Annals of the Missouri Botanical Garden 2009 * Volume 96 Number 1 '.'■.■'■'.■■■ • (i t ri ] )i • the Garden vviJ for information to Authors are printed in the back of the last issue of each volume and are also available online at Editorial Co C. Hollowed Editor, itanical Garden Belh Parada ig Editor, Allison M. Brock e Editor, otanical Garden Tammy Charron \ssistant, Missouri Botani Cirri Moran Press Coordinator, Roy E. Cereau Latin Editor, oldblatt Missouri Botani IcPherson For subscripts Garden, % Allen Mar keting & Management, P.O. Box 1897, Lawrence 8897. Subscription puce loi 2009 i $175 per volume U.S., $185 Canada & Mexico $210 all other countries, lour issues per volume The journal Novon is included in the subscriptioi Tin: Annals or tin-: Missoi ki Bdtwhcai. G\hi>ln 26-6493) is published quarterly by the Missouri liotanical Garden. 2315 'lower Grove Avenue. Si. Louis, MO 63110. Periodicals post- age paid at St. Louis. MO and additional mail- ing offices. Postmaster: Send address changes to annus or mi Missoi hi Botwicat. Garden, % Allen Marketing & Management. P.O. Box 1897, Lawrence, KS 66044-8897. stract/Global Health databas, „ iu<» i 'OK, Research Alert®, and Sci Search®. ..'.:"■.. ! ! ■ www.bioone.org). © Missouri Botanical Garden Press 2009 The mission of the Missouri Botanical Garden is to dis ® This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). share knowledge about plants Volume 96 Annals ^j||k Number 1 of the ^"5C 2009 Missouri Botanical Garden THIRD INTERNATIONAL Petra De I M. Taylor? and RUBIACEAE CONFERENCE: ^ Huysma,^ INTRODUCTION 1 with worldwide distribution, but most of its diversity is held at the National Botanic Garden of Belgium in sWematists, ecologies, and conservationists at a basic problematic tribal and generic complexes; the full level, and their important role in these tropical proceedings were published in 1996 in Opera and intensity of this ,th increased 2005, when a half-da. symposium focused on by conferences spe, amily. Rubiaceae during the XVII International Botanical The First Intern;, ference, held Congress in Vienna (no proceedings were published). at the Missouri Botanical Garden in St. Louis in 1993, This symposi i : terest in and need tropical groups for the first time; selected proceedings The Third International Rubiaceae Conference was were published in 1995 in the.; . subsequently co-organized In the Katholieke Uni- Ann. Missouri Bot. Card. 96: 1-3. Published on 23 April 2009. Belgium from 18 I . et tribe. .. ■ :- '.-:'■ ■■■:.■ concludes that the " fossils (i.e., ninckx et al. enetic relationships those useful for d; are from the within the he; macoceae s.l. based of the family. leaf and petit acteis and e\aluate netic evidence, and reproductive biology of the Rubiaceae and show the taxononuc effoits in this Neotropical tribe Posoquerieae, and cites their f ted here shows that A REVIEW OF MOLECULAR PHYLOGENETIC STUDIES OF RUBIACEAE 1 mam molefubi prnlo:-,. ti< -h.li.- Aan- iV IV P. . .u.i. 2001 2002. lih.i et al , 2002 Wln-cn ..' ■ ..'..' :.-,•■.•• summarize the mail :iave not yet been tested by «n Miuii. , ,l ,« ti i i . i It t ml ..!>].- lI i . it hi 1 , ,, In Jiei numbers of Coptosapelteae. The Cinchoneae tree has a low q levels, from closely Jo L< H . on 23 April 2009. aNScr^ 1 — Posoqueri ' — Cremasporeae Bertiereae "Gardenieae" Pavetteae 1 Coffeeae 1 Alberteae relatedspeciestotlRvli, Mii.nl i < M ,t f,.i ihe l,,,i Pun, i in Rlk.nsirihik.ns analysis of restrictio r studies have d The first attempt to reconstmct the Rulnaceae been used, seven from chloroplast DNA (cpDNA) 1991 . Bremer and Jangen in , 1|e imenc „ n ;„.,„,„/ 1 * ""' "itk /../ s/' nil / S ip>i ]ht ii(ltj | J)nl „ slll1J ,,„ u „, (,l and six nucleai DM ll f- IH ...mtiansoibed of cpD ^A. I.arl,,!, Km Ifal mH.„„h hn. .. , , puiposes olhei than pin 1. - n ,,| Of these 4400. Btidson and \ t ,< nlil<) x 'i m'P.Iu , t This paper is divi, .rts. The first /,-„ J acq ., Hoffmannia Sav.] and Ixoi : ■ ' '" • • ■ cl.-Ml" ,1oi (I, 1 .use il 'Mil, Ih.Mi.f u. Vi I ,< ui'Koiil e , ,pectneh J 1 111 < IK ! u I Jl m 1 f / „ i, i, ,„ VHu 1. ,,5i. credibility, except h 0.92; the in Bremer et tudy included out- Tl» Ii „.< a in. ... ,,u ,i i-i«, eut, il n n,J V I- ],' e(01msltdd,tal.l<)9?)andani.iphol„g, J taxa and thus could not be tested for monopliAl • .1.1 m | „ i I , an , , m,(, „.di„ I (F ., m ■ ^ xou le si (me lulu l . Jtei I n 1 in, I i, I former ' mr] i..n.K,. „i J IJOl'l], Km.Lmiin ,7-U-1h. IjM -•■•.,•:.,. .. ■' ■ . the iamih, and the genus flz/ifon-u Bullock «as -. ". t<.- ^, c.l I »» , < ,, 7^ ' : . ■ .••...■ ■■ ■ • ■:..■■ ',-■ lie, (L > //r//^,li ,# ' ' < 1 tf» i ml I nl.J( 1(1 i.ul • ikttn leae (scnsu Rob- ■ -^ : ■'. : Rubiaceae. In the study, 43 Rubiaceae genera p Antirheoideae fide .. .. i. ■ .:■• ...■■■ ■ ^ -v . v Cinchonoideae, and I >kU v u .11 1 , i. .pi, Jh.. deletjne) <7i7oro DC (close to Ham/ lie --Hi Uw » Si| neeae), Neo- blakea Standi, (close to Guettardea.-Kondeletieae), Rova et al. (2002) performed a phvlogenetic l'huil.mlhm Grisel,. iChiococceae-Cal,-) ;eae), Phylla analysis of «mL-F f . .mn "» »K ,h ui r„ b .k,L to //,,/, .,,n/.,„, U,,.„, <.ys . I \mj-1iT tf|".»,. -,1 ln„ -u Mart. & Zucc. RichartUa L. [85%1). The tribes B m- il'-"'!., TIi in W <' ' j pi di.i.M. h. in agreement with Nepokroeff et al. (1999). lar results. The tribes P / ' / \ , il, , -t „ 1] ( , , Palicourea, Spe lamlia L., La- lii il ,,',,'< 1 ' Hi i^.f^n-lnp |.^.,i t l„ i.„i , We- Natali et al. (19 equences to the ih- a ..Ll.^itl.t'ul .ufUi-^uj.thm th.w hi mil, ■ '/»'// .. .. IV .1 li ,J (1 95) divided the V ,.,/('. !«! i\bi n m«' M.I ill U<»>«>> , u, ., I sp,^.-il.,...n.-Pi.-, fl it,!tnes of the whole They rooted the published tree between family Ixoroideae (Coffea and Ixora) and the rest. , ,1 '..Hhn-p, ,,„m.\ 1 • o- H< ',..ii I M,.mV,. 1 \<1, i .-1. 1. '1. f.lii' f liimti Th t1 , mi m>1 ,.Kl.is...n is that the lack Ill h siil t . ,nl ( ( ,i ulu b ' ip, [,. u« the, u lu'ug e.> (aiapithti \ubl , (ha < ', also analyzed the R chotria s. str. ( = subgenus analysis in Natali et al. (1995) but divides the et al. [1999 clade (including , ■ -,■:. ■ ', :■■.■...'.■ Nie et al. (2005; le genus is Carapichea was reestablished as a genus by mi -li.tt-.l U tit tn I tun. 1 is i okioeft et al belonged to th- -,„„ j „ ,, flu (1999). They analyz and rbcL. The eluded in the is Carapichea were . • ' :.'•,•.-•■..■ was also shown tha tube was monoprn- member of the Psychotrieae but closer to Coccosypse- letic. In a sec Inthospermeae taxa I]. /, >-.„.,/ .mui.lt s The it suit tht mtein. was very much ir: .kroeff et al. of Anthospermeae formed a monoplr, letic but ■vcakh inni .L.I. i-ilh <,,>),,„<_.• -.11.1 i -i.it.lr ,-n , lu-le, lie ii.m mi ffiastom, It was excluded. The latter was instead sistei to the also suggested that the ancestial aiea of the Lewis, Stenw 30 taxa altogether. .piosminae kept distinct id North American :,..■■ phiasma Biemek and />•,>, .,, ,s \\ ndh and loimd impoitant lole loi hi, ton of dneisificahon The .. ,■: . . -..;. . ..-■ ; til ..... lil', t >>H, Fill ■• i- il-o ..ntiim.d ],, i m. .pliol.^K ,1 and y ... he 'an Hi..,. I ml. * illiin a.,.1 study by Thulin ai ,pecies. The^ found a clear association s].,iila ! ,tusl„f. 1 .,n th, c,,,,,,, i ' ■ 1,,,!,. i-snli .,-,( Hi" rhao.i, ,n i tu.l. .... i.l
  • >„ imtstigat-d ljjlh :•'!,> P Pi / <> 'i,«i< k = r„ ,, ^/„,,k SIhm.HU I7,..s t »,. ,,„,„!, ..„ JItlll I.itutl., ,s , hs , ltl( Hoc!,,, Ou^/WD< flm-olo, ,„' «. , x , 1]l( lux ,,„ h „ ,,,„!, , ,„ „,. I Kruusszu Han., Pe .., Tarenna. circumscriptions of th as et aLj 1998 ). !'^ t . W< 't, ,l„,Mn hd W.n.l r' 1 ,L 1 "|M7, I ', 1 ,i n D ! ,i , | l ,'| , |i t !!' .e in.Vni, i - I( ,l,. ,ithu.C~i,l.i. Il.li, !'».%, i~ , ( t.Vr!>,i.',,J,| ',,,,, Jitl.lMu*,. lu'lth, -.n, yueriu, Bertiem. h d genus belongs to Rubiaceae in the subfamily ..... *..'. Vlbeitea (ilbutaE Me S ) i a, li ,i . > ti if s^n.i I Lili P. s , . 'n'Od) continued his ....... umesohed plrdogeny of the group; he also wanted to and A. hispida formed a well-supported clade at the ■■,■■■, study (Persson, 199 :L the taxa. ■.■■■■ : ..,■■:■ • • " 1 1-'~, tLir "iKr) moi-.pkjtii. pi. -up !L.-,dl to the Andreasen, 1997). He further identified a core h frican clade (CaZo- , .,j,ln „., - ,1 a'hm Benth ami r« U |in« pu. s ln!', J u. )•) .h ,s 1- nh dtmonsti alt d that »'' „i id > nli i » I -in, ij it mj i, , , , r p, -nun _ioi lit 27 genera of the with tetrad pollen. tribe (ITS, trnT-F, and morphology ). The) found m.I d.snn IN d.vj.l .1 int, Uu inn d.,', „,. ,., /„/.,>'('>,'. Hot i -.- (Vell.)K.Schum. In the combined „i. l>,i, 1„. '< Caiaio, Pcponuhum (Baill > litnt- i' : .'•■'■.■. .. ei h.tl.i in ^rm , d m | ltl *,j, ( „„!,{! h.,1, , „],„„ t() ' [,,u t< .1. study (Lantz & Bremer, 2005). Sixty-six of the C zed trnT-F and ITS estimated ca. 180 lyzed for the data and den tribe Mussaendeae s the Ucinin of Mrectaria. The. also performed in the tribe by don rbcL and one of rpsl6 (mainh sequences from f.ill.Ji -it I) L.U...S- Iumis mil. .1.-1,-1- ami the . ,., | hoU i, .1 ,m, -tip-in-m uei. calyx lobes (with exceptions). Taxa of the tribe In a sense, the : ■ ' :: . ^ ■■■■.■ ■■■■ -'■■■■ \ndeisson, 1996). In a study of rbcL data from d ;mended to include i in, -<■;•. ,. uvmoplrMrti, suggested In Bremer, 1992) than to the rest of the species of ' sum and E. may- 's s. str. was nense). monophvletic without the Portlamlia group, he Erithalis is an endemic Caribbean genus, the included the Portlamlia group in the tribe Cates- 1 )2). They investigat- baeeae. Therefore, the rest of the Condamineeae ed seven of tl es with two nuclear mdtk.n-. JO ...„, FI> Tli,; t'» , / - N seim, P-.emlv.ah- Lamb.) Roem. & Schult. Hook. f. salmeoiJes Correll, £. offon/era Jacq.) that were ed In And- o pinpointed the ,al..-t,, l v ,i',s, 7 ! r„. ,u<.i ..,1 Hiiilm M'»<'.>) h. .Ii«,i,«! > f in tribe «eit hist anahzed with nioiphological chaiac- - .1 i-t-le*' instead belong to ,ii('Hiii.'iUbtiu- ' ' Il d,ie. 1,1 unl se Lt „,i,^t'l -uM b„ ...111 ( 2M. | i. in (,»ated the leiationships of Rids hi ' i r d <> < ., ', >u in, I hWuI.mk > : • . : : " ■ ■■ Hill., . |',w ,/, t tin J F< I Is mr t [\ 7 ,, ph,!, j, U llt „, i M , (l vi t n 11, _(\r, t , „ H , , lt Ul ( , » , u j i p „,i I ,|t ,i, ■■..:.. to fimbriate wing maigins of its sn .1- " I ^,-i >, & oj th. -nbtiibe \,!,nae with • . » 2005: 26). (see Hon) « :ated in a study by Tube Iseitieae was fust anahzed In \ndeisson Razafimandimbison et al. (2005). They analyzed ITS ., "mu-j.1 /V, ;,..x;o,ilh- Africa, and tropica v. The sister Drupes were ceanic taxa and on s ', , Mi K , i il, t ih I,, tl . i i Hi i .nun I Ik,. , ! ].. otneial enhance. >f \ariation in species richness. Instead, certain three species as monophyletic , bur ,t , is i„m,d to !■ r„nii.m , lions ot dispeisa] mode oi life foims «eie • :. disl m i « G ne i ' • ' ■-- ■■ . : . .. . ■..;.. as a M,K,|ii,,« t a ;.lolul .shmaie .m ailhiopo.i it can be used fo ly, species. -.,:•..■• , • . < , (K.Koi,,],!.,^,) ,,1 Ik It- ,a,o,i w li,.-e N..H, Jre...- a (on.paiame . oule i W ,rl. tl i- -piciesn.h and species ,l ( /„m- /,v /._'■'. (1 n- aries in this gi ithin a species come Bremer & Eriksson, 1992) in which a Ri enetic signal. flisp , ~d |„,is ,.l,ui., i.llni,.-) hm , unihi ! V, ,m, I),]... f, .«- liul .1,! < 1 ill ill lln.e 1m )-« ■ ,■■ ■ ■..,..■■ M s, j,,. , l.».,h. .Im, .ana „hh.N d M a., «,. . 1 ,J0 >^) concluded that bers, and corolla col< origin and that the Mi,.- „ i; '. -, , . i > 's, /„• /. U. ,, (, ,1 |,.M .. of different fruit and seed ft pes and the al.ilin to colonization and dispersal. Both parsin.om and complex. Motley e :r species from Hawaii together with eight extra- ■ ■...':..:■■ ..... distance dispersal. • i .20H, ,. K,n,„ I ., 1 ,,. su.m .1 ,,,:„,-.,! J,!*'*^'^ L^K '^m. d and ^IpLVogKaJ v ' j.IimHx |,i„ ,, .. .1 ,2>*M ,. -sen. ,1 nut-™..] ^i"^"' I.TllT.V.'! i"",,,! ""i,. u'l •I t. m ,11, s„ m ». .«,,! m ..II. «, 20.1 , |K „,..,,. , alc themolrtll i al eeae, Lasianthus group, Morindeae s.L, Pauri- , ■ ' ■ ■ 2002). Mitova et al. bid patterns within '1 .1 Ul '. n.-v \.,.,1 .s.in, vX s,,,, ,, u . L . mlikI ll.jl <•,..-„ t-ol -„!.!. nle-, 1 hr I,a,,-H ,„, ,,,,1,1,1, plnWneh- .Lit.- A ill, ^.m,, tribe Gardenieae also di et al, 1996). I ' ie, 'fail,,, , j,l. n.^ln-.l *! lid,- , ! ,.lxe. i Ps^chotneae, and variable in Argostemmateae. informative th cters are as the PCA I _»)<(»,> lu.asmed cncentiation of onh sho«s smulaiities between species, «hich can pattern «as foi aluminium, and theie «as also a f, ared to a nonphylo- Volume 96, Number 1 Bremer 23 2009 Molecular Phylogenetic Studies of Rubiaceae genetic tribal cla ht, 1993), so it is Literature Cited RublLr l/pu's -ib^rU^TTb.d 93: 103-121. i »lu. . hH. , ( iR , , , i [ r «._ ,u- b bb'^i.t i, 1 w !Zi»*ZL\ r\„Mi .".'."r h,i 7 1?'»_1(,4 Fnrtl.,.-)...U.,ih. -t. «!i -..{.]ii r M,Mt] oipioblen ^ : '= \ (\clullt tt a], 2006), Isuhotua (N.hmn ' ' ' ' ' 'l 2007), and %>a/< |ogpm of thp tub eae 17 " 28 - id («,,t ...V, , .1,1, ml l.i *, btimil - s/lLMy"*' J,V ' u n rh< i i .u i I > ibt ..ml I . if! . " ] ' 1 1 I ^ 4 i ' i f r t 1 f-. I„.'i,i. 6»/,.,»i, /. • ,. f,.;,-7 fl . .,V ; ^,f . mut chattel- Ehi.iidorfei. F..1 F. Manen & A. Natali. 1994. cpDNA 245-248. 1 <>J r i ! h lu'.i nl r x > 171-1184. Bul1 - 55: 85? ' I . js J - i, „ . I, ■, , > .(I. V i~ M. hf ™ i D 01 .,,, 1" te). Rev. Pa aeo ot. omy o t e u mceae. 51-1076" , L. B. Smith I & . 2005. PIts legem of the comply i. Missouri Bot. Card. 80: Lillo'a 17: 243-288. ;. H. & A. P. DdM 2(fir V,u .,„. 7 v, ii tl I! -Uf Rubiaeeae. Bull. Bot. Etat 28: 200-281. a, J. H. E., P. G. Delpre A. Albert. W REVISION SINOPTICA DE Eha L. CabraP GALIANTHE SUBGEN. GALIANTHE (RUBIACEAE: SPERMACOCEAE), CON UNA SECCION NUEVA 1 ' 2 ,„»i J<)..,,mm-, <\, .listnl,.,, ...ntioj.i. A^ snl.u.i,,!. J I. ■'A.i-.i-rju. A U.-, Jii'K.' <...., fio„«zu G Me^ ■ ■:■■■■ fastigiata Grispb., designado como tipo dpi gen , n i,' ( < B ',!)