F d. 2 r i o a e s d . Prin S r i i ham un ni r i s r s Pr ah . ( , ev e ) t e v ce , B g Yo g U ve ty , ovo , Ut Phy tologia (Feb ruary 1997)

ND X TO A UTHO RS O LUM 8 1 I E , V E

- 6 Al an r I ur i . 2 73 V ibran s H . 3 9 ej d e t b de , G ,

4 W am ock M .J . 15 1 A har M . 1 5 t , ,

ni z R as . 382 Be te de oj , C

l m ns . 69 C e o , D

’

D Arc W .G. 382 y ,

Denharn, RA . 5

- Es arz a S an al S . 2 73 p dov , 4 Hensold . 2 , N

- H rr ra A rri a . 46 12 4 2 73 e e et , Y , ,

Hirsh r . 69 be g , J W 5 H lm s . C. 38 o e ,

a rri er J . E . 12 1 L fe e,

MacRoberts B R. 2 8 35 , ,

MacRoberts M.H . 2 8 35 , , 365 McIntosh . , L 2 2 M a h r W L. 8 e g e , .

M ral s J .F. 36 1 o e ,

R al L 42 J . 1 eve , .

- 4 R sal s arrill . 6 o e C o, O

il a 2 2 S . 3 b , J

- 4 Si u i r s l a ME . 12 q e o De g do , mi h B 44 S E . t ,

T rr ll E . E . 108 e e ,

Tri l 348 p ett , K . T urn r B . L . 1 5 10 1 6 2 2 103 e , , , , , , , 1 4 4 15 , 1 18 , 32 9 , 333 , 3 0, 3 3 , 348

W AILABLE PHYT OLOGIA MEM IRS lum 1 1 NO AV , O , Vo e

THE COMPS O F MEXICO , V o lume 1 E UP A T O RIEA E

B .L. TURNER

- - - 0 2 5 2 0. 2 72 . illus. i a 964 08 1997 8 x 10 i n . , , pp , ,

i i ated ten lum seri es on the A s ra eae of Mexi co hi s The second of an ant c p vo e te c , t her lum s ill in lud the r mai n r of the volume includes the tri be Eupatorieae. Ot vo e w c e e de e x i an r r n a i s the 2 700+ species of A steraceae known from Mex ico . Th Me c ep ese t t ve of H ni l antheae d Eu at eae ami l are art i ular l ri h i n the ri s ele eae He i an ori . f y p c y c t be , , p

ia M m irs lum 10 e m s of Méx i vol : imi d su l Ph olo es Th . 6 L te pp y of yt g e o vo ( Co p co , Tageteae and A nthemi deae 9 (The Univers ity of Texas Herbari a Type R i s r and 2 A S ix h S umm ar th e r na a A i nni a a eg te ( t y of Ve be ce e , v ce ce e , S i l a a Dicrast lidaceae S m horemaceae N ctanthaceae and Eri aula a t b ce e , y , y p , y , oc ce e of the rl as alid Taxa ra hi is ri u i n and S n n m - are als Wo d to V , Geog p c D t b t o , y o y y S o l avai able .

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