■' 1] i» Hit HBP. IV IT'J [i N If i ItB k III J P I PP \l , MjM Mi \-< MO M\FV MAM N\ OUPR P P\( \ PI R RB il sp SPF 1EVLL UB I'ti !'P< I. !'- U»/ tie G. centranthoid.es (Cham. & Schltdl.) E. L Labial & B, . r jluiv 2t)0l . ...... tm t verbenoides (Chai ;eb. [= 5. aladas y granos de polen zonocolporados con verbenoides Cham. & iuras multiples. ■ ■ .■ E. L. Cabral (Cabral, 1991). curso de la re\ision de la trilm Spermacoceae Schumann (1888) no acepta la propuesta de e- i L. y Borreria sect. Grisebach de cons .teres no se ajustal.an a la Bonena romo sect Galumthe (Gils. I. ) k ^ Imni . „n V,/>. ,H, Kl , i 1 din .i„ a ,., .,. . , (MM!,, b i. Con el andhsis de .' i ,i hut. »nt. ii .1 u.u i. n . \ , , „ , „'., i , -t ^ v i<> In i Sorreria s.L, se o riirhs, como el tipo de inflo- Borreria sect. Galianthe. ■ 1 H _, t „ ,„i », 1 , 1 , m,. . fl)l i I I | t I.,- < i) .< I mo, kiehn, 1080, 1005; '•, .-:■•• ,' 1086, 1005). Postenoimente Damn i L l.i il I TO I) Gahantlie Giiseb SmiiI. Fl. Argent recuentos cromos. i Galianthe. Los siguientes < .ales: inflore- Galianth Publ. Field Colum- t -!• n> Hum A —. JiJl.JJ si monudoiK peitmbada F\ p. Kii..lm iile ,< •. i ui! lofih, ci -1 )'« AnA .,-/.,'..,-, , , (. I.i.o (■ h.mi X > Mtdl) E. L. Cabral). Galiaiillic subg. Galianthe E. L. Cabral, G. reit ', souzae E. L. f 1 , lU ' ''' '" ' M """"I"""' f|] h,u,< hnl.,^ |k|,s smill jwcuMiK ladas 'estigm°a 300 ° m ' Fi § ura X ( A " F )' 3(2). Sufrutices con l-2(-5) talks de 0.4-l(-2.5) m alt lobulos 29. G. thalictroides 13. I > mm lat 3 papiloso . . en e l tubo y en los lobulos 3 . . . .16. G. liliifolia hojas con 3 pares de 5~61acinias 1. Galianthe ham. & Schltdl.) inconspicuos 24. G. peruviana jjpQ. , ig . Minag Qerms, |uL„l. .Hh/ a del :tr. II,,,.,,, "l".!'," irias, glabros o pi i",I; 5 J.n„a ^t;:: la c,' I aiuh :s: , 2 (_,»-,, t ,,, It I.,, t i I 2 ,m„ m.i, lIi, mIlJi.Ios „1 ! n, ,ol. ^b^ c. ♦ - Brasil, en el ' planalto central (Goias, Mato Grosso, Minas Gerais, DiscusiSn. En el protologo del basonimo E 2. Galiaiilli.- <:uiiii ( l ' -'.ll M .u... .,.,,.ss m^lm , < .Inn ^ ^ Kit II \= (.1, ,' i i , i, ' "' l r 1 » I )li.h r,, on™ .1 MO, PY). ' ' higado y via ne resina odoiifica L. Cabral, Bol. S. -4): 240. 1991 i ,, hi , \) ill » 1 ( 11 m [, II ' ii i >zi ( mini \t I nml > , i .,> I m n i >,) G-DC!).' ' I'.iM.ii. , (\iii..i~ • ..1 > lllt i. U.„,.l . L ,v ,11, |a Mai fo »i m' , spi f a^JlfjaiP, 111 i J' ... . '" " ,M, -9WJm. De Paiaguas si iono< . ln-i, ii I. GiiliiinllK- •■lio ( l:ili:in:i lSlan.ll.) E. L. Cabral, bre > fmctlflca en enero Y febrero - mo, 24690 (RB). Figura 4. I i /„„/, , gb- ( Lhoi!atmn < disco entero, papi la: corola 6- Cabral, B 27(3-4): 242. 1991 ) persistente. L.nl, , li u- , |HUl ,Bm CTES,MO). El unico 6. Galiaiillic clcgans E. L. Cabral, B, (Corrientes) 7: 10. 19' 'arana: herban0 B > Vila Velha, en campo sia, 15 colecclonad< ene. 1987, A. Krapovk 40875 Abruzzl 164 '' (holotipo, MBM!; isoti 31!). Nombres es. Se caracteriza por ser un sufrutice " balcuni " ( F jas Material re ei.j.tK 1- .L.ha- ui,n .stipnl,, « . H Umi-s «rr,-s : I„,m SI) BRAML Parana: N.Jlaui ...I. . .Jh IT") I (_>„• -, l,o|>.,M... \ I -b 1 > >u « Bum to 5<)lfi4 (HBRl Sai I" » * I, i .... P,(].( J ( u ,dl r ■•.-:.• Borreriafastigiata(G Bias \' ' 2 " ' ] 1401 (CTE (Martins) 6(6): 68. i date r*™?"'"\ { \. ,' ',; l' iTu 1 \ 1M , 'i,,,,,.,.' ! ■ / , to^Jackson" (Griseb.) Niederl. Boi lit ,_ , ' M \ \, , '"'/,""[' " , , 'jjW\ :^ 8 . li ' "w, a, , '.»-'( "- (K Vhimil Kunti foto F 879). Distribution, habitat yfenologia. Norte de Argen- (Rio Giande do Sul, S>!.1,i oriental (Alto Paiana. \ mai.il, r.U, n .71. 1 ... - leuo In . tl.iih 1 ...ill i MjUui,.!.. !- ,.>.mn<-. (-M'OOm; florece d* octubre a febrero, hurtiiica dr ,...,. o . Observations. El caracter sobresaliente de est. especie es su tallo simple de 0.25-1.25 m alt., con rsistentes). 7: 14. 199. linas Gerais: Rio hepaticas (Bacigalupo, 1974). Observations. Galianthe gramliflora se reconoce ^mb,es ndgaies En Biasil "sabugueninho do facilmente poi sei un sufmtice eiecto con tallo de .in ], .«.i ,l..h l.s . ., snl ,uim . nt. N l.s ca. 1.5 m .U-u,»t.li 525 y 1300 m, rupes no-pedregosos Distribution, habitat i fenologia. Paraguay (./ I, 1991]) y recien Paulo- lli.nau i, L">> / ,/ ,J, „ >, 7 i-Pi __' 1,"» ,,",'!> ! <)r 1W7 4 Schmm et ul U5IU i (CTES). 11. Galianthe gu: i Hassl.) E. L. 12. Galianll odat) E. L. Cabral, 186. 1904. TIPO: Paraguay. In campo Ipe hu, S: .!.■ M.ii.HMvLi. Dm , F tfa-/« :T(,V ifc.»l.iu|.o 3 0.5-1.5 m alt., t ml H h.s to-" i ..lo.ilinU', ■■■. & Hassler, P or se ■ bai os < -■: :ay, Sept. £ ffii»/ CT ™ <,«&*, per 13. GalianllM- k.-mp ni i ■ '., i !.n,i .,„ ■'..'. ! 57(2): 145-] iolivia. Santa y omb , e lu! al "\ loe iimha" (obsenaciones de etiquetas ■, . . r fructifica de marzo a junio. 15. Galianthe latistipula E. L. Cabra si 7i I -4 1: 18. 1993. TIPO: BrasiF. Rio Grande do Sul: Vila Oliva, prope Caixas, 8 feb. 1955, B. Rambo s.n. (holotipo, PACA 56661!; CTES!, SI!). le 80-100 cm alt., m indarias desarrollada, [ . ' ....,!,- \ P.. /<)!,), l'->')u>. -i : > , ,.,11.. -J,,.! J. l.Vl .1 ■■ulloru Marmuri 27227 (holotipo, CTES; isotipos, G!, MO!). 16. Galianth Stand! i ■ li 3.1 Sol ;nt. Bot. : ri.i-t. 21 > 92 11., Publ. Field n Mus., 1 ): 392. 1931. , 31 die. 1911, >2 Figura 7. «"i; in;: alt., 1- 6 2- Tollad.i,. Ull,. Mi.^m, ,..| ii ■ !0 6 : »0) . thalictroides (K. Scl '■ 1 ; e^r;* ■lus.i'tilh! : : :::!:::,: ■;"::;.: de 900-1370 m; floi ,ficados x 5 _-, <)!,sm.,.-iom ■ont/rc fifijfoKa se n os simples, puberulos, t 1 ,,,".! 1 '"'If' Soc. Argent. Bot. 27(3-4): 245. 1991 [1992]. Basonimo: Boneiia thalictroides \ar. longifolia Standi, Publ. Field Columbian Mus., Bot. Ser. 8(5): 394. 1931. TIPO: Brasil. Parana: Serriha, in campo, 840 m, 7 die. 1908, P. Dusen 7303 M . ,|. ,|-v_iJi-65 :nias de 5-15 mm! 5 , terminal, con pedu , !!„,, ,i, ervaciones. E: a Ga/iW^ gn,„5 V""" "\S '^ se " tatll ° '" s '" c/z " c/ « B R^ S "- ^ OIlis: 19. GaUanthe longisepala E. L. Cabral, Bonplan- 9 • US 2774977 )- PO: Brasil. Agua Ric; yfenologm. Centre de Brasil (holotipo, n< L-2-caules, de 0.8- a especie se reconoce facil- i m ak ^ con ho j as de 55 _ 90 x 1-26 mm, opuestas mente por las inn fas y por los sin braquibL labras, con vaina 3. / f»»"< ^'"'^ /' 'J-*'" (UBi PUUUU .,.il,i.n„.,as, aladas Es semeianl, Syn. PL 1: 124. 180 V,, , ■ ; < !/ ' , ^/,<< \R<.\ \Tl\V Misi- i>' om 23. Galianthe parvula E. L. desi nado a u (Corrientes) 7(1-4): 24. 195 Chl8nd ° d n L<,hAh,o. 8 hh. 1951, C J. Schwarz 11791 a variables. ., .. Goias: RR 040, 12 km al S de Luziania, ca. £• A. Furlan 3366 "' ' ' ' ' *" ' '' , '■' ' ' flu 1, tip > Hl'( R' jsitiuj i lf^'i corymbosa (Ruiz & Pav.) E. L. Cabral, Rol. Soc. do Sul), en ujeto a incendios nl.t .... onK nl s f ,1j ^// - , ■: f Al » BR 199 ' 1 . ' s^<>. ', r 7870 (MO). Huanuco: 5 p. 1948, 1 ; M.i.t... il | ■ 25. <.:ili !.<■ |.M-i>.f(.|>c>i,.i.ii.i i i Paraguay. Amambav Siena de Amamba), ene. F!, LIL!, NY!, P!). IP, Mai.. U, ,,--,. do ii 1 At... .'.i 1> K <, '*.. (Amambay, Caaguazii) y sur de Rrasil (Mato Gros inundables, en terre zii E. L. Cabral, Ronplandia y arroyos, de 200- : noviembre a (Corrient. 126. 2000. TIPO: rocosos de 600-1000 m; flore ular 1-1.5 mm, pubprula, con 1 lacinia central d de pelos desdt la l.i- d, 1 >- ! 1 nk - k.-t^ ' ' •"!«• d- sj„ u. I > a,(i i 1 ') deltubo). Suiruticp ramificado de 0.60-1.20 m alt, con florece y fmctifica de diciembre a abril. Matenal ,ep,ese„tatuo e.tudmdo _ BRAML. Santa Cata- pl 1.5-1.7 mm, g ulos de 0.7-1.7 mm, triangulares, > to. Flor breUstila: corola 5.5-6.2 is cortos que el tubo, 28 <,ifamllie-..nzi. I I i 1 - .1 M, ,o ,1. ,.., lllt „„,,„ m || ,iU U,iU si,!< J ,-3 2mm;anteras raiactenstira es poco hecuente f,i fl ^ ii^... I i i .,. .iljna .Mi-ir n. s ^ [,m„ilsi f wis, I \l\» M u IV, ?ps y 2-3 pare ccion clasica dp Brasil, Sao Paulo, A. W- Material repre.sentatu •o estudmd o. ARGENTINA. Cor Dist, dbucion ha bitat y fenologia. Nor, Jeste de K^o^TetlrL "ffidfLrCon^do u 1 ) T 1 al. 470 (CTES). BE \ - Cordillera) y Brasil (Distri 1957, G. Hatschbac: Girua, 18 abr. 1974, I Paulo: Capao Bonito, PARAGUAY. Ainamb, enero, fructifica de febrerc jemplar citado en el prot, 1821, voyage d'Augi - lakrianoides ha desaparec 30. Galianthe vale r,'o|!!,liK,.k, 1J en U elT! L. Cabral, Bol. 1991 [1992]. B MO!). Figura 11. En Brasil tiene utilidi Cachoeira, in palude, 12 Ian. 1902, G. 0. Malme 1060 et A ■■■ nm>, h j ,,. - pJ. ,...,< u^.v-s,, ,„„ 2-', / r^-WWriBBUBIP pares de „erv 10 s se, lacinias de 5-15 mm. Inflorescencia terminal, P. 4 mm, triangular-su i u>\, i;\ : . . .. ■ ios inconspicu, 2 (Corrientes) 10: 119-121. 2000. TIPO: Brasil. ! ■■ 5 NE of Diamanti: na, ca. 1.5 km frc cundarios' ' 900 m, 12 abr 3 (holotipo, UB!; . 1973, W. R. isotipo, NY!). ~1 Dutnbucwn, hdbi i afl( : m •elii E. L. Cabral, Bonplandia i verbenoides Colonia Ind, i 2111 (holotipo, CTES!; ' fimologia. Parage (Corrientes) 7: 4. 1993. TIPO: Argenl La Vina, Quebrada de las Conchas, entre la Salamanca y el Hongo, Ruta 68, Km 77-78, alt. 1300 m, 10 mar. 1990, L. Xovara & S. Bruno 9630 (holol ipos, B!, CORD!, """ m ' 11U1CCC y llucullca uc "»*" c """ c a J""»- CTES!). paraguariensis (Chodat) E. L. Cabral, por el habito, Observaciones. Se la reconoce por ser un sufrutice aceas con la haz puberula y enves pubescente, vaina v . con 2 sepalos, raro estipular pubescente y el disco nectarifero bipartido. 4 y sem iH as c lente aladas. Por el specie tiene tallos y idora, /luiz'fuf i fenologia. Nordeste de icd, Santa Cmz), Paiagua^ ( \lto Paiaguav, Mate, al ,euies, nt , . -. i < „ > > \U E V H , T \ * lit i BnlHu (MliAdTjla H„ u, „„ u , 1 «9 / -' ., ; lUL^i UloPuiuit Kuntze. Revis. (Jen. PI. 3(3): 123. 1898. TIPO: s < l " t - 1 J ' ' ^ ' NE <* J Hl '" ' <^ 1 " " _ "' '^ i-,i,j Minns (-■ ( li^r V' n^\, M ' B ™i P^J^7u'"t' b " l |] 1043 1 Uut <:(/ » t s , , i [^ (, (,B MO M) Boqueron: De auncion, 18 nov. 1992, i de 37. Galianti Schtdl.) E. L. Cabral, f^' Borreria hltdl., Linnaea 3(4): rt al 337. 1828. '•■.•.'.• a Sufnitice erecto, a \eces apo^ante de 0.5-1.5 m " alt.; tallos glabros, pubemlos, pubescentes, de ' vraii-ei (Mies-, ) E L Ubial Bol CI hda-, ehpticas u cnal ehptita- discol - > ., i v " ^ "^ -■;-■>• -j' wi [iw2j dg ; (kl ; dtPlluarlo , lMse dgLlfla ; n pseil d„ F ;,„]„. hasta la base de con suelos pedi m; florece de noviembre a febrero, fructifica de marzo a octubre. 1901)] es nom. illeg. MO, PY). 37a. (TjliaiilLf laNH ((nam A N |,| t «' '" •>'"' « ' « "HM subsp. laxa. 1-00 ! I .,h .1 P. I, ,„r„ (| 1 ,„„£, <■ ,, „, Observaciones. El material citado para la Argen- afecciones del higado (Bacigalupo, 1974). '. n , ,(,„/,. -IH-! -fllM Buc- • l,i ton I « 21 ,ul WUk R Palauo, 71 en relieve en : surcadas en la haz; \aina estipuh >erula o pubescent e ■ ■::.■■.■ •,. M.Mmn R J. mwl, 7SO-.7V 1 MihU bhos . , b, ,n, j ,1 -, 1 ,- , I, k i . :tesi, P!). ja.lo. de :;o- cortos. enlanntaddeltuboyalguno :,_!._>, r.,,.,,,1., 1.7-J »lo penristente. Ho'll'J,, para i ,u. & Schltdl. i L. B. Sm. & Downs, pe , trata eada una de ,h> I IIIU, SulLm- ]ls „„ L , „,,,. =f . ,,, ,„ ,„., , , V- C, viv, l .',,!„ l<;«0 ^ ' .„„„,'.,, 1= r , ; , ituuoiJe*) cm enti 39. Galianthe verbenoides (Cham. & Schltdl.) aqui. El ma: or esos autores e Griseb., Symb. Fl. Argent. 24: 157. 1879. identificado como B. leil ,i/.;.,»i'.> (i l],u, i\ Slili'll ) \l-<\M. , Mens. Mus. Pro. 306. 1890. Este concepto fue seguido por Delprete et al. Spermacoce verbenoides (Cham. & Sc tarina, Brasil, y 958. TIPO: Por considerar que el original de ' il„ , |i Ml „l. I ' h !,,( ' '' 1 . \, li,. I I . i 1 . ^ U Mnli.i' i HI ' .inilmcdo « f,iudo das Ha 1-34. ffiirtz, Paris. h.s (Chodat) E. L. Cab in li'i D 1 -( ur.rMn - * . , lift Ul > - i , .. L.lffi\S \ _'(«,' (2-1, i/F\EPi> U L 582(10). SPF 32772 (24). ' »l i in ,1 ' 'jili i,{ r r ,MRI" ' I « r, (35); 1114 (37a). 1115 (3). 1117 ,8). 1118 (39): C1HR (,*> i ' i -,21 , > , ,'.<> i W) UULIEITI, A. M. 13601 (1); !U< (J,dl HYISCHIULH (, Hi;.!') UP i ,7„, l. 1 ,!*' (37b), 45444 (35), 45529 (37b), 45672 (8 7638 (4); KRIEGER, L. G. s.n. (14); 401 I.WI'III \l. ' '_••;..: ,1,. M. s.n. (8 (3 I.UKVE^. E 24' ; MAGENTA F.'/M.p.H7dl. U2 34); MANTOVANI, W. 186 >J vh ((MIL 28602 (31), 34117 (26); I* VBELLC. M 1 MSJl'M > U. Mi 2^2 ( '-, . \l\\] \ / i JM i«2jc r , i'2.» - fi .i MrM>u «.m m i. - -, (10); KIRKRRIDE \1 . ' ]'■!( 1.6 107, !I T Pl,[], ,r,<> dil 6492 (15) 65?0 a I) 710 338 ! (3 .) ■ I2i E 8) 2-1 (37b), 14117 (35), 15 — rr« la I> W" V J6 )4-,U> uKMil MS5I (30), 10523 (25). PHYLOGENETIC PLACEMENT OF THE TRIBE RETINIPHYLLEAE Timoth y J - Motle r iA AMONG THE SUBFAMILY IXOROIDEAE (RUBIACEAE) 1 in. I.nl L (hiinuh i.Mf.Il, »bi,t.i« I ji!i,hJJ«li I ., rkuJ, , I tmtJ and isolated located at the base an involucel in the family, ason, its placement Iii . I is H ii il 11 b I Hi i ( ( \V LOI F I (,H UTiN K M MO ]\\ R U T \>Hx \ - \ L >U> M,v,i ( „il,ei ( .1. . r,,, , ,1,1. H f ,!„ ]'i 1 ll-jiKio]'>, _ — i (Kunth) DC. mcolor (Spruce ex Bent Persson et al. 1930 (GB) AF 201048 1 AF20] eira 10407 i - -MR. Table 1. Continued. GenBank; Taxa Voucher trnL-F C. F. Gaertn. Larsen 43134 (NY) AY555098 2 EU821636 l I,m'i|* nil. R tmi, I H. U ,whu' t K....I, . r. ( ,1 subiamih and ii.l Ia ik iu. .... -. t jut. !]... , ,. .,, .,j th tul . Retimplnlleae «as placed in the supertribe m , 'mit^.l . L 't : , .: . .. . ■ .. Ii i -M,iaii. ip complex In thes. anahses, Retun plnllum «as located in an isolated clade of the sister to a clade with m i b .s ti].i„l. L< Si.. . n,' I m tl -"1 I .mih 1 <. i h . .....'.,'■ :: '■- ■ ■ Henriquezieae (R lerieae (Del- al. (1997). The PCR conditions weie: hold 94°C prete et al., 2004), : & Cortes-B., for 3 min., 32 r 45 sec., 52°C for e 30 sec, 72°C for 1 min. 30 sec, and hold 74°C for (Andreasen & Bren ieae (Lantz et 7 min., hold 4 ation was controlled Pavetteae ( Vndic isui \ 1. < n 00! 11 ( | lt / , t „l ' u ,i bN \ t . i L« i> < i < i Is pel species was group (Persson, 20 -ae s.l. (Rob- extracted, am. ced, when possible, brecht, 1988). Amplified pre 1 with spin columns ■ ■ . : in prep.), most of t ,lved in three Valencia, Ca following protocols ■ .■- .' :■■■-. ' .;,>-, ,l;i (S ]JU ' -"U> T>N\ Giade F columns (\.m,i -.■■.:■:■• iflorum Bonpl. : ■..-. < gen W) In the second authoi, oi obtained fiom (cpDNA) sequences of the tmL-F intergenic spacer .Ulltl.. •/' "5J1IH- 11 til. [..Uj.l,H!lb . ,ts,,|,l l,f - ( 1mi,1„«1, 1 K m u ,R,,ik n.,,i P,j* -m 2(01.', L. are members of the si >.Me h ~i edited in Sequencher 3.1.2 (Gene Codes n, Ann Arbor, Michigan, U.S.A.) and pre aligned with ClustalX (Thompson et al. •■ They were then m ted using BioEdit (Hall, 1999). e performed with NONA i« lobofl L< >3 with WinClada (Nixon, gel, or from herbal! ,hich were parsimom infomial CTAB methodology (Motley et al., 2005). heuristic search resulted in 54 most par: ,.V(;<;m n<;TcccTCTvrccc-3') and -r (.v- 4,^ the strict -,»n-e„ , ATTTGAACTGGTGA ?S and JK support va using the primers rpsF (5'-GTGGTAGAAAG- ma Condaminea Hippotis Limnosipanea Sipanea Swaneopsis Mussaenda Pseudomussaenda R. concolor R. rhabdocalyx R. schomburgkii R. maguirei R. secundiflorum Scyphiphora Ixora Vangueria Fadogia Keetia Polysphaeria Pyrostria Feretia Cqffea Ibetralia Borojoa Botryarrhena Stachyarrhena Leptactina Pavetta Randia Tocoyena Calochone Macrosphyra Outgroup Tribe Retiniphylleae ;d iml-F and rpsl6 ai (BS = 89%, JK = A. Rich, ex DC, and Borojoa copln Hum DC, and 1 ested an affinih bet 56%, JK = 67%) ,ers. comm.), providii Retiniphllum species. All the species sampled of su] ment of Botryarrhem clade (BS = 100%, . ween Botryarrhena ar ■K.„.„iea. Lilt tut u, r il .,; I . lj i u llu!, ttuhses ,Ft g 1), the monoplnh of the J " ' * .'"•-'""- ^ »" >< ^, t ,i«. • -.. 1%6) AJlli., r data, placed Scyphi- addition, Robbrecht ; hypothesized, that these similarities were consistent with their . plast D . 87: 1731-1748. . L. K Mmth \ V II kl. m 2«* - ■ )loboff, P. A. 199 A GLOBAL ASSESSMENT OF a. DISTRIBUTION, DIVERSITY, D ENDEMISM, AND TAXONOMIC Jmtm M ° at ' 2 md Ned A Brummite EFFORT IN THE RUBIACEAE 1 ical Diversity, 20 ;l "a wideh enable broad-scale anahsis of distribution and I I S u» unti, .1 ,t<, s< ,ti .,il\i h J U . .' ,K> i„!m b-mlut, . i ■'■'.'.■ ....... impicne om piospect- 10 >< l,i. un. 1 •>> ( ( , w I «, iPkx.Ui t e( al , 10 >0 1>";«> : ....■., monocotyledons inclination is accessible onb ua list, ioi different parts of the woild (Banl.l, a ,1 A the Internet (e.g., Govaerts, 2ft but often without ».ih f .,i^.i, „ pe.i -,, horkli-n .i tli^ll. J,i<-. ^ t I. - «,, 1,1 fj, ,klj,t ol Faibiaccae (Gmaeits et dl . 2000; recently become available (Govaerts et al., 2006; < ?ae>) to analyze the and sMith, sis of 1 x ... , ,ir ,w ,k ' Mh , ', ,1 i m ,K ,n.l i< ll « >s ,-■ „t s, nk a j )lt ,,„ >f ,11 . 1.1 1 I I Untmlohl. .., H lx << i Mill i ill wil-VU. 1 mil bution, but species diversity and biomass are c ual computers. The Hl»Hi[ir m .| c„ ■ ! Ij\ ill -J .-< i. h*t ' ■••• ■'-■•'■■ : ' - !l ' ' !l ' T " '» -"'«" l I' 1 " "" U stems with a woody . * TDWG Level 3. forms (Robbrecht, 1 dson, 2007). sized areas Rubiaceae includ. Inch is by far ^^ ^ for bo(h ^ ^ ^^ for ^ eas) were extracted egaeta., . pplicable], othe: [.'ilni . < (' .. . .t M .1 1 H^Uniur^iM J, LOT Pi i.l„, !, . . .. . • ■■■■.. 1995). This \alue is then standardized to a size 1 ate at which species using the exponent value z. so (see below), more synonyms have An important co; value of the been created than retired from suiomnn, all relative to other regions). The \a!i i, lii'.n a , .Hi'.iluJ >1i -tin it m th i (JIM >-~u00 range >i."i,..W .K In kineial ( 20 -5 hi. h empm ih fc.,1 1 - till J')U,> . ) .It). ->„> ■■,.•■■. - F ecoregions. In this study, the intercept values of relative ° ;ources that exist in .■ . ■ ■■ •• : ...puVintd n, k ..sulhnch „„,!, .. I,u. -— We. 1C e ,e ,23,000-30,000 spp., 1535-1700 i , ,s ank, „. as M .,KM 1 ... » 1 ' l ' " ! ; ^ '^ 1 ' »'_ ' ' / ' > f " for all 369 TDWG Lei I •> ,,. . ., , „l ,1,1, 1> u„ th- ~" MJ "" ' ' !l J-'* 1 " 111 " >' l " , . calculated (Lewis etal, 2005). Parent!.. ■! ' : 0nreqUeS ' five largest eae, Poaceae, and \- .. ., .. h lb . s ,, p (I.. :;o Results and Discussion 1 1 < imii. li' < Cu!.;, v ii r 1989) and Smith (TV,:;i L-lwuf s t r. .,- I..11.,- , Luimd 'lopiral fo.ests and. iih.ii 06 genera. number of f le species is still a I 8 1 ?, I 1 ■, ■((..!.« I|iel.>r..li.,!inl..-!uil' species is estim (A. Davis, M. i .ns (K, P, and L) and an liti-.-Kir Ul.V >iL .ui,l L.t.d.1.1 ,\ <1> ir. ' TL- 2t .specie [uii ■ to . iota 6, lOl aveu- oJ (■>' j ^ues p, i * as So, tins ,»„ .<] Tli .-.e- (, L !,h,hed since 197f fact, it is very difficult to at the total number of \a 04, r year. The .la 1834 357 (Sohmer & Davis, 200 ; Ruhsam et 5 0p 2 317 313 260 and Davis et al. (200.1 ; the world's 8 Spermacoce L. 257 third largest genus, < ;uminosae) 9 Oldenlandia h. 249 228 (Orchidaceae) with c n, 2004). U Far " : 208 There are 30 Rubiaceae genera with over 100 ^ ^" e "' species, but most cont 182 162 iO species or 17 Guettarda L. 159 fewer (72% of gene raxonomic 18 Gardenia Ellis 143 133 1972, 1974; Cross age of mono- ^ Canthium Lam. 13 ° ■ ■ ■ ■ ■ . er such a large n lie genera in Lees, 2000; Ga very long-lived ' .-,. : ■;'• - 1 : : t is evi ical TDWG Level 3 a: , and usually fewer tha in subarctic reg entire Subarctic Am. y eight Galium L. spe vever, specific areas n tro . 1 belt thai . . noi ha ve high numbers of spe , 9( : 19.0041 1 : 17.5181 1 ,. 76 ' 23 Table 4. The i >0 highest areas f * gross „ Um be r of endenuc Rubraceae specs. Rank TD» W-.Uu.rK.liv,) No. of spec Endemic spe, endemicity ba; dies, New Caledonia ing is,!' M,^. > Hj.'di' iL- J , H \\\ , ..iv| M„li jv -i tJ) h .1 Loi-j [2»7\I1 h mb[M^ \| A .IF uil [»,i„l, i_'i I (,Ai Mtli 212 -pi.^^ eu.l micil len.i. -i> <]>. ( ,,»!,,,] \, 1 „,i.iK piil.li U"M \r,. witb. large islands or island groups, with the first five falling endemic ih , and Nigena 1 22 NL •■ , ■> id, *>(» ^.p ( r / into the island catego endemicity. . ■..:-• ■ TDWG Le\el 3 areas are islands, both small and Of the 13,1 le, there are 8456 endemics at T h means that 64% of enai ras, and, in the >J. M.uum et . ■i.. i-.«.:i 1,,. b..n du .r ,je lOjMOi. l. f i-,f,u.ltf t]..,| ti^'te^mil! 1 "!™!" 5S it will take us 45 years to fully enumerate dhersift in Rubiaceae. This calculation is supportable assumption*. ises of plified, as it does not take into account other l^:\Ti"co :s such as names added to or removed from (IUCN, 2001) have beer ed foi Ru! 5 be done and an indication of where to focus neasure of re) ,ity (Table 3, Fig. 2) analysis inch lion of areas that Based on estimat and Le species endemic at I Level 3, and t'ul ..,1 < In , \ li i, _>■ .- .-tdhl.-l. '« In it .<•..„ mis rtl tl, luf.itai-i U \ ,„ Pmpi.ii U .s i) [!,„ = nil ,m mi'lui .l.'U- ,..,! '1 ; ■■..,.■.■:■.■■■■. '. ■ typical laterally berr ; enus were not the points of i rs in two pairs (the De Candolle (18 queria in the Karsten (II genus Stannia H. tribe Gardenieae (as "Gardeniaceae") and, more Karst., based on „S. formosa H. Karst., which he iflom.P latfiluRo ... ,S. -< h, It ,' , I M , il, , u > . \,tit,„i} J Mill lutli'i li-lui ,omi . .'.x.'n l I ,.UI i ill m<- I L. otli . > \\ n ronsidtit 1 [hest the notes and material sent to him by Midler. characters triv . synonymize the two t «! ,> „ t m 1»x »i >th n..l u< l < n n Iki, I et ,1 _J)U , 5:» Ihe mthcis rescence and hermaphroditic flowers, rareh poh- ha\e a wide ightly curved from peculiar pollen c M the same first to notice the tnerall similarity of the pollen t.i,,.,. m-1 -,m ,., n.M.t .1 ,r rh. ., « otm il,. ,. i i„ ,1 »( V ,„ , „I, , , , I t I, . jun iJ',10 l.ri loU', l.'.'T) ...I Fl.u.l. ,n ,J ,Mi' -il ! .i.uil.1 tiif.„,lt ,1m,i.,iI ,»,>!,<. [„. tl is. It-it i-,o„l, Delprete Taxonomic History, Morphology, and Reproductive Biology of Posoquerieae of Posoqueua, and I was imalh able to defiiiil.-h aener;, Gardenia Ellis, Randia Houst. [ft» !( //a L.], ascertain that it was exactly the same as in fl yena Aubl, Gempa •■ . ■-•■'•■'' from German; Delprete et al., 2005a: 58). ubaea], Duroic some other ones." In Schumann (1891: 9-10) positioned both genera addition iw mjmr.iin-rl dial P- ::oqueria is excluded t..,.t n,.p,o. ..M.-nt- 1.. i.llm.n he .,, i nil„.,i- 1 .11 th< .ii lu.l. h uimiis ..'is «. .11 iik If. ud ..II .1.. PJn. c ml.m.i.n embedded. The. are not rareh dioecious, in which results, sep, -dated with Port- which the upper puts i ,t ,1 i , ( II n I , i u « Mil tl if. It ti ( t ! i ,M'l..-t Ltin -. .imk.- lespite these recent results, I chimarrhis Ducke and AWru Aubl. These results p ieae and Posoquer- (Rondeletieae s.l.). ■■■:■■ the tribe Gardeniea ;al chaiacteis s ho« some m ters, e.g., the half- parallelisms: apical < E Trib£ p 0S0QUERIEAE «idel\ dt limited Bondeietu a (i.k lii.n i < n ,„>t, , , t „ , , mr]l sl „ , r |, j,, > \\ lih th .iljlli.un and Condamineeae p.p.) as related to Chimarrhis a gene ral view of the morphological variation in the the close phylor between the Posoqueria is a genus of about 17 species Posoqueria and Molopanthera. In turn, they were d opics, from Central .... " '■ >>>>! 1.,-,, i .... >■ .Hphol^ioal, pah no- Pieties distinguished In the hpe of \esfiju,.- -I I he tribe Posoquerieae, as shrubs or pollen grains 3-c, ocular; fruits tall, with the -80) cm DBH, and Volume 96, Number 1 Delprete 2009 Taxonomic History, Morphology, and Reproductive Biology of Posoquerieae (Fig. 1A), which is tl tion in the STAMENS family. In most species of Posoqueua, the two doisal pollen catapu ly a few species of vamined for the lire observed dm ig in i< >elr.>~e m.- len similar to those of /W/ Posoqueria, the anthers are initialh united in an ellipsoidal sti ly oblique (Fig. 2A) nam'-fWied On the othei hand, those of Molo- a ture ' the tw0 dorsal hgulate, 01 lanceoL.tr >,n.l .... i. .i.liK , In. < ... ..hi] In both genera, the inflor pollinated In long to..- rd .plwid moths (Bava & lutijoha *as hi-i 1 ,.,,1 ,1 Jjlinm l <1 [ ,.! K Mull.. Beach, 1983; and pers. obs.). lower buds are curved, Puff et al. (1995). In addition, I ha\e personally Delprete Taxonomic History, Morphology, and Reproductive Biology of Posoquerieae . , , • ' , ' , . • -I' mt-.l, Hv> i»\e. downed, as the fltroeis aie pendulous; ho^ei, lepiesented by the solitary anther folding backward (in accoiding to peisonal obsenation, the. might also \ential position), probably due to cell shrinkage, which exposing the anther ellipsoid structure (Fig. 3B, C), or exserted (e.g., P. longiflora, pers. obs. in Ecuador, may remain closed, with the anther structme enclosed Sminame, and the Biazihan state of Tocantins), in Santa Catarina). In flow aid photographed in P. longiflora (Fig. 4A, B). structure exposed, as soon as the hawkmoth touches Molopantheui panuulata is a species becoming the anther structure akes a violent quite rare in nature, as the Brazilian Atlantic forest is forward movement (Fig. 3D, E), tlumung a dust (01 no« almost completely destroyed. I was able to find a minute clumps of g. e visitor (pers. health) population at the Feliciano Miguel Abdala In flowers with the Juded, it was "Caratinga Biological Station"), in Minas Gerais, observed that some hawkmoths forced the entrance of Brazil. Studie iology of this genus their proboscis at th id (pers. obs.), are planned for the near future. The pollen catapult sudden movement, a for the first time at this locality , and although the dust of pollen onto the hawkmoth. At the moment of floweis aie much smallei, the process is practically throwing the pollen, the two lateral pairs of anthers identical to that in Posoqueria. The five anthers are after, «ith the solitan stamen remaining erect above oblique position, as the flower bud is curved the corolla mouth. This stage has the obvious function (Fig. 2A). As in Posoqueria, each anther has basal Volume 96, Number 1 Delprete 2009 Taxonomic History, Morphology, and Reproductive Biology of Posoquerieae 1993) and Huysmans et al. (1999). tht poll n P j>u, ^er^ smulai. The. aie spheioidal (or oblate), ceptive aftei the antheis ha\e folded backwaid. mg the However, with the 111. 11...VH, „ t tl)l. (Fig. 2D). The fruits of re capsul; SEEDS The seeds of Posoqueria are attached to t] outline, with a central hvlum, and wind dispers< The testa is shallo re and the subd, elongated, «ith radial orientation, and interspaces ; ng within the Cortes-B., R., P. G. Delprete & T. J. Motley. 20 ,■ ■ . close relationship between the two genera. Delprete Taxonomic History, Morphology, and Reproductive Biology of Posoquerieae 82: 357-382. Hooker, J. D. 1873. Ordo LXXXIV. Rubiaceae. 1999 [2000] 86. A survey of the Gar FOSSIL RECORD OF THE RUBIACEAE /; , li.k, there are approximately 134 taxa described or f Oosperms used here l < ,n,l,i<„LvT. I ,<,uia ( V t -«J „...! W.I ' i]l,o !.. and none have ben i t ,i-.,l > w< die .„ i-in ,J phi. (oih., >i f „.,,., \,,l,j ,. „vl i'J> tU i-l-i.titi- : the literature of the not included. tion (e.g., mo -termination, better 10 1006.165 „.< ,„mpl.u ,|...ji.i,.i^ airi'w .u.^ui.i /, H, , ,U.ll Di- Ion G, mm., lUiM, .'■'■:: • ■ .. ■;■ reference is made g., the fossil town, South Africa; SM, Senckenberg Museum, Scholtz [1985] con an L., Nenax Company, Den Hague, The Netherlands; SUPA, ■• not found in or characteristic of the famih. These Netherlands; ULP, Universite Louis Pasteur, Stras- assessments are by the present author, unless C, Universite Pierre et Marie otherwise indicated. The abbre\ iations are also C Geological Survey, .'..., ....... ... , ■ = • . -. ' ■■ '"-■'-: author affiliation at cation, are as follows: ASNU, Aust ; of Fossil Rubiaceae berra, Australia; Cl> ial Recherche , um CRLUCLOLS of Natural History, I U.S.A.; FSTS- . MPUC, Museum of Paleontology, University of (1985: 72, figs. 17a-c). Pollen, southwest : !,.-,,.l.,, : ,v«! P„!omtt<.Wu P, ople"- < ivt tl , oous, to M Ma ihnrh Paleocene). but .......... ,. : Czech Republic; NTU, National T.,p,-i, Uui-.oimTx. ,1k - ij.,h tin u- pif.M.h (authors unknown, contact: G. Liu, pe 2006). PoUen, People's Republic of China. Both family by Liu (possibly Gardenia J. Ellis). NIGP. NA. attributed to Gardenia In Kmtzsch (1970; as species." SAMC. NA. -6. ^Trivolpotopollvm ■■■■::. ■ ;r, and Psilatricolpites coprosmoides Couper. Berry (1938: 133, pi. 52, figs. 4, 5) USNM. P. 'r..|.r«- U...I lr..il. M.-I.M •, 1994: 80-81, pi. 36, figs. 1-11). The •ry. Leaf, Berry (1938: 3, fig. 5), Argentina. USNM. P. antes camerounensis Sal.-Cheb. ialard-Cheboldaeff (1978: 246, pi. 6, g. 4), Late Eocene to Early Miocene, Cameroon. 'andia uliginosa DC, but this is regarded ■ .■ . ■ Wehr & Manchester, 1996: 25, pi. 2. fig. 6). NZO .,\ SfK DH Ohgo,- !e T^rx In-, -io> 1, n , Uln A FV .,, •tistej ham col !fe.< , Imeyer sp. PoUen, (1994) and Wehr and Manchester (1996). FLMNH. A. t^U.^olp,)'.-. d.-sf.il.-dln Sfl 76) no.,- tl.,1 ■ i p(»s,l!. ( ica. Scholtz (1985: i (Rubiaceae) and 8. Exoslrnia +p*,;,<1oc,,rib,u;,m Rnrv (1916: 349, pi. 106, fig. 3). Leaf, Middle Eocene, scribed, Berry (1916) thought the Wilcox Forma- tion was Eaih Eocene, but it is now considered i Benth. (Bomb; icaceae), amongst Middle Eocene. Similarity is cited by Berry with V", ■' tl.< vf.il /"•,/"* ,o,.'«.-.,M],Ml,.«] >" ,is jni'I.mifss e lFs\ |; (lr i l')LO cora- Schult. of the Caribbean region. USNM. P. the Caribbean region. USNM. P. (1985: 9-520, figs. 64, 65; Fig. 1C). MO. A. stedtensis Krutzsch (1970: 412, pi. 48, figs. 27- 32). PoUen, Late Eocene of Germany. ZGIB. A d). 11. Guettarda fei L916: 348, pi. (1938: 13 Argentina, P. Berry 106, figs. 1, 2). I SNM (all). P. MacPhail (1999: 205, pi. 5, fig. 7); pollen, Late da. ASNU. A. Roth & Dilcher (1979: 1203-1205, figs. 1-22; Fig. IE, F; Paleorubiaceophyllum sp., Dilcher & Lott, 2005: 17-18, fig. 9a, b). Leaf, Middle shown that the leaves are alternate (Manchester, pers. obs., 2006). FLMNH. A or P (because of 18. l\xthol,w ruai,d>folij!i.,u,, !<>;r> -zo 69); pollen, Late Eocene, Panama. MO. A (as family). 25. ■\Rubiaceocnpm,. muiLs-ioll Ki i, - I i l«M<) 108, pi. 1, figs. 19-24). Seed, Eocene, Egypt. SM. P (as family). (1938: 133-134, pi. 55, figs. 1, 2), Argentina. (as family). 134, pi. 49, fig. 19). Fruit, Middle Eocene, Tennessee, U.S.A., family but no generic af- pl. 45, figs. 9-11). Fruit, Middle Eocene, generic affinity mentioned by Berry (1930). USNM. P. , Fniil Mi.l 1. Eorru i.nu. -s I 30. +Rubi[>oli;s Mam-* >[' .rknpll 205, pi. 11, figs. 26, 27), Late Eocene to Early Pliocene, Murray Basin, Australia. MacPhail (1999) associated this with Canthium Lam. NZGS. A. 31. fTricolpore reticule, without atti Gruas-Cavagnetto (1977). Pollen, from the late Eocene and Oligocene, France. This was listed • (1981). UPMC. NA Mildenhall & Pocknall. Pollen, Argentina (Barreda, 1997: 286, pi. 1, figs. 10, 11). Barreda (1997) compares the fossil to Garde- nia (Old World tropics; see Miocene). MACN. NA. (1883: 130; Ettingshausen, 1888: 49). Leaf, Nevada, U.S.A. Ettingshausen (1883, 1888) compares the fossil to Cinchona (Andes). USNM (Lesquereux material). J\ C. ,Ynn<« 1960: 59, pi. 9, figs. 1-3). 980: 215), NZGS. A hall, 1980). I'.iJimi it J u/n]. 1 252, pi. 7, fig. 3). Pollen, Cameroon. Salard- extant Macrosphyra longistyla (DC.) Hiern that grows in Cameroon. UPMC. A (fide Midler, 1981). Pueito Rico Guettaula piesenth glows in New Caledonia and tropical America. USNM. P. 7. cf. ^Guettardidites Khan (see Eocene). M,.,;^»hr,; ! look. f. (see Tnpomletradites hoekeni) in Gevin et al. (1971: 386-393, text-figs. 1, 2; pi. 23, figs. 1-8). Wood, Oligocene and Miocene, in), Laboratorie de Paleobotanique (Lemoigne), 10. ^Pnl,u;nopn>snia,liH-s zvhuuliav I'orkn; Basin, \ustiaha MarMi„il IV**), « oh,|ui.- 1 Oregon, U.S.A. This is an extant genus in the Michx.; Fig. 1H) in the southeastern U.S.A. FLMNH. A. Salard-Cheboldaeff (1978) compares the fossil to Mitragyna inermis (Willd.) Kunt/e, which grows in Cameroon. UPMC. A (fide Muller, 1981). la Repository unknown (Vienna, Austria). P. 1i,.uL . n ..HIJ 1 i'ii i. M.H II 11 . - 1] l . HI p it .1 .in .K Ixi urn 1. Borreria G. Mey. PoUen, Medus (1975: 576, pi. 10, fig. 3, pi. 11, figs. 30-32), Senegal. FSTS-J. A. 6). Leaf, Colombia (also Pons, 1985: 241). Pons (1985) compaies the fossil with the extant rial). P (both). Leaf, Croatia. The genus occms tockn in the is. UVA. P. Mildenhall & Pockna (1999: 205, pi. 5. figs. 5. 6). Early to Middle Miocene. Murray Basin. Australia;' Canthiumi- tlites cf. bellus Middle Miocene-Early Pliocene. Falkland Islands. MacPhail and Cantrill (2006: 610, table 1; 613, table 3; pi. Ill, figs. 39, 40); Randia. ASNU. A. 5. Chiococca P. Browne. Leaf, Axeli nia, U.S.A. The Mint Canyon flora is found on a 212-213, fig. 40; Fig. Panama. MO. A (as the family). 7. j{,i,irlmm,lm • . ■.. - • ger (1850: 4c L, pi. 3, figs. 1, 2, i. UVA. P. 7, fig. 2; pi. 9, fig. 1). Leaf (as 17), leaf, Peru. USNM. P. O Mai J. .11 Klih.l- T'»<> i ><- Rubiaceae; see also ^Psilatricolpites copros- moides, Paleocene, pollen). 2.1. Gardenia pUraiahx l , aU II,. i. Calyx, Heer (1876: 58, pi. 12, fig. 11), S] ZOOZ. P. gelhardt (1895: 12, pi. 1, figs. 7 [as 17 in Engelhardt, 1895 text], 19, 20), Colombia; Berry (1929a: 91, listed only), Ecuador, under Rubia- ceae but genus belongs to the Lauraceae. RN-D material). P. 15. f >.»/(■».« .;,,„,(.,■■ .„„ 1. I\ L<-.|| \i ,,N (1939a: 132-133, pi. 18, figs. 4, 5), Cuba. USNM. P. i"\ I .: 1'^lLl. Pal........ I USNM. P. Faramea (see also -\Psilatriporites corstanjei). 125-126, pi. 21, figs. 5, 6; Tertiary), Dominican Republic. USNM. P. Berry (1923a: 9), Haiti. USNM. P. Berry (1923b: 26), Oaxaca, Mexico (as y). USNM. P. 27. Ixora cf. Ixora casei Hance. Poll-., (1969: 1174, pi. 310, figs. 13-15), Marshall A ... i. ..ii f'l'M' er, 1981). (1969: 1175, pi. 311, figs. 1, 2, 9, 10), Marshall Islands. USGS. A (fide Muller, 1981). . cf. Mussaenda frondosa L. PoUen, Leopold (1969: 1149, pi. 311, figs. 3, 4), Marshall Islands. USGS. A. . Palaeocoprosmadiles. PoUen, Zamaloa (2000), Middle Tertiary, Tierra del Fuego, Argentina. (1969: 1175, pi. 310, figs. 16, 17), Marshall Islands. USGS. A (fide Muller, 1981). (1936: 65-66, pi. 2, fig. 4, Colombia). RN-D (Engelhardt material). USNM (Berry material). P. Hoorn (1994a: 102, pi. 4, fig. 35; see also Hoorn, 1994b, c), Colombia, compared to ,i,!n<>» f » nT.-n::, P i no u - -u ,Ji M.u-K'HIsUhK |M,s ViM.illf. I")i,| but 4 lh i) 1, «,poJ,i i oiuj.ai.^ tli' ..J I, k> R ,' > > t [ Mifll . Wijmstra. Pollen, Hoorn (1994a: 105, pi. 6, fig. 63a, b), Colombia. Compared to Rubiaceae. UA. A. ■€ iwif>i,, r,< lii l.af k. .ib.i >t (1895: 40-41, pi. 5, fig. 6; pi. 8, fig. 6), RN-D. P. Lent' ( lull- P. B. li,...l only), Chile. RN-D (Engelhardt material). Marshall Islands. USGS. A (fide Muller, 1981). • , a- 2L-20l.Wa.-kJI kiwi, ! S<„i V ler, 1981). Pollen, Martin (1978: 191, figs. 7r, s), Australia, similar to Randia chartacea. UNSWK. A. 57. IjTriporotei Sal.-Cheb. Pol- len, Salard-Cheboldaeff (1978: 253, pi. 8, fig. 1), Cameroon. Salard-Cheboldaeff (1978) compares the fossil to Gardenia. UPMC. A (fide Muller, 1981). i.l, to Middle Miocene, Muna, Bj,ih, Australia. MacPhail (1999) compares the fossil to ASNU. A. : Pliocene, 16 fossi (1918: 43, 44, pi. 18, figs. 9-12). Panama. (as family). la Perkins. Fruit, Perkins Traverse [1955] as Rubiaceae?, and under Pollen Not Classified, ??Rubiaceae??) and Tiffney (1977 et seq.; fruits, seeds; see also Tiling & Traverse, 1994; Rubiaceae not listed). P (fruit as family). (1976: 813, figs. 196, 197, 200), Veracru: Mexico. MO. A. Khan (1976: 766, fig. 29), Papua New Guinea. . >j ,,ui K1.1 - !i -V l.kl (as Canthium type fide Muller, 1981). (1911: 335. , U.S.A. SUPA (possibly consolidated «ith collections at the (1976: 813, fig D), Veracruz, Dearie (I igs. 7, 8); Copros- Mexico. MO. A. muephyll, le. Leaf, Dearie f. Gafturo L. Pollen fig. 4b, pi. 36, fig gisches Landesamt, M< : 9 s. 9-11), Gen -lil^M 11. Mullen, 1981, "Me ik, (1976 ollen from th cene of Engelhar RN-D. P 891: ,1 :L (k-, i l..L\mmir an Republic Banks & Sol. ex Gaertn. Pollen, USNM. P. Crosbie (1979). (1900) compares the fossil to Psychotria. GSV. A. dt, Menzel (1913: 10, pi. 1, figs. 20-24), mlarioides Berry. Leaf, Germany. Dresden. P (as family). Berry, 1922: 42, 111-112, pi. 7, fig. 7), Bolivia. as family). (1880: 49; nom. nud. fide Andrews, 1970), Nev Mexico. MO. A (as Terebraria type; now Laugeria Rubiaceae. 1. Borreria ■ ncana Ferrera et al., 1-4. \Coprosm, Hum Deane. 1990-1991; Belize: Hansen, 1990; Panama: . ■: ''" ' ■ .... ":.....'. t al, 2003). 1997b; Co tra, 1984). Rubiaceae (Argentina: Heusser, 1995; Mancini, 1998; Prieto, 2000; Quattrocchio & Borromei, 1998; Brazil: ian der Hammen & AIjst,, 1994; Behling, 1997a; de Oliveira et al., 1999; Chile: Paduano et al, 2003; Heusser et al, 1999; ia: Hooghiemstra, 1984; Costa Rica: 1992; Islebe ).. mi l^i i . Jackia Wall, in Rox 'Hi \,.'i1m'wI, \\ (Washington, Oregon, U.S.A., Emmei Fj.1. LlMm.ll. Mi,.. F.Ui.i m iIipL.mmu u.1 ,i~ i u,i,i Lou, ,* c i I II L. .11 (~V Ma). cation and radiatioi ism is often associated with :.;.■■•'.,. • eocene (33.7-23.8 Ma; patterns in the different anther lengths (e.g., Rudgea 9.9 Ma), and Pli. les (Cham.) Mull. Arg.; scabrate in pin paied to the Eocene 1^ «-*-, 7 M„ 21 f M .1 m.l tl. fi lW , -, , • hi.n' ... tl.. n II .\m\-; Bakei, 19%) In i(..l..m J"»i 20< VI In Hi. } < ,ie, it «as a almost ill _ F talaman sigmfi.ant chanae fi in in P 1. « .. -£ en II, 1 uin -Mi.,11 1 ... k,l 1 • N , M« ■ ' ■■■.:■■■ Optimum (EECO), dunng «luch time tempeiatmes G„seb , Cuba, Webster 3837, MO), while occasionally also corresponds to the time when an in own by the diporate vegetation. the Eocene, t< rersity and distribu- te epoch (3 i M. lil.l u) Ihe Ml, Pll . em ill. I. '., both continental gla. d The history of the Rubiaceae is based on vit.it .,Ui.,m, iki tl. s „e L s Mh, .. „n „, ,„ ,i)s I m rh ,n«|,i ./ all kn.mn lepoits thf Mi Vli Mm 1 i l r Mi \ 1 11 th~i mth \m ... >,^ „, , 1 , _,, „ , n n h ,„ , ] d ,„ lltt lUII M [ l tl ,\, » jh.t n '.«.' T " ' 't'U . !. 11 id he H.inln, roup iM tl tl,. los il , . o d ! ul I, ill. , , t i V»ZZlnn^t fvlir. 11 ",".,.' ,V '"J, -; l(S ' t ; * u * i t i1 ,, ; ,i ( i , i '/, r \! ;, ! ,;';!,;;/ (eg IhclMnuhlnms Scbiui Tlnhnomim) Ho«n Wien-, H t\ J 1 1 I 1 ' 1 , w ,.- 11 il , D. I. 1940. The Min • ••■I u :-!-•-• 1.1 , II , si! hi i, „l ,t I. .ih m 1/ lr I I B..H 22 1-77. \t Olnma.P E. \ M Fiancj BaiKiciV K -u- p> . ]'>.'» btt..,, I \\ ,,i, k „1 - , I Dilcher, D. L. & T. A. Lett 2M , \ muldl, P , ■ Lo- Geyer, H. T lt.37 Ube. . J til,-,, i L 1 i ! , H i >> I '(., Vll . n. . 1. 1 1 1. il P . L-77 n Flora Fossil is Expen. Vetensk. \ibrtrn 4. i ..-.,.. 72: 504-534. Vol. 7. J. Wurster & Co., Zurich. i >V ium '!m ' >1 s. 47: 1-140. M , Late Cretaceous and Latorre, C, J. L. Betancourt, K. A. Rylander, J. Quade & 0. & \ M ( , J „ I > • > ... | i Til 73: 449-461. 95: 1-109. •ewald, J. T. Jr. & E. W. editors), Packrat Midder ■tud. Geol. 1: 1-117. Press, Tucson. I M . ., i£.n Jtt .,^ , II.. PHYLOGENY OF THE HERBACEOUS TRIBE SPERMACOCEAE (RUBIACEAE) J BASED ON PLASTID DNA DATA 1 , iij.-.m . ,.e s. st,., are characterized by species and genus levels (1 Is with an apparent adaxial has been the subject of debate. In the last a .lecular data show comprehensive classification based on mo y nested within the >3529-0266, U.S.A. I.,- ]'|-i|l,'2Kir..'t] ■■'"•■v -.'.■•. ■ ... .••■;... ■ "-.■. .■■:.- Knovieae that indue! 4, t > ' id it H ,l> hdei. - /;, - - , 1 1 ip.ik.mMioi,!, I l, f Nm - , :. . : ■ : -■; . . ip i.ui.l. nt t.-JSwii. nut anient, |. .1. hi- on, e as N tenum, L , the Upe spe< les of the In -l1 ,i n> mohf In -T K I ImuI.H ,, | , „ m Ii in . -utli ml i ill i <> ;< majn,], ' '■ : , . ■ • ,■. • ■ '. . ■. ■■ 1 • l^«i,fc'..« .ml i.'rtm.^',™ (mr hiding leca- ■'.''. ...... . -s s ,{- ,t,l ],„[!. „e l.iohi, VI.., t t, . i.^.ibl. .1 Mllun Speimacoceae si Thefii.t ,h. ,1 ,,,(,* 1 „n | iiIki.1„ ln.,i ■» |i,],Im„ milt,., i, IT, IhiU !•>- ct, I, ,/,//<„ T- ,11.1 ,- Mill- A !. iclataes Most of the li.ai,. Imnh pl.f il m H , h t „< 1 i- In. r , _"),< .mac oreae s 1 as .. ^ .■:■■ ■■■■ .:-. ■ Neotropical taxa by Terrell et al., 1986; Terrell, 1991, mo, , [,1a 1. 1 1( (J V, hat aie tlw relationships among •'■,-•:■' . ■- 2003). former tribes lanettieae? (3) What ceae s.l. is Spermaco is character- dia group? Speimacoreae and to enable identification of the prin :i] a] .■ il i the ihe S . h duded a tol ...1,1 M.i», ,. u'00<., i,.! kv bed u,l For rps/6 and trnL-trnF, we used 40 and seven newly generated (10 ices, 67 rpsl6 paisimomous t,m (Ml'ls* >.-,r i n .eat,-dh Johlld 1 AMPLIFICATION, AND SEQUENCING mateiial using the CTAB method ,.- .'. s, i,k ,1 I,. ,, p l„ „,. . > ,tl, 10 u„i...l t,. - h .hiiug fne trees per v.,-- spacei«a- done with <.h .mtl^tid s t ,u, anllj- J- , 1 UK; I,, hold 1000 ti -es, and calculating a consensus an annealing temperature of 52.5°C declining 0.5°C (Posada & Crandall. J9W1) mid ..111... Wl]Kl- 1>tl- iM,tit...n< , , ■ Dren, Geimam) avoiding to the manuiaoiun s M.K.nt- M4 Hi Ui.l,< 1 U Ru.icjuisl, 2001). Four 310 Genetic Analyi: usly for one million Belgium). Some PCR products were sequenced bv generations, sampling trees at every 100 generations. burn-in and c version 4b 10 (Swof- SEQUENCE ASSEMBLY ALIGNMENT AND GAP CODING ford ' 2002 ) Wf 3 50% ma J orit y mle h,! n,.K ...si,,, i ,„<", ,1.1m.- ' u< < •>' , .) J. H. Kirkbr. Spe Arcyl„,,l,yl(nm \\ ,lh! . s v. | Schult. f. R. Forst & G. For Diodella Small « Hed E Troj India, Sri Lanka, Thailand Polyura* Hook. f. Hed E Hiir Hed Mexico ( )b): < 3 > = Cabral (1991); < 4 > = Te r i U ,1 - .u i_i»H> Results Clade I in F BS = 77, PP = 1) includes a K, autia Verde, clade . /. ■-.. :■■■ :■ . ■■ ,■:■■■.■.; (Figs. 2-4). Compa es of the in- ( L .) R xb., a. Schum. Our results support. The Baye, , 95, pp = 1). resolved than the consensus of the MP anahsis. In clade II = 1) of Figure 2, but more resohed lineages rune low posterior a is species, except ff. Spermacoceae s.L. as delimited m rf„- .nlnidn.-lwn //,.,/, /.\ . .,,. clad.- iJS = 100, BS = 100, PP = 1), hst. (JS = 100, BS = 99, PP = 1) is unresolved. Within the Agathisanthemum clade, resolved as sister to the rest of the tribe (Fig. 2). The A to Lelya osteocarpa significant ,ackknnV ,11 1 IhmWi ip v.ppjit .,1 ! ltw l m , ,„, 1 \f 11( , m ,0/ < L / , , „ ,., . /,,,,,„ K S ' ., 1, this clade, stars wi s I to III are ican (0. um/Z, species (JS = 100, in the following paragraphs. (Figs. 2A, 3A) er relationship lacks pohtonr, is a clade with species of Kohautia subg. Within clade III (Figs. 3, 4), the earlier derived Pacfostigma Bremek. and Ohlenlamlia species, clades lack signifi -pecies 0. corymbosa L. (JS = •-.:■. stilus Biemek is sistei to a clade of \fiican ■ .. .■'•'.■'■ . / ; ... ; sister to O.fastigiaU oxieae. The main = 1). differences are listed in Table 3. with modetate lacklmie nicl hon'-fiij. -. p,...i' hi . , .dint- rh- i>i„n..j.lr\h of a nnmbei of geneia \ush limit im(0 M'l •' «T Ml II r M,. ll' (' n ,,',„ oi ,) >f,.l ].. . I/W, mJ fF Mi -II » If Lf! h U.-U. \- ,-i <.-,/-/,< s h|,,u,pmi tenelhfloia, and the Afncan species 0. lancifolia (Schiimach.) DC. (JS = 91, BS = 86, PP = 1). spermacoceae s str Infum, t - t..'x>,t]„, ii( hil. in r't .'.In if, ILl » a, borea, a species \m top folium serpyllaceum (Schltdl.) Terrell. iLMStJE^R © ■to 1 • o © ® ;«^f.^3 © ® ©. ^ ( Bremek. — B. Hed : ipermacocinus (K. Sch ,/./,/;, Martins. — H. OldenU Schltdl. —P. Olderda di Pit. the difference between the MP and BI analysis could supported, the relationships within Spermacoceae s. or characters. analyses corroborate ^"easUbs'peMesV^l'.ll".!^ 11 ,.l,"s|y !w !l 1-1, Wdio !,. N,th \u.. ,.. all W ,„* , than in South vttum may be one of ND M4MTK4 tHe few eXamf ^ tHat HaS reached the Andes by >n (Andersson et al., -^ ■■■'.■■ ■".. i pi i 1 , n phi , i 1 ,1 M.utlme^t Nee hi Wiofcs i H Lems In l.m 1« m. se M . . no I ; :,■■■. i ><>;. > . -"...<' hu n-i * „ \ ,. t, ,tl, . . m- i.h i i, .di - - - " - - U - U - --[ w 11= If 1-1 11 II 1 1 0//V,W,,ms Ten ell & \S H Lewis (Terrell, li lit 1.11 1 '/ !',,.'.,',,»<- ..■■■. ..,.,..■.. (Lewis, 1965; Terrell, 1991). craterifomi and pollen are 8-colporate «ith a their closest relathes among the Baja California ,-.-. , .-•..■ rell & Lewis, 1990 n Terrell et al, 1986). KADUA Lanka). Sevf considered the genus F . I i li f 'i i lit I o* - 1' 'i uililh mhn ti ! i r, , s .. I .i. ,T i II : : ,. •: ume,ols«l Onh section ffi^raunnra Me^n « L gem s , j ,,'„ ,H, V I t^, L m,, m 1939, including . ' ' : ■: . • . ■• • /,.'■■■•■ ■ . \ Inoadei samplm ecu l a r data of Pleiocraterium ... •. • ■ ., ■ ': ..... ■' ■' . (bireticulum). close relation ihocalyx and Phylo- ... . '•'. .• • • This subgenus, as descnbed In Biemekamp (1052), (2005a). Almost all taxa in the Hedyotis-Olderdandia , ■ ''. :. • .': 5« is distributed in Asia, Australia, and Ainc a u,l h ih m in ,., of, hut , PE\TA\npm OADE (sub)shrub, th: alyx with eight lobes, 1,1 ,1..., j>, •],- -I In Ruin, i.l le, ,., «, -, /, / an I > „' - t , ,,, JI ^ i ™, Conosto- se\eral morphological characters distinguish La- ,-> v ai.dV »/»» , Andersson and Ro\a (1999) and K--m el .il ,,.,!>, ,4 1 ,/.,«■, 1.1,-1*1 i-, Mmounded by eight .■■■■■■■. .:■:■-..: .... studies excluded rmacoceae s. clade (JS = 99, BS = 99, PP = 1) together with another and to the Pentanopsis clade (Dessein, 2003; osttr u > , ( n .. ih-.' (Ho- U . U that is larger than th era within the the same typ scence (loculicidal Hethotis-OhlenlanJia group (Biemekamp. 1952; followed by septicidal dehiscence), seeds with testa Scheltens, 1998). These characters, however, are c. tig radial walls, and 1 / „ i >m f ,i i i ai.l «nJ If l Ij i "d I in "/ .;'/»' Biemekdmp (1952) • ■ , . •.■:'■. .-.■• - //. Jfn and V-m.ntLUi.' I ,bo^ igatluwithenwm has a distinct ectocolpus, ai ndocingulum, and a ■ ' ■ • ..'•■;■-•■ MONOSPECIFIC CENERA WITHIN THE HEDYOTIS- ...'i 1 v.i-li I! , II- Wurtl, til' .1 ,»] ti.t - ^ n, II m< Mi , liun, n.dfu.i- «,lh jh.ms,,,, , I , .MlM.s-tb, r. ■-■.-: and Uiella aie ( h«uv u ^, A [,, i!^ piesem e of a , haiactei that ail,, \ i.hnuh ,1i.,n > 1 tli. -jh°> w , , ■■■■■■■■ ; / - ■ .• . ... .-., .. ,: . . ■,-.. : .,■•-.. . ':. ..•: Araofts DC. species, lera (Lewis, 1965). Xeanotis (Lewis, 1 t«een the two subgeneia aie .i°'.-il, iuo< t .Kvl.)l,e.. ,, -1..U.,- .< . , knmosome number. 0/7 /„ / j ,p u.,1, „ wU i ,,J T' I ,1, ,,?,,< l' i th I si]j,cnfii miy have diverged and sequence data of As; nth of one another fairly early .-].. iK <-...i a tlir //,.'. ,■,-(■,;- tli t. .l! v . -lit') ..11M1 , k ct!/uh(/zu group are Carterella, Dolichometra K. f> idely defined genus ^ ■ -:. .- ter ^einham, Pohuia Hook f, StephanoLOtcus molecular dat. rt the recognition of discussed in previou ; scriptions. position of most of t! enera remains : ■•'-■• IJium i „ Mm ti -"()!» O, t / I k» r, [ I , bin mu I i n i.itln I >/ t / > 7 t / polyphyletic. belongs to his group two. which is characterized In mek., and 0. wane, nuis, form a clade sister to the \,.. I ^l',^ 7 .// , ^1. . I. - I i < IKim. -imiIIcl , .fl i U ... ,„, , ,, [, d , , I.. Bi m Urn. d' "- lUtm " > U "-' ' li I ,l ', i, I to ,i In , , t. Tt is, whereas 0. herbacea in the Pentanopsis clade is sister to a J VNS AND CONCLUSIONS the rest of the clad. mcea is sister the tribe. Bes character sampling, enlamha, Halford ( d five groups seeds that die shghth lateialh compiessed and Eu.l 214 161-1.% .. 48: 1-197. , A. \ les Uitrasacrm 27: 159-166. superior c» New World hi. i... el 1,1 - < S ,.> !', .7 2VI-JH " Ffn' > 1 f 1 I ijsmai : . UilS-1 Mr.'. o ton, and! J -mi | l i >t .' rh L > ( Annals of the Missouri Botanical Garden | l| | 1 || | 7 V T^: 7 :: 7 7 7 '7 7' r^ :: 7' rrrrrr'r - . S F" :'- ;i : - FOLIAR AND PETIOLE ANATOMY I Teresa Terrazas, 3 OF TRIBE HAMELIEAE AND and Hel s a Ochoterena 3 OTHER RUBIACEAE 1 The taxonomic- usoluW- ,,.('.- . 2006). The tribe 1986; Kocsis et al., 2004). ixanthes Blume, Brignolia DC, Evosmia Kunth, that divides the fa, jmihes: Anti- Gaertn., Urop all, Sabicea AubL, doi: 10.341.- !00619i and tetiameious .ello« flo«eis «ith a usualh for each genus in raarcraa, Omiltemia Standi., and Pinamphsllon Bran- each genus. For comparative purposes, 13 species of ■ . ' ' ■ ■ :■■:■■- ,-■.:. & Pav. and ffi//», ided in this study incertae sedis). Ne\ertheless, in Robbrecht and in the state of Chiapas, Mexico, is only known from Pinaiophllon, and barium material. of Robbrecht (199 >) and rh is tl n j< II , < I i' , , , , ,„ f, ||.,, u ihWi n t I present study. addition to t ctions, leaves from -iwia , U-,11, d l»\ R<.!.|..r. lit ,1V0\) stimuli. ilj'k judixidraU sampled pel specie- aie studied [Ueppca, //,./•-■',.-, lh-'/m,;ima. Onulte- individuals vei. s„ m >d ( .e, ,, , ties) Leaves from ubfamily Cin- at 60°C for 1 h. and fixed in FAA for 24 h. All •. Terminology follows Metcalfe i])") ant' ND Methods Mentink and if, Wilkinson (1979) Deppea (11/27 spp.) refer to the central trace and lateral bundles. aJ,. l'V.i,.k s"t t .l,2«)()li U. II,. ll".„~,~ll„ ~. •. », M.l.l ll ,<., (1979). Results superficial. IV ,mata (Fig. II) with IJNDIMLM ipeci t ,.!.s llt ,„A mHamelw lion MES0PHYLL they have 10 to 11 c it All the material studied showed dorshentral De W .eo, simple multirellulai tnd.o. j. - ]. . l'ur L< , t ,u, ] , eis m the other genera (Table 1, Fig. 2B). The ■■■.■■:■■ the thickest walls (5-6 jim) (Table 1). arrangement is loose in all material studied from Hamelieae, 5) :hotria, and compact CUTICLE in the rema) es (Table 1). thicker cuticles (Table 1). Exostema an. , while druses and (Fig. 2F) and issooated with the ■..:■.. undulating antKhnal walls (Fig. 1G, H), vanmg Am \ p^rholn t , , . r.stals and dark- In transverse sectio lis are mostly In Hameliea mionly raised on the -, ss hicker and more \ariable in members of odier (Table 2). In Deppea, Exostema, Hintonia, and I.- ,i , i! ill ih . - ,li,.,|l. - -, ■ cm.1,11 ■> ..,-. i leaves and thick cuticle are the \asculai tissue ronf.M. ^ vi vi vi vi 4^4 w 4 1 55^32^ 2 24 4 * co iri« W "^4 4 4 4 u w 2 5 3 vi 4 c Rubioideae, Cinchonoideae, and koroideae, namely a ob,fue.< m Hi- p.< -< nl „lhK I map u«i, of trichomes, a singl. a confirm that ttpe I, described for Deppea of raphides and tannii rs from mem- es among \a- recognized in this ■.■•...- .-.-.. ■•■ ■ in any other member of the family. It is imp pes I, III, and VI. nal species of the genus and other I ction in the number members of tribe Ps. chotrieae to confirm the of traces is a n petiole vascular diagnostic potential of this characteristic. tissue. Notabh f the foliar trace was tlia group (Cinchonoideae) share the presence of Raphides have been considered a taxonomic labels of palisade ],„..,., |,\ l , . 1 1U ml h,!H]u'•} • n>^ ,t ,m.ml) . .ftl.f tub (i-i-li.i. ,e impoitant to stride then ontogen presence of raphi, eaves of this The single palisade parenchyma hner, the taxon adds support to the ide ; ular tissue in the midrib and Hi-) ir.pl id n. t ti di«, 1 l>< , a illtl ,l (t,, J II oi III) and the piesence of from Rubioideae. raphides and que combination of ence of druses are a i of characters ■.rn.h.d Fuilhe, s I, useful. in the petiole is luglifi lb .•! ili' i. ]> >n> d \, l\in ,11~ , '" if ^ ( , ^ ] l p u ' \, ^V'lVV™ ro'owus'lB. '") Bot. Neerl. 15: 1-33. & L. Ci >f the Dicotyledons, 1st a, Bel g- 1: i" 271 - L. Pro enga & M. Gragas an. E. S. 1999. Plant Micr ' »' / u diil i .. r i . J.,!l. HIM I Wilkinson,' H. P. 1979. , en, < 4H'h lML\lh > I t > , ,• .,,_/("t>]\l P„ 1, Maya 7773 (CHAP A), P. 7enorio & 7?. Torres >>, . /', / Jl. »'/iqL\[ , i> i /', ,i » .»"/. •,•••'. < .•«,'.,. if. 7W, ,',5/6 .MKXU). l ,>«™.. .7. /'«> (MEXU), M. G. Zola 0391 ■ x \ t, ', • D l , V ME 1 M k v 1', „ PARAPHYLY OF IXORA AND NEW TRIBAL DELIMITATION OF ]acc i ues Florence,' Joel Jeremie* and IXOREAE (RUBIACEAE): BtrgUta Bremer* INFERENCE FROM COMBINED CHLOROPLAST (RPS16, RBCL, AND TRNT-F) SEQUENCE DATA 1 ............. ■ Ixoreae initially consisted of two large Linnaean time that Ix, tot closely related doi It i ^'K'JUk.l"! nil) llil ', f II i 1111 I I I III JJ .lj sister to a strongly supported (JK = 100%, Andreasen (Robbrecht, 1988). Malesian Alei Yersteegia Valeton (Valeton, 1911). Accord] nera have contorted a Andreasen and Bren nces, and capsular (Halle, 1973), Dorice )00). More recently, Re 1989), horn, Mvoj, a; they tenta " '-*'»> ii • "I' : the '.,s ol Jwoideae s.l. (Andreasen & t ..w .,tl. -.i,H ^(.'u.^^^lftJIlSMft,,^,^! ,. three Southeast Asian genera, ileisanthia Ridl. oi agieement with its i ,..-,^ performed with zl/1 , III .1 -rem. lit 8 ■ bidre , Brem 1 rbcL, with at .: /■■ .{J c All sequenci; is of the performed with Big Dy or vl.l Cycle ator v3.1 Cycle d ly analyzed on a 3100 Genetic An; 1 pp] ed 1 ha-r X HlUl- , 1991) o r The rpsl6, trnT-F, i ind rbcL s. ^quences were al. (1984), as n I beta-test (Stat ,er 3.1.1 (Gene ;an, U.S.A.) and Doyle and Doyle (1987). Exti ' ■ ' '• ^ :. . . ..: ti. U^'Ahli l./) 6 , n.t .1,- lliw. ,U ,„„„.-, .,1- !,,!- !» U1M W'L.lr phase of 7 min. at 72 ,, one reaction primers d/e (Tarberlet et al, 1991) to produce and all sites ariable (GTR + G) ■•..■■■,:..•:■.. at least partial ov i). The rbcL imariant site constraint (GTR + I) for the trnL-F coding region was amplified in two parts. The first segment, and GTR + G + I for rbcL. Unambiguous CI = CI = CI -- RI = RI = 0. '•'•.■• RI = -■• 0.851 ' .. 1 10' M-lin,,.,!]- ' ( ,in,l<,I M,i.k«,wl,,jii . . -n i'( ..!<.. I dl. 2(0]; Darlu&Lecointre, consensus tree was ed trees from The results ducted in this study 0.95 were regarded as strongly supported. characteristics of the indhidual data sets (Fig. 1, p,.| K M^ i„r.,i,'.l I,...,, lln ,i,m 'JabU 2. n« < , i M \\ A MllLmri^ tin ,<>,L ,Njlw,.U lf - u.tli. -m-l. -|.i v nUli-- i. jml lolKiu, null hypothesis that our data sets were the I , . „ illiw. \ l.jl Ik ru ci » < n.l «/ , ' i epilations weie peifoimed. The lesults of the spoieae, C. , this clade was partition homogen d in the rbcL tree (not included). The relationship was collapsed in the rpsl6 tree (Fig. 1A), the similar larger monophyletic groups retained in while the Aleisant .roup (repre- the separate anahses (Fig. 1A, B) with strong as A and B (1 as highly supported COMBINED ANALYSES ( BS 100 %> ^ led the Ixora g r <>up, data, we merged the three matrices for all 34 terminal /„ drophy laeea C. F. Gaertn. (Fig. 2). The relation- samples (represented in at least two of the three , 3) and the Ixora- chloroplast DNA [cpDNA] markers) in a combined A iopsis clade (A1-A2) was data set of 3872 sites, including 288 (7.44%) strong (BS 100%, PP 1.00). In the combined parsimom informative characters. Of the informath e parsimom m a as lesoh ed with characters, 255 (88.54%) were nucleotide substitu- poor support as sister to a clade tions and 33 (11.45%) were indels. In our study, the c es Al, A2, and A3 (results not combined rpsl6-rbcL-trnT-F tree (Fig. 2) identified shown). Finally, clade B (Fig. 2) received high Aleisanthieae .1.1. hi...].]. Mini , mrl nnu.li J <„ i .i.ym. -i.,r.j . all «■'.! -"^ -.n. i Uai > of Ixora. SEQUENCE CHARACTERISTICS Our conclusions on the circumscriptions of Ixoreae se, 000), represented . i..i,», -J., .a,, S/, ,..,„ aH „ai .....ft a ,..* at...., . ...it. t.e , U li.t J, tl I [u.l.U-. . . Jy relat. e. :or. .on observed in Rubiac, a Ixoreae s. str. (e.g., subclade Al, Fig. 2) can easily Scyphiphora trnT-L (s rele^Js'hefeTlimi" 3lade A4, Fig. 2) hav sported position withi .2), i a remains both unrt a putative affinities within clade A tribes according to d The tribal concept of Ixoreae is better understood sarmrtioRi the name con reae to Gray (1858). sensu Robbrecht and ,ot corrobo- rated by our results, as it is placed in an unresolved x. I xoreae A. Gray., Proc. Amer. Acad. Arts 4: clade A (Fig. 2). I e F1 Brft India 1: 337 186l5 as su btribe from the taxa in clade A (F. ;enfam . 4: 1891; ag ].. -.,.(< m tv, ,u U V \'h I entire. Win,),.! ,> u, •„,>, , , / - ,n< («j,S, • 0.i,,..!h. .« i ■. Il -uppoited In unambiguous , .,(„„/.,. 77 ,„ .,■.,„ , \U Ixoreae (subclade Al, Fig. 2) as presenth , ,',')'' 11 V", t 'J. , ) "i: u r„ 1 '""!!' I',. '.:--!-,' ,\" I ll,'<.',".i 982; Tange, 23: 443-466. 1 >'. 1, is ulmti. ll u«.,,)l,,l.,^u .llv U (,, , don which it differs only splits and rolls \ndieasen and Diemei (2000) '■ . b .1, , ill. .! Ifa it, ] 12001. 1 .- 1 11 ( ,./<>, , . - ! „ t . . ■ 2001) \ serresol n.tnt pl.xjrn, 1 Iv.il, ,km j. ,. jup j\ f.n U Pi«l,n l'»:.7| by two of the autfa 002; Lantz & The circumi ia have always been ■ mi *usv \en.oun ami R.i.ison c jicvmw nphon (iN.uk of rlv . lost fa i. bt«U n. ia di.jecic.us group), «hich contained all sequenced ne\ ei been tested before using molecular phylogenies. Bremek. (Bremekan ,, identity of Pwstria and \iewed the Malagasy dmnu rsemlopeponu's > t It-moll - . \,e,v- iW fa i..u.j.litodite to Ca^aco (1972), „ nes, 1960), Pyrost, all other dioecious Malagasy ...... Trukia Kanehira Smith & Da Ixora pudica Baker 1 - 2 Friedmann (1994) irviflora Miq. 1 - 2 Burck (1883); Naiki & Kat< dium) (see Table 1). No species of the Indian < ), an Ivr.lnig system in Rubiaceae. Monoec), se| J04, 2005). iiu.i A Iurll.1- VK >.,L„ m ,1 P.,. M i,nl ) ! an! (I «ms „n ,e dioecious Vanguerie al flowers are moipl.ologicalh hentu|.liimht.. bur tinvti-.xjlb di,.- l ^ m ^MPLING cious; the functi reduce func- We trfed (o sequence as many representatives as rt"'bi"""l'l' lu-i'ii ..in,,, p) ^ il)]( n jj ll)imi]] ,|.- .ii.Ml' t i, i. in H.^ ' *" ' ""'^ n 7 !l 1 "'" l,u , *'" 1 ' .i.l til, ,- '(«.//,...,. 'A. ,.. t -.,,,1,1. i, ol.t.ii. K . linn «n, 11 M.e .i,.lm.i /J( ,„„ ,]| .„»,,.,!, ,. n . I- ..» r', st,,d...l.. .11 n...j* (Bridson, 1987). In \anuueneae, functional il. > o , 1 1( j m , , , d[ j/'j | (1 „ ', > , ' j ?'/ , , t .. (as Canthium odoratum (G. Forst.) Seem. [Skottsberg, Doyle (1987). \\ , .mi|.lih. d jikI -eq., nc nI put, .. L. tMBud-,1. 1<)Q, 2l> ,,! ) -i in, nm (i !,„,„ 11U11 , t | ,v r , -, ,, Us 1 r , «, V< ( , M ,T* i» 5 & Nlill.ll i Ciilz |B<. The ITS of the i ) the 5' border of 18S and ETS, and ETS-HL (5'- .ul.n. is . ,u,s, <,u«luii ,. |P i- on :<)(.«, j , „x R.^.fxx.H, I !.,'., m i >i Naucleeaesl [Ra/anm m hmbis. n ,'v IS, ui-i 2»»\ i_>,Km, [V . nhi- II -,, ">. n (ok lu.li„ t ih. S ,>-—,„ > .iM.-lla.lM i).20«U|. Mi-^iUei, (>t I , ■ un.-„m..lifi.r! 1 n,!,r, | u.n. .1 iimbison et al. (2004). Direct sequencing of their ;d PCR products consistently produced multiple al, t.«, !,..i!i nnikus, m Ik.. pu> ,ii- The resulting plrdogem «ill then be used to: (1) p ..rphism. As a result, ■ ' . ' ■■., /.■;.. ' ■ ' <-/■/"' '"'■ -'""I i" v .iiaueneae, (2> e\aluak tlu la,.-[,( i> riL... Sweden) and .ubsequenth analyzed with the 3100 (RI; Fams, 1989) weie calculated to e-tnnate Genetic Analyzer (Applied Biosystems). homoplasy. Ja et al, 1995) values DATA ANALYSES TREES on, iping, five random additions, an. to assess relathe «u!h IK p.afuhed.-.i:. ^K.n.n.hsis.-t-li^TISd.U ,NNJ,l,ia 1 Khs il ,. 1 , l , J n g It ik pj <-1-..].j]u v .,t ..l-tumm: 1^,1 K iWqir-i 20011 ||„ l..,tj.t n^M .,. o^lel.lT-ls'is'u'. 1 ,' erpreted as i et al, 1995). -'-.- ■-■-. l ZZesltl7^S7^iZ l T, tZfftt^zivz A total of 84 ITS sequences were analyzed and 58 l 69%) are newly published here. The ITS matrix nsensus (length [L] = 790, CI = 0.439, RI = ?27; result not shown) and Bayesian ITS trees had Hilar overall tree topologies, which were not in nflict with those of Lantz and Bremer (2004). In the yesian trees (burn-ins excluded) from the four iependent it el, all sequenced .1 .' ns uu m'm .1 M-ITN 1 j„ ,iii i . . H = > . 0* In'. -. Jihs,M,|ll = earned out Bayesian anahses of the ETS (PP = 1-00). The three h. ■:•,•■-.■.,>'■■■ ~. .si,, - - Iv utl. ih. Ml I I lit F - .07W,' urn sp indet (,, PP = 1 00), the . , P i huv hHh.m- l «, 11 .r. H.,i, ,TCR) bia.i.-h • la le 1 , ,,„ hull • atj.l. (1 /'. r „,a-Jnjn .... -e.j,,-,,,,- I,. ..II >„., -Iukh Co m, , .i ,1 . , nil ,,li 1 . . n ii. ilh inf hi,, t, ui.l. I . ■ , .lid ( .nn ■< -ill. n s l\hlaga^ species, PP combined El iced identical tree ..ik/edi-d-ik-i iupuLpi.-s. \ ,,;,isim<>m .malw-< ! le aligned ETS E ally MPTs (L = 822, ipport (PP = 1.00) as sister to the mo= .antz & B. Bremer), and the ipported (PP = 0.' s.L), all with high )<[ Bk «. iJHHl Wahm |l > .li« mi, mm .11 -i.| ><„i i Ik = < 1-J00 1 P - 1 M,) m ! ,' eep nodes received only poor (PP = 0.60) to c . JK = 55, PP = 0.64). The = 57) \ho-Malaga a / j w , ,> / -lib. 1 ., . iiionoplnletKgiouj. Fi.Uh Ix.ti, i- , w, j>i,i «eie deeph ne,ted Autlnn the /' , w., , ETS clonal seqi: irai sp. indet. 6 M\rfflI/MS.L. . u. >hl!",r <\il;,t. , i. , .,-ui i,t-t"i , -»,.'' ) fit* •■.'.'«'., '' , t . Cvl /•/. //,-, and iV/.- ( /ram clades all i. - . . -,1 tl,<- sine M 1, ,1 : : »i ' -,Kj-. 1- ic at. ! . „•< !>,> n,.s o.ohe.i and „[,. h,.,l,Ji.l.l au.up Tl,~ in. ] .Lei- of the /?«//oc^o clade have two- Brac « ^ e many other Vanguerieae genera. The combined ETS, ints to single Bremer, 2004) md the dttidu.is -iilII < , . i ■ ..,„.,, .l"7l.i) M. .,tr, IsL-nd iTia.ce) This fmthei suggests that Leroya and Neolaou J 1. nh' e, lt t, ,„ s.uo i, . i i »< > < i.lK.ll. I ].. 1«V> < * od'^ i' I ( i <«) . ,<■ imnl id ,! M,7 / / n"i - [ if-, torn I .»i M , t )l. 1ml . hand, we cannot yet rule out Hi . . in -. il, n, » ,.,o,hl !<, ,m - ■ ■■■■- ■ ;• ,-■ . . restricted to Vsian spim specie's LIMITS OF THE DIOECIOUS GROUP Bullockia clade. The Bullockia clade, retained in New circumscriptions of the dioecious group I i20P~ ,((■! inj, v t mrli.. J' ^>' full i- I'^lu.. .1 . Ii , ithei, ■* itli u>il, tl, c/arfe. In the ETS (Fig. 2) and !!.( < .rime in 7' ,< '«»■>." Vim)., i \i .i !').,( . iud h - \,l.|]. .n » <(». '?//•/ (Baker) Sumn 1994) have recently urn clade. The ETS and combined b e (Avino et al., ETS, (.phi-Hhu 'Ivi- „ill. t.u- ' D,J-i. l..,.t- that aie alua.s .... ....'• ,,:■:■■„ , V i-i „. ,i T IV (I.udson, 1987). and .1, Mat mt»)Hi>] 1 il.. Iilf idil „ \ ,,., n „, , „, ,,!,', « u , IITJ. , ,1 ,...,« vl.iL <2<>',1>, ,11 d i„.mI „",]„„ "j.,1. .Jeueju MiChvL,,," 1 ]^ U'™l Jj„li'.., n.'lvi. I l-V.T ,-.„», ,.',../,// ZV'/,'.J i', r ./ '.If i.l"ws'/.r the :'.:- li -lu. J lU I, 11, tt , . „d , u«..yu' !1| o,d ,n™ ML mkImIiik- mh ill M • . .) i* -.,,. >i>u- than Puostna with ii^rk-.l J. si In t Ko t< i im Ihnn.orlLum t . .. ,,ii l i..U ,.1 , >i< in, „U ,U .J. rn IikI, i„.. l< . 6.8-7.8 millio Sodrigues Island did PRELIMINARY B10GEOGRAPH1C HYPOTHESES OF THE not emerge ul llga ll & Chamakun, n a phyllum, and Peponidium s.l. (Fig. 3). 1 The prese i further disintegration Bridsons (1987) piedi. no,,, -.hi, I, .iu., 1,. I,.a< ha,! & ^ ,_„ f| ,, fw mm ^^ f> V>n„ t -..r„ 1 ."^mlt™,.! ,e . i.tn.. ' 1o d. »„„, t] ,„ 1 ,|i *r f> •.„ I , m, ., I, . li nanth Malagas, m ■ ' 1 i I [,-[«., i tin su.o.gl, -n| ],<•(, i P.utUI « .•).! tl. ih.. n' - ..]„■) u- MAi- m lift.. ' „,,!„ Bulloclda (Bridson BuLhckia Bridson, TYPE: Bullockia Lantz & B. Bremer. 42: 633. 1987. TYPE: Tanzania. Lindi Distr., "Noto-Plateau," ca. 450 m, 9 Mar. 1937, H. J. Bulloclda moml>az<-ii*i* | Baill.) Razafim., Lantz TYPE: Kenya. "Cote orientale d'Afrique: Zanzi- bar," 1847-1852, L. H. Boivin s.n. (holotype, P!). Bulloclda |>M-ii ( | ( >M-lilloi:i (Hrnl-oiil lia/alim., Lantz & B. Bremer, comb. run. Basiomm: Canthium psemlosetiflorum Bridson, Kew Bull. 42: 635. 1987. TYPE: Kenya. "Among granite rocks by watercourse," 3400 ft., 1952-1953, /. Bulloclda setiflora (Hiern) Razafim., Lantz & B. 3S Bremer, < ym: Canthium seti- ix species in mamla Ld florum Hi Oliver et al.] 3: 134. 1877. TYPE: Mozambique. "Between Tete and ,^t^"oi the sea < 1860, /. Kirk s.n. ishland, woodland. 1S II. Peponidi , Notul. Syst. (Paris) 16: 25. 19( ihium sect. Pepo- Diagnostic features. is distinct from tl 16 Peponidium horridum (Baill.) Arenes. le For a description, see Arenes (1960). ■ the corolla tube of Number of species. There are at least 45 species, ^fleeted hairs; core mbescent but n ot all (except tw an species [Arene, ^fteiie'r:; r^ttz 1 !:::'::: l')<)0; M.iiK. £007... m,l rK- S ..Icll s (h„ ],„„„, IV(H m >ir j. Llnn . Bot . 16: 14 _ 15 . 1877> , 1 ■ Annals of the Missouri Botanical Garden L m < , ) .. Mm cm. ,.- ]<> .. r . 1'H. |\-l f tuml „,^ii!, ,1 s| U m (ETSI ui l!!v>6-, iDNl ebola & E. R. Robinson. 1996. For a description, see Bridson (1987). n h f ■ tv, * v * «n L M - van Me< Number of spec™. ILu. .a .t L ..,1 . I- s,„ , e. sm _ Vi]llnPlinrt M , , .„„„,„ (55 species in Mad Dordrecht, species there, with in.l-oii. h M V). Seychelles, eastern ca, and South ■.',... ..'.... leae) in tropin i 7: 353-401. Buis-Pi ^ H • \\ [,nl.n H*'" Floia I oxalic ra* 167-180. iru.l < . I •»!.!« -. . Lull -Mir- Nitl &st Nat , T. J. Mo 2006. 38: 700-702. n uli i II J r J j ii , ii U ii I I - In- _' . .. i ill Mi \ .] His i , Dl -Mil Andreasen, K., B. G (editors), Flora of Ecuador. n G. Harling & L. Ander, P. 1 ! ' I I lk 1 to 1 ' \ Lid 1)N V isohtion x. Amer. Bull. ih>,>rt< efde- (,mvn< ' ' <, JU\ „:J I< ,vA " a Jdtc & y l^!r m p 1!;;::i:;;: fylander, J. A. A. Uppsala University. Uppsala. t i i« / pi -,w r >i _, j >-] b u _> i- i< 369-381. Volume 96, Number 1 Razafimandimbison et al. 179 2009 Functional Dioecy in the Tribe Vanguerieae H tt tt TT TT tt ri ri ri TT TT H Hn H 77 ri ri 77 77 ^;:Jc,^^j-« i § 33 3 : Annals of the Missouri Botanical Garden 77 77 77 77 77 77 77 77 7? 7? ir 77 ri ri Q Tj" 77 77 77 77 77 77 Q ^ ^ 77' 77' I I 77 77 77' 77 \ f §i f I % 1 S|||iii||||^l^s| i^d^cg-SJ^^j THE RONDELETIA COMPLEX Johan H. E. Rom,* Piero G Dek (RUBIACEAE): AN ATTEMPT TO Bir s itta Bremer ' USE ITS, RPS16, AND TRNL-F SEQUENCE DATA TO DELIMIT GUETTARDEAE, RONDELETIEAE, AND SECTIONS WITHIN RONDELETIA 1 mique et Bioinformatique de ['Architecture des Plaiile-, (\M\P) 1\-V51/P>2 Bhu , The tribe Rondeletieae (Rubiaceae, Cinchonoi- is mosth distributed in the New ^ o. ,.nie< «1tl,vi,ih ... ,l.— :i, ,M Vurill. - \ the tribe is found in Delprete (1999a). The largest genus of the tribe, Rondeletia L., is n Borhid. & Ja.a. K i .] 7?,. / Plane h ifoz^//u (Ste > ermark, 1967: Borhidi & Fernandez Zequeira, 1981a, b; Bo.h.di. l<),;2, I'W'J, l9 f <4.Bo I...I !«)•'> ,s f>„, ' ,/ , ,,,,,,1 , ,„ L>, ||, K t Una L999: !.ova il 2002 id e 1 ..< ,. t ,l R„ulim Kit (VI\i,n- r The cock tai | ,1 5 follows (to ca. 25 ul): 2.5 ul 10 X buf l_ _' til .INfP, |_T, t .J I ., L> for amplifiea:io. were realized k.1 . \in.wui.i Hi,,.,™.. Br. !....< h, m,l M .n ill. -,!.,*- ,on. ;ACE 1000 DNA amplification. For the ITS study, 50 new ITS seq. (2) gaps (i.e., inferred insertion/deletion events) were correifolia (Griseb.) Borhidi ,X M. FVuuuJ.v 2Vq. « 1 li. m ' pi s ,] V mpts \5 e were not able to , i I. HlblO i-v.U i 20-1(1 , M I)( ,-t.Kmt.l t , jl)i i... uU f .llcft i .», .trinl horn St \mcent each matrix. The i as a heuristic where the spe, d it was not possible HUMO .e|.h lrf s, .„,.! J.,- , s 11( „,J t .ml, - . Lei paper (Fernandez Zequeira, 1994): R. ■:■■■ .:■, ..■-.- " : :: ■ the trnL-F study of Rova et al. (2002) and from an Dwjer & M. V. Hayden and R. purdiei Hook, f it, s (i i) , ul , m i'i , .-, , In s .1 t >, - <>. ii 1 .i.i.nii i, uuk.I I in separate data set resulted in similar trees, which The first I i coding, included implied that the . iracters, of which 174 were parsimom using the command DELETE in the PAUP block. The w iters, of which 198 least ITS and ipsl6 sequences weie a\ailable. The strict ined from the ITS ping, and MULTREES option in effect). and 805 (CI 77) in the heuristic Results trees were id lata sets. Jackknife produce usable sequences weie mosth madr iiu„. K ,.rh rtjth indel codinj resulted in two clades not found in the jackkni search without indel ipport for thes clades was low in all cases. of Fernandez \ that is supported by ■ ■ ' ■„ k. <-, ,]«,,! .,' . ,.! r, II I,, m rn,.. 1 f.itUi 1,1 ..pjr nmmn,! ,1 « I i ] ,h.. i-l.i|. I. llif A Mpe^es. Neverthe eral interest- Our study does not sho« am support for a ''"' ■ I >< >iiii. Sin.) SuhkII. i- li position of this Central American species has recently f rom ot l, er actions In Laving 1- to 3-flowered been under debate. Lorence |1'X)<)| supported its m ||„ r , .„.,.,„,., ;lll( | i,.|i„i-.,-,,ilose flowers. this species si .f£ ^ ■fl^^ f| Chiococca alba Rogiera cordata Rogiera amoena Blepharidium guatemalense Guettarda scabra Rovaeanthus suffrutescens iryx leucophylla Cuatrecasasiodendron spectabile Rachicallis americana Mazaea phialanthoides Mazaea shaferi Rondeletia deamii Rondeletia alaternoides Rondeletia odorata Rondeletia pachyphylla Rondeletia intermixta Rondeletia inermis Rondeletia pilosa Rondeletia portoricensis Rondeletia miraflorensis Rondeletia apiculata unknown Calophyllae Odoratae Pedicellares Rondeletia Leoninae Leoninae Rondeletia Hypoleucae Hypoleucae to Hispa- (Rova et al, 2002). The fourth aim lobes, five to seven us revolutus Urb. ar i Urb. & Ekman (nc irasearh dhersifiedli na could be that we V ieletia s. str., pending future studies with bases shorter than the other Rondeletia sequences. 1 sampling. Another possil '.ana does not belong icrosMianthus ,ent anahsis differs d aim was to compaie a nucleai ITS maikedh fiom the lesults of the tmL-F study from >f the Rondeletieae with the results from a Rova et al. (2 mthus was found in nL-F chloroplast DNA (cpDNA) study a position equivalent to basal in clade C (Fig. f). A . ! '„/, „ v K11 W.- found tki I's 6 ;;; 11 ;;:^ included in Phyllomelia l„ne,<< l.„tl. l/ f .,i.„ .mi ['Inl'.melm aie easih distinguished In the peculiar fmit (pseudosamara, sm.su Delpiete, 19991.) .md . U i .Mil i 1 *.H ! j./ "/ ml tl.s , .is.. 0<" i- lit., it. t ph ].. i .1 . I ^ t ,s tl K , us . .n.l ill If. ! 1 ti . s si. , 1 <" but .gam theie is no iq s tint. ti.i.nw 'Ju / < i H ,. i ' - /< ■■■,■■■:. .:-.,:• Rova et al. (2002). es and retrorse-pilose flowers. This clade could s- t m L. .,.. , I.tln.lih, m.l. »,«/!< '/ / ... ™.i / Roi^itlu Ri laianthm aad Sub- -urm, w..sl..-U t ,r.,t| 1 ,.lt 1 ,.ul.l,t, f > r -1 Imlh.i ,m. i^il... I. , iiils], out, Ik l l].,tli. . ..libit, d T.al is„,i. i K i.tj.ii. I is !.,',.?. <>n 1 A' from ITS and . . is certainly needed. b. t« e. „ A'. lf /;. ,./,' s DC , Mu (It* 2, mi. li „r.i , I .. 1. . I ..i . , ,ilo < ,|,1- .il '.t.i] ii ]j II ... n.,.1 t . logical data) that HodgUnsonia is part of the tribe s imbricate, peltate. nW m 1 !. -, |P lj j t J , »>! 1 I) «) R 1 f. ii 1 ' Komi..!, , M>, H h' , i i, Tribe Rondeletieae (DC.) Miq., Flora Nederl. Indie ^"^he "description and delimitation of the Rondele tnin- H.-tKoii-l-Mt. R.m,i,-l,-i. Der Deutsche Botaniker 1: 77. 184] Rondeletia L. (1999) ai i !.« va \VY"« i..i- i i. -., upn, .. ,t il,. caducous; lobes often minute, sometimes foliose; a eeae belongs to the h ..nil-', . 1 it , „tl. ,11. .1 v'.lu-,. i t ,.,!,,. ..,!«. v , -,it .. ,1 ... .', -m ,1.' analysis (Delprete et al, 2004). The monohpic genus Rachicallis, endemic to jy Delprete, 1999a) to the present delimitation of Rondeletieae, which was placed close to this trib Bremer et al. (1995) and shown to belong Rondeletieae by Rova et al. (2002) and in the pre Ai.uluu.lhux 1'h-,, I,. Fl -iii.-J.ii.l Em 5 14-2. 1849. TYPE: Arachnothnx leucophlla (Kunth) tions. Barco, 5-8( 944 (fl.), /. Cua- trecasas 17165 (holotype, US!; isotype, VEN!). calyx lobes, capsul I seeds, ovary lad. El Km :: .,. I" I' 1 1 (ff) /. ill' ^ llt - ,h,), "I,,- .- ... ,-.1 'a s. ^. 3 duce^ in tb v Z7^i!;Tzrrtt^ etlard"; 5J; • I..1.WA 11 M...1.IN.I (1'll.H .list ingui- ,'.l ' .rsV i IL,' s | 1)t -.ik ! Jo i. ,'l| . esb. ■ , s, . ssed -] os. - i. 28 "amc u..ii< ..lie- LM.ns, ■ ■ to separate the two tax, . I..II into > ! ,i t!i- \ ,11. .1.1 r in a De ,rl , from a kn Rubiaceae. Opera Bot. U Vm Him. -VS r-Ls; PHYLOGENY OF TRICALYSIA j (RUBIACEAE) AND ITS ms > 2 M 4 lke F - Fa r" RELATIONSHIPS WITH ALLIED Enk SmeU,%S md Elmm Kohhrechf GENERA BASED ON PLASTID DNA DATA: RESURRECTION OF THE GENUS EMPOGONA 1 Tli c un, In A <, \ Rj, i, i ,« ,.! il„ I,,-.. I *jj-dt, ,1,-a.iil k.u ,ti.,i ( .jlKLje In the presence .i.numn'il lu . I, . V N „ , ,« s Mvlagascat (12 loberl cahtes. .„> 1 s^cl, vith a shallow hilum. of the tnhe Colfeea iBnd, ,n ,S \. .lou.l -'00., numbei of spenes arioss a hioad geo ..;,!.], ml Davis et al, 200" Ian inflores- ecologic range, often separated In minor and ,■,<,.! -m.- T\V >\L (T„ l(l lK, I, , tion within subgenus Tricalysia. /> ?w/ ' I,i I it! b lioisi , ,/'n' / n ,,',,] « 1 l lu .,U„l.. I, I.. lulM.lis M ,fU. , // // , ' > <• ,it. .in ,1. „', Species group Species Tricalysia se acidophyllo Robbr. sensu Rob T. junodu group T T »gafe™« Robbr. T orofi/bfe. Hiern Nok Em " """"■■" affiliation thin ■ eoncofor N. Halle T gosswdleri S. Moore Tricalysia se brecht, 1979 T crepimana group T H«^ta De Wild. sensu Rob T ruandenm group T auomfeui* Hiern rwuuferui. Bremek. Species group Species E. A. Bruce ..Schum.) Hutch, ft ,brecht, 1982 Tncahsu, sect. Tricahm, 40 s,, : , r. (^oieiau group ■a K. Schum. Robbrecht, 1987 - i .,.,. j , - . T„ c , ; > „ T oMensis Hiern T. pollens Hiern tzsch)Robb,9spp. 'ft Hook. L. 7 ' ri s^vdS£Ss^ye 1 . 1 r , 2^r Block -;oly X Baker De Block ft De Block Homolle ex IT f.A CG \u n;\ \(.r >'Ar\ t;v(. \ AATTTAGLTCTI \ " e pimieis 1661r (Jordan et al, 1996) using a Perkin Elmer (Wi I S A) GeneVmp 9700 Theini, I < ,rj* . , Amplification of tmL-F was carried out using the UENCING Amplification out as follows: 96°C for 2 min.; 34 or 1 min., 50°C for :s were obtained from silica gel 1 min., 72°< ; and a final extension lively, from seed, flower, or leaf f 72°C for 10 min. herbarium specimens (BR, K, For the tr 16 regions, 25 ul PCR leaf material collected from the mix (2.5 m Six; ABgene; Epsom, with tht romm. inlwin mis , 25 u] P( R mast, i pies, DN A was isolated using a mixes were >taq DNA polymerase ii Extraction Protocol (Royal (Bioline, Lo of 10X NH 4 reaction . '•.-■'.■' lerbarium material using the 2X dNTPs (Pro isconsin, U.S.A.). All x ! ,1 b lM.1 ^u l a--, 1 i- \ uii.t ,. ! h -hi , 1 K i 96°C, 5 sec. at 50°C, and 4 min. at 60°C. Cycle into five discrete Clean-Up System (Promega) using ™ ant. ih.H,' ■ ' tualh info, u,,^. ^ni . (\ M Ii,I.3j ,s, 11 , In d.lili.n -mini, i in 1 |. ,.' it l,i»s, n su K ». ', ,e cold) „ .ere run sim 1 2'y c ..1 nee- VjJ^-'hu,, ,m,L-. C^.fo. t..l. L'U'i, Ml . s.Mo^euatnebum-in. After confmnmg by eye that iKMll^l.hjlilh is analyzed with gaps ing data and deletions) coded ace mple indel lit analyses al. (2007). In ; luded 79 accD-psal ■■.':, - : ..,.-■ : d, I ,^r] han-foimation (DELTRAN) optimization, a com pleie sequence for rp/i6 in problematic taxa. •' '• . ' . •• ' ■ . .... fo.-.t t.aj, llld lw- fleWnstem. 1985) One thousand mgU)Up (Coffeeae! taxa and outgioup (othei lxoioi- pill II. 1 •,,>]!!, 1 'lliopil 1. plM ,[« ncl, p ll.li], .1 tlR ( Jt ,] , „ n \, ut ll,.|ll. pi ,, Vl< Mi. i * ('. ii, cf, V ,1/,-,/f Jl. s tl t.j. l.u.t (Posada & Crandall, 1998) under the Akaike a xamined by eye and '. . . .■•,■■,...- KY + I model analyses. TVL.li.lstu ,„,!,,]. the combined plastid data matrix generated 8853 parsimonious trees with a length of 929 ste] consistency index (CI) of 0.816, and a retention : (RI) of 0.908. Table 3 summarizes the tree stat th. .■ dh 111. D^.sm i.u,|i.i. x iH. ice (Fig. 1) were cc •t (BS) and branch 1 p (BS 99%, PP 1.00). Bt MP j 1.00) and is sister to the ingroup (BS 79%, PP 1.01 The clade of Cqffea and P.ilanthm is w. supported (BS 100%, PP 1.00) and is sister to t remaining ingroup taxa (BS 93%, PP 1.00). There also strong support for the clade of Argocoffeopsis ai Cahcosiphonia (BS 99%, PP 1.00). The sist relationship of Cahcosiphonia and irgocoffeopsis (BS 50%), but is supported by a PP of 0.98. Both MP and Bayesian analyses recovered a cla Tricahsia subgen. Empogona. Although there is bootstrap support for this clade (BS < 50%), the cla 0.98). Within this monoplnh of Sericanthe (BS 99%, PP 1.00), cospermum (BS 100%, PP 1.00), and the grou[ 90%, PP 1.00) and Tricalysia subgen. Empogona 98%, PP 1.00) is confirmed. Within Trical Bremek. (BS 85%, PP 1.00); and junodii (Schinz) Brenan, T. ovalifoh ophylla Robbr. (BS 98%, PP 1.00). The clade of Belonophora and Tr Tricalysia is present in both the MP clade (BS < 50%, 1.00), and a group of predominant] it,'j,l / .ly- urlith" inclusion of T jmmwifloi , .... Tricalysia (BS & Hook. f. ex Hiern (BS 97%, PP 1.1 d Wit] in Trica- moderate bootstrap (BS 75%) and I roi ps n c. . e strong for the clade of T. acocantheroide . ,,},! ,' (I. i i s, ' lt E>„Ilu ,i,l tli. ill' i i ,.1 1 21)07 Just . .lid not recover an aforementioned Ma. ether with T. Asian clade. a formed a well- Discussion strict consens yesian majority rule * • ' . I i ..• ..-,cir I a rlade containing Senamtlv [),«<«•••- 1 . i nl in Fi . tht iiisttmi., but theie «as s„ P| »oi> 1o, ll,e s. < molecular data. f. "t-uiii >.-,. , ou-ist, w is is otten the genus 7m. ','/ n ,>„ hi i H '.,„n I o, , I) , >,» i .jlnh.f i' . i '/ 'i . .in; /i I .!>,<< lit (1979) enumerated foil. - represents a ne«, though perhaps unsi ms Empogona are *• ,,,-'- i; ' ,i ,« , .ml in \ j. n I..! in. In.l Mm. I, ,h in a .i.iml,.. ,., ,iii ingDtp/o^oraandfts.os/OTn/um) The studw.t Da, i- (pl... .i,t.,>i< u jh.Uh, muphofo^, plastid legions (t,nL * <, O w mil ,i>f>. 1. >v, . h > p. i nntu .,i teims of taxonomic stability, as sUll.lt hill ll, ill s,,t 11 II llh 11 hl,.s th IS ,1 'l \ 1)1 . l)t. s , . , 111 V , i.lt, <,,.,1„ t.-u, ,Hh lW stnU In. I ii,.l Ir-.f . Ik ii,.„ m F!,.!„...i k-I, KIU, hi !<><>]. h, ,,|.lnn,. the, ,s , t...„- '"• -i—1" r..H,i.Mu ,19! • IX, K>.r tendeno to«aid unisexual lk t . j.i * .„,. - . ' " * t 1 I, , „ , ^tnuiis^.ui,,,,!!!,,,, i.-.v^.tu, .1 ™i„,h uU sfh In < ( '/,„r, 7, ,-,-/,. ., .onfined to Madagastai (Ranamelo-Ran- ( Table 1 »- ,, ^ lll '' u ' Jl1 ' I <—-»•«<" m fruits in Dncuspeimwi , fh. .Ldion [. iim.n-it, ,! -<-<-'■■»' Pi» «/ |K> '^"r. PP !<>(>, In I ,t Is, -up, t !! -ii moled. i Is ~e« 1 fphe-l-i.^he-a, here represented In unstable since its initial description In Hooker Je in the corolla throat and r . , ,11 t-.dl ■ ,ub |.e, l,cule on the innei surface canthuoak, „id T foirfsomaiiu aie situated vitlirn '.ti 1 A - ±ji tin p.sr ]\ n Ri.t n, ,fl\ • ,. nil «u i.np , • • Hi i,i s rf„ „ ^ , ' , 1 (2003) and later sup ith.) Hiern and the n.',i, Ip-j'i, "is , R.'li.cl.) I'l-'l ')i,t-li ,]. I,"t ,,^i>hi -it N .*,-,.,.. -"i.p,. Irk- V, IT liM The i, ..[..mi. .„,J, the dacle of T /»/,<«/», F <- „/'/<„'., 1.00). | | . 3 | b.echt j.,.( Mann. |3H i> ),.(.. 1 t-> pJir. L ■.„<„,. ,.. | ~ subtribe Bertierinae, sister to Coffeinae, as the g I g> | g| the .aracters in their • Mtilmi -I ],L«.l..nLi n.,.p)i«,|.>.„,.l a. i v ,,e _.,,-.. ....Ill, ,.'„!.>. .1 ..t /. -.',-< !s .2 jl.e| Inthethn alysis of Davis et al. ship between the v d . . 6 StnCt crepiniana (De Wild. & T. Durand) Robbr. , ,„,., _ the -,,, , i- nuiubeied .",, 9, 19, 26, and 27 in ;1 and below. EMPOGONA GLABRA SPECIES GROUP , ( r; .Mm h. I )?> -ij' , ' E !!,, h 19" > _i,\ r N „ ul] oil „ , „ i, spe(les (sperles uum l )ele d i, 7 21, and 27 below) analysis. molecular analysis (see Table 1). EMPOGONA DISCOLOR SPECIES GROUP The checklis lie novelties for all species, i.e., 34 ln f "' «■' ' * *'"'/<'U mi - COI!, EA!, HBG!, K! (Sim) Robbr., S. African J. Bot. 51: 331. 1985. TYPE: South Africa. E Pondoland, Egossa Forest, Aug. 1899, T. R. Sim 2386 (holotype, NU!). 4 ..l>ll.».)H-M.l. l Ml I I. cosperma Robbr., Bull. Jard. Bot. Natl. Belg. 49: 296. 1979. TYPE: [Democratic Republic of the Congo.] Congo beige. Musenge, 20 Dec. 1958, A. :•> ■ ' l!, uohpes, EA!, K!, MO!, WAG!). quaertii De Wild., PL Bequaert. 3: 157. 1925. TYPE: [Democratic Republic of the Congo.] Congo beige. [Kisangani] Stanley die, Tshopo ) -,. I l.' V l. WW(lU<1W'i BR!). 6. Fmposoiia l>raru~ita (Hi iu< j Tosh & Robbr., comb. nov. Basionym: Tricalysia bracteata Hiern, Fl. Trop. Afr. [Oliver et al] 3: 120. 1877. TYPE: [Guinea.] Senegambia. Karkandy, s.d., Heudelot 855 (holotype, K!). Bull. Jard. Bot. Natl. Belg. 51: 166. 1981. TYPE: Gabon. Moanda-Franceiille Km 23, 12 Sep. 1970, F. J. Breteler 6431 (holotype, WAG!; isotypes, BR!, P!). 8. Empogona buxifolia (Hiern) J. Tosh 8a. Empogona buxifolia (Hiern) J. Tosh 'i h <«, „i Hi ,n II li>p Mi [Ohver et al.] 3: 119. 1877. TYPE: Angola. Ambriz, Nov. 1872, /. Monteiro s.n. (holotype, K!; isotype, W!). .Natl l>< k 19- «!,. I.u.,,..-.,m, i.,i%n.,li.. ,i ... aii,lralis (Robbr.) Jard. Bot. Natl. Belg. 48: 465. 1978. TYPE: Jensis Hiern, Cat. Afr. PL (Hiern) 1(2): 467. 1898. TYPE: Angola. Rd. from Quipaca to fortress near Ferao, Oct. 1859, F. Welwitsch 3112 (lectotype, designated by Robbrecht [1979: (*) 10. Empog Halle) J. Tosh & ,oloi N Halle H Galw i I" .,» . l<*7(» T\1j- Gabon. Belinga, mine de fer, 21 July 1966, N. Halle & A. he Thomas 119 (holotype, P!; isotypes, K!, P!). 1953, R i, 4 Dec. 1937, /. Louis isotypes, B!, BR!, C!, MO!, P!, PRE!, UPS!, Hiern, Fl. Trop. Afr. [Oliver et al.] 3: 119. 1877, replacement for Kraussia coriacea Sond., non Randia coriacea Benth., Niger Fl. [W. J. Hooker] 387. 1849 [= Tricalysia coriacea (Benth.) Hiern]. TYPE: [South Africa. KwaZulu-Natah] Natal: Durban, s.d., W. Gueinzius 100 (holotype, W w,t p - KM' l' K' P . Empogona crepiniana (De Wild. & T. Durand) J. Tosh & Robbr., comb. nov. Basionym: Tricahsia crepiniana De Wild. & T. Durand, Ann. Mus. Congo Belg., Bot. ser. 3, 1: 120. 1901. TYPE: [Democratic Republic of the Congo.] Wangata, 17 Feb. 1896, A. Dewevre 740 (holotype, BR!; isotype, COI!). . Empogona deightonii (Brenan) J. Tosh & Robbr., comb. nov. Basionym: Tricalysia deight- onii Brenan, Kew Bull. 8: 112. 1953. TYPE: Sierra Leone. Jama, 10 Mar. 1948, F. C. Brighton 4723 (holotype, K!; isotype, P!). . Empogona discolor (Brenan) J. Tosh & Robbr., Kew Bull. 2: 72. 1947. TYPE: [Ghana.] Gold Coast. Mampong Scaip, Feb. 1933, C. Vigne „'„ stipulatum De Wild., PL Bequaert. 3: 211. 1925. Kew Bull. 8: 112. 1953. TYPE: [Democratic . i'. i filii'm-miMipnhila -ubsp. epipsila Wild.) Brenan \ar. epipsila Robbr., Bull. Jard. Bot. Natl. Belg. 48: 465. 1978. TYPE: [Demo- Mus. 3: 122. 1912. TYPE: [South Africa. KwaZulu-Natal:] Natal: Durban, W. Gueinzius 68 (lectohpe, designated In Robbrecht [1979: 313], W!; duplicates, P!, S!). ■ ■ phylla K. Schum., Bot. Jahrb. Syst. 28: 66. 1899. TYPE: Cameroon. Bipinde, Zenker 1569 (lecto- type, designated by Robbrecht [1979: 339], COP; duplicates, BM!, BR!, COP, E!, G!, GOET!, HBG!, L!, M!, MO!, P!, S!, W!, WAG!). 21. Empogona on & A. E. van Wyk) J. Tosh & Robbr., comb. nov. Basionym: Tricalysia maputensis Bridson & A. E. van Wyk, Fl. Zambes. 5(3): 475. 2003. TYPE: Mozambi- que. Matutuine, 8 Aug. 1957, L. A. G. Barbosa & F. L. de Lemos 7807 (holotype, LISC not K!, P!). K!, P!). 18a. Empogona kirldi Hook. f. subs Hooker's Icon. PL 11: 72, t. 1091. 1871. Tricahsia junotlii (Schinz) Brenan \ar. kirkii (Hook, f.) Robbr., Bull. Jard. Bot. Natl. Belg. 49: 271. 1979. TYPE: Malawi. Cape Maclear, Oct. 1861, /. Kirk s.n. (holotype, K!). Empogona allenii Stapf is the only species validly present taxon (Robbrecht, 1979: 272). (*) 18b. Eiii|>o«.>n.t kirkii - '■ i i Hi .- I,. ■ 67. 1900. Tricahsia junotlii (Schinz) Brenan, Kew Bull. 2: 60. 1947. TYPE: Mozambique. Baia de Laurenco Marques (Delagoa Bs\), s.d., H. holotype, Z!). N of Chilumba, 17 Dec. 1969, /. Pawek 3095 (holotype, K!). gueirae Robbr., Bull. Jard. Bot. Natl. Belg. 48: 466. 1978. TYPE: Angola. Musenge, 14 Oct. 1966, /. B. Teixeira 10701 (holotype, LISC!; (*) 24a. Empo liern) J. Tosh & lit // Hj j U Ti op. Afr. [Oliver et al.] 3: 119. 1877. TYPE: [Tanzania.] Zanzibar: s. loc, s.d. [ace. K Sep. 1868], /. Kirk s.n. kirkii Hook. f. var. glabrata Oliv., Trans. Linn. Soc, Bot., 2: 336. 1887. Tricalysia ovalifolia Hiern var. glabrata (Oliv.) Brenan, Kew Bull. 2: 58. 1947. TYPE: Kenya or Tanzania. 40-60 mi. from coast, [1884], H. H. Johnston s.n. [Kiliman- lolotype, K!). (*) 19. Empogoi a (Sond.) J. Tosh & ar. taylorii (S. Moore) 1925. Tricalysia ovalifolia Hiern var. taylorii (S. generic rank. The imestigation. fill required to fully Natl belg 18 46u 1 >7a \\Vl 1 , , qnue-, the „ b.sj W of Niapidou, 20 Jan. 1959, A. /. M. nbo ( /™«s Bremek., Bull. Jard. Bot. Etat Bmxelles 26: 253. 1956. TYPE: [Rwanda.] Mayaga, Mutema, 19 May 1954, L. Liben 1416 (holotype, BR!, WAG!). 27. Empogona bbr.) J. Tosh & liensis Robbr., Bull. Jard. Bot. Natl. Belg. 56: 149. 1986. TYPE: Somalia. 17 km W of Badade, 30 June 1983, /. B. Gillett, C. F. Hemming, R. M. Watson & H. Julin 25153 (holotype, K!). (*) 28. Empog iiIi.iiiii J. Tosh & botii Wernham, Cat. PL Oban 49. 1913. Bot. Etat Bruxelles 28: 291. 1958. TYPE: "- ,, _, "lb 1 1 ]» bM i^ot^pe K!). witschii K. Schum., bot. Jahrb. S^st. 23: 449. 1897. TYPE: Angola. Near Ponte do Felix Simoes, Apr. 1855, F. Welwitsch 3106 (holo- type, LISU!; duplicates, bM not seen, COI!, K!, within the ger ,ru.,1 /, „J.w/ avis, A. P.. M. Cheste r, 0. Maurin & M. tives of Coffea (Rub: oyle, J. J. & J. L. Doy le. 1987. A rapid DP »!..!'! : M2-K.1.. na 311-339. Associates. Sunderland. Massachusetts. ^s^^^^^^^^^i^^^^^^ Acknowledgment of Reviewers ■■ ■ ... • : ■ . • ,ik N -!i d www.mbgpress. org CONTENTS Third International Rubiaceae Conference: Introduction Petra De Block, Charlotte M. Taylor & Suzy llmsmans A Review of Molecular I'hylogeneiic Studies of Rubiaceae Birgitta Bremer Revision Smoptica de Galianthe Subgen. Galianthe (Rubiaceae: Spermacoceae), con una Secci6n Nueva Elsa L Cabral I'hvlogonclu Placement ol tin lulx Retmiplivllea. \mon«llu Subfamily Ixoroideae (Rubiaceae) Rocio Cortes-B., Piero G. Delprete & Timothy J. Motley A Global Assessment of Distribution, Diversity, Endemism, and Taxonomic Effort in Rubi- aceae Aaron P. Davit, Rafael Govaerts. Diane M. Bndson, Markus Ruhsam, Justin Moat & Neil 4. Brumnutt Taxonomic Histoi e Tribe Posoqueneae (Ru- biaceae, Ixoroideae) Piero G. Delprete Fossil Record of the Rubiaceae Alan Graham Phytogeny of the Herbaceous Tribe Spermacoceae (R11I 'lastid DN A Data Inge Groeninckx, Steven Dessein, Helga Ochoterena, Claes Persson, Timothy J. Motley, Jesper Kdrehed, Birgitta Bremer, Suzy Huysmans & Erik Smets Foliar and Petiole Anatomy of Tribe Hamelieae and Other Rubiaceae Dorismilda Martinez-Cabrera, Teresa Terrazas & Helga Ochoterena Paraphyly of Ixora and New Tribal Delimitation of Ixoreae (Rubiaceae): Inference from Combined Chloroplast (rpsl6, rbcL, trnT-F) Sequence Data Arnaud Mouly, Syhain G. Razafimandimbison, Jacques Florence, Joel Je'remie & Birgitta Bremer Evolutionary Trends, Major Lineages, and New Generic Limits in the Dioecious Group of the Tribe Vanguerieae (Rubiaceae): Insights into the Evolution of Functional Dioecy_ Sylvain G. Razafimandimbison, Henrik Lantz, Arnaud Mouly & Birgitta Bremer The Rondelelia Complex (Rubiaceae): An Attempt to Use ITS, rpsI6, and /raL-F Sequence Data to Delimit Guettardeae, Rondeletieae, and Sections Within Rondelelia Johan H. E. Rova, Piero G. Delprete & Birgitta Bremer Phylogeny of Tricalysia (Rubiaceae) and lis Relationships with Allied Genera Based on Plastid DIN A Data: Resurrection of the Genus Empogona James Tosh, Aaron P. Davis, Steven Dessein, Petra De Block., Suzy Huysmans, Mike F. Fay. Erik Smets & Elmar Robbrecht Acknowledgment of Reviewers drawn Annals of the Missouri Botanical Garden 2009 * Volume 96 Number 2 '.'■'•■'■'■■■ . ' ontribub ; outs (i . ' . • ililie< "I , Hinted in the back of the last issue of each volume and are also available online at www.mbgpress.org. Editorial Co C. Hollowell Editor, rtanical Garden Beth Parada ig Editor, Allison M. Brock e Editor, otanical Garden Tammy Charron \ssistant, Missouri Botani Cirri Moran Press Coordinator, Roy E. Cereau Latin Editor, oldblatt Missouri Botani IcPherson For subscription information contact Ann AT. oi- the Missoi m Boiamcal Garden, % Allen Mar keting & Management, P.O. Box 1897, Lawrence 8897. Subscription price lor 2009 i $175 per volume U.S., $185 Canada & Mexico $210 all other countries, four issues per volume TlIK A\\\I.S OK THE MlSSOIKI BnlWKU G\l!l>l\ 26-6493) is published quarterly by the Missouri Botanical Garden, 2315 'lower Grove Avenue, Si. Louis, MO 63110. Periodicals post- age paid at St. Louis, MO and additional mail- ing offices. Postmaster: Send address changes to Awu.s or tup Missoini Botwicat Garden, % Allen Marketing & Management. P.O. Box 1897, Lawrence, KS 66044-8897. stract/Clobal Health databases, ingenta, I !« OIL Research Alert®, and Sci Search®. www.bioone.org). © Missouri Botanical Garden Press 2009 The mission of the Missouri Botanical Garden is to dip ® This paper meets the requirements of ANSI/NISO Z39.48- 1 992 (Permanence of Paper). share knowledge about plants and Volume 96 Number 2 2009 Annals ^||k of the W® Missouri Botanical Garden BIOGEOGRAPHY AND PHYLOGENY OF CARDAMINE Reldar Ehen ? and Chnstlan Brochmann* (BRASSICACEAE) 1 politan genus with at least 160 to 200 Arctic, alpine, Al-Shehbaz, 1988), appioximateh 95 species aie and boreal species, and is one of the most species-rich Eurasian (ca. 48 of which are in China and ca. 25 in genera of the Brass: .975; Hewson, Europe includ; There are 43 species 1982; Al-Shehbaz, 1988; Webb et al., 1988; Al- in North and Central America (Al-Shehbaz, pers. varies considerably among different authors, illustrat- Arctic areas. There are fewer native species in the ing the notorious taxonomic complexity of this genus. Southern Hemisphere: 20 in South America (Al- The center of diversity is clearly situated in Emasia; Shehbaz, pels, comiii.), 10 in Austiaha and New indein NO-0318 Oslo, rfrica. gave rise t Barbarea dherged , such 6.0 Ma. This be an underestimate With., by Heads (5 ed the dating of the pollen. Howeve data set of Koch et s,116 al. (2000), I ? (2001) performed a :tions. more thorou ious evolutionary rate deca. assumptions. i divergence time of Lterest 6.2 Ma. ,. (Schulz's Eu- five number is x = 7 (Elven el nnl> known with In = 16, a number is In = 32x = 256 l^n.nmp.,1, ■,l. ill „> •jin-t.i.-.ul..: |. - , 1 1,.' i tmL). The on - ' 1 1 i Nh, J- (100-», 1W,> ,„ ,n .-id ?,,!,-/• s i 1 .. Li e ;ne> ; ; PCR ampl f ITS was pe ' ; primers ITS-4 1 30 cycles of 45 sec. al r 45 sec. at 55°C, and < primers c and 5 (White et al. 94°C (first c . ]'*M l-lll ? .l.-t r.,Jp Mat Fresh leaf material was sampled iLdd.j d .n.iiic, >•• , «|5 < rfl , I '«) , „ 7: ( ihsl,,J gel in the field. Vouchers are deposited in the 10 min.). P( led with ExoSAP- ■...•■ ..,■:.■ specimens in ALA. 0, HBG, LE, 10 sec. at 96° i& 240 sec. at 60°C. 0, 0SBU, 0SC, S, UPS, and WU (Appendix 1). Sequence- mcher 4.1.4 (Gene «hei. (uK/Wizne oc i , W 10 ,* *'ie H actions positions were coded according to the International described by Schu I: Union of Pure and Applied Chemistry (IUPAC) 0. E. Schulz (Table 1). MARNA (Sir i, 2005) to detect Kit or DNeasy Plant 96 Kit (Qiagen, Hilden, st led correctly. The V. Giraldiella 0. E. Schulz" Yllurn 0. E. Schulz" damme Godr. ,!,, .Kl.'ls. ,.{ aligned ITS mi *hich 188 were p ;';;_ ;;;• luding the outgro excluded hom the , >l itk.t,,* Tui tl>. ,/,/ i pi. i ig New Technc .logy (TNT) (Gol id teti in Figure 2. ITS trees into n: -UHiillh ,,)■-< Ill (2ra = 16) Eui 20u<» Ilemsf], ^.jhI,..- rt ,. 1( (f s Jsf < 111 ih> HJutvM Us 1J(>) I ,.,, ' /M 'f //»< \\llll <" MM toll, ( . ■:■-: ^ :: • - longer, starting wil rees one step Group B wa = 1 and comprised one step longer in 11 steps. Jackknife (Farris et al, o jrrhiza 0. E. Schulz analyses were perfc 1 = 5, and PP = 1.0. The European species of section taxa, and diploid ar; ita 0. E. Schulz, C. bulbifera set in MiBa^es (Hi Is n r' cV h mqmst 2001 Ui. N and( - , |V L r b n*h iV Ronquist & Huel- h the model Hook. f. ex Sc th the Asian diploid the default settings tarting trees BR = 4, and PP = 1.0. Maiko\ chains foi ion. The first the East Asi; amine macropfolla 25% of the tiees «eie discaided as "buin-in' torum 0. E. Schulz and the steinii Dyer of s dition to two DC, C. lacustris (Garn.-Jones & P. N. Johnson) = 98%, BS = 97%, BR = 5, and PP = 1.0) i nth the inclusion of Schulz). The North American clade (JK = 100%, BS The ITS data - | \ - in< .,i < lusne about the .■.,M ) i..i.olil:.n •, reed C. Ursula of section Cardamine. purpu7-e«, C. jWufa, Group F wai d by BR = 1 and PP = 1.0. ,,, C. victoris, and C. sphenophlla). Ho, gi ever, most of the MPTs s from Sou! lerica, East Asia, and Africa. -.„.. i.sas.1. oup with North Americj irou : ;maining trees support! This group in separate groups, but bo tZZZ^Z esTbt^ref^S h ... .e- ,,..11- of C. scutata npling analyses. The South Ame The South An , i. ,., . o , , I nth giouped with was most closely relati African and J f .,l,K 11 i group J, C. bonarient Pers. and C. flacc o or nested within the d C. ecuadoremis Hiero to In = 56. 'ere resolved as a sist Group H wa ulcdbyj] 65%, . & Thomson in group BR = 4, anc lit t] , >. „ , . , „ s did not form a mon I, ,l.d . ., „, up Mill tl„ olh . < ,, u iirnn, C >.],.! * due II.U ex A Rich was different taxon. A n from New = 2 and PP = 0.96. A specimen from South America (L.) Crantz). The Oceania two distinct clades. Group I was supported by BR = 1 and PP = 0.94. I %, BR = 4, and PP . ... .. , ' . ■■■' : A extensheh studiec et al., 1998; In group J, 1 Australia and New . . .. ,. Id n n r.Lt .ui hi u i , 1 t l I.MMil,: ' miFI' = ( ><. , l,« \, ti ( umhdlata and complex from our fi C. glacialis. species related to C. pratensis to simplift this The Berin occurred in three .-:,.■•...•.■■•. "■' ■■ ■ ..^ .,... : i America. - C. tenera 2n - 16 [Card] - C. matthioli 2n = 16 [Card] - C. uliginosa 2n = 16 [Card] 1{ fo c. acris 2n = 16^32 [Card] tis [Card] g n° excluding the outgroup). There were 11 potential of the plasti, suited in a poorly with CI = 0.694, RI = 0.807, and RC = 0.56( uted a monophyletic them, as well as the strict consensus tree, is presented group with 91 BS support, and a in Figure 3. In terms of initial similarity retained as BR support of 2 with C. glauca Spreng. as sister (JK = synapomorphy (R[ rs were more 67%, BS = 55%, and BR = 1). There was also self-congruent than the ITS characters. The analysis support (BR = 1) for monophyly of the Oceanian < « i — in r Q H S inr Canlamuiella spe< i. - « ,,.-,»(/>,, Hi- < -I.- American C. cordifolia A. Gray, and the South Although . n the ITS data set : ■■•.■' ..■ • ■ .•: di*^, »t ..1m, to-uu.l ,. d.H.- ,JK = 12 = ,A I BR = 1). effect of hor, s. The most likely rapidly. affected the resolution of the ITS phvlogenv. Our ( i i,m s<\\ . it, tU C , , „ t ae maene.al independent], i., , o, Inu •, ji f—C. cordifolia "ft ' ^'olalls " -C. appen'diculata -C. Jbrewerii -C. flagellifera -C. flexuosa -C. lihengiana ^tlz! ^E; \ microphylla '.. purpurea :. blaisdellii :. digitata :. sphenophylla i.'li.lls .in Mti-im,,,,,,,, „t r l„ most spec les-nrh c lll.ll F 1 -J, ,.(-! 11.., 1 * Hi, Ih 1 pattern observe d ii ol lei irge j .... ra ■ i! is family. lyses of the ITS ca. 350 species; Koch & Al-Shehbaz, 2002). l.tt . lit). 1 L . lll.tl , t ,,l ,, ,,, ,|[ ,. plrdogenetic signal from frequent alloj'ol , tion. Mranant speciation c ... 1, u! .1 out became the pi-Mit « .iHtellation of continents ua- o ert-.it (fi-Mem t [ M . -ZWi) 7 i..u\ jiiJws to. Il,i- .huh ,ni,h 1m mm late Tertiary forest in the Cu.rdu.mine ma; prove fruitless. ")' , pio i,' .1 I!, lust htbitat foi estab- 1 ' "li. >K< ,1,1 -,,U 1.1 ,t ((,'./.»- /, ii 1 possible that the Intel submerging of the Bering Lnnd s on M. t], , distinct clade sed Northern Hemi- UtTmoNiNG sphere taxa (group H; Figs 1, 2). Thus, Oceania appears to ha. I least twice from the Schulz's (1903, 1936) large sections (Figs. 1-3). results aKoie,ect the monopbh oi section Da I u " '" ' ^ dle moiio^letic ">* »« tilt °f ; ;.,t\ Lit "' ,;" "\n i". !l F f- -- ;-" » « » *- ^ ^-» 4*1 In S.eene, and P„. (2000, ^ ' '" ' - "« ' " " "» " U - "™ «*«*™. - 006 > \\. !■■■> .il-« Jim,,. i)i t V„ ,./,,///„. .u.i '.*;»< - V, .ioir.) 'Imr.il SiH U Australia, Tas Zealand in both the = :.:^ DF{ = J.ri' = o.')a)tr.v,.„„[iun-.i J , .. RAPID COLONIZATION AND SUBSEQLILN 1 DISPERSALS ^ ^ ^ subc j a( j e ; n gr()up j^ Because of its po< .ylogeny was The South American Ca , ,'■■..- to reconstruct an. the European C. Despite its typiralh shoit-distan. e mam mode of diiection is impossible to determine as both ways are Hemisphere (Asi , America). one old dispersal e\ent to South America with a ,1',,..-.- J,[. Ih. -, i N,m. d- r,u t ilT - U 7/ KH = I, Up,, i ti, l,/ in '. / t /,< •> ,- in Figure 4. group F and d from Eurasia (not Oceania. One example of ,en long-distance "' Zealand taxa, whi ;=an species did not form a ,■,■:,:■ Ln..-, rf ,i .iium ,au . . in Fi„ 1, 111. Uu.au (' Thus theie aie two distinct examples of European ouu., ,^|,t 1',. MM- ,',«,t si,!e, t , til,- -,[!«..! iln , n example of a probable Noitli linen. . , mi u.ucana. «lnchoccuis both in bur . I. Ji '.s,n .>ijoi, mto i„ -n -,,„ , cS m Benngia, but Literature Cited BS = 88%, and BR = 2 in tmLF) to European • A. Fr» diploids, specifically in the Alps and Pyrenees i nn " " , l T . b in Fig. 4). This ™e species ( rassica- 0.96 and BR = 1). a, \,'.V \ZnuZ '«','""*,.' i.'.i T i ' s!;,!' ..' 'f- a ; ,,h ;;'' "!£;;", ' h 'f 1 " V /,' Y'^ ■ ]1V 1", ^' ' l ,sT !' ,| "dl u ' f "n, ! ,"4 K F ' 1! ' "" ^ ''l' 1 " R [ ■ling analyses. •> - r ,n Z»> t li< | i ,1m,, .1U1 ,(.1 i . H ri LII„,11VP , \\,| ,_')).(( r , , [, U, li \ 1 ,1 V 1- -nil - m, i lf -, IV. vl m vs T » A \ P (> I iiu.l-i.i,,^,. I P .S T IS ,,-ij ,-t ..ctl UKL^I - M, „] ,,,1,,™ JBm«po»i « 9-312. - C.D.l.es M M,tM in , ' Bl, 1 , M I A! ^ H, I 4(. 1" ■2-1043. Bot. Jahrb. Syst. 32: 280-623. a,J.,I. \ dUeVk l\h i U J>«i >n Liuf J t ' i J' > < i i il i P -'-,, ' .IiliiF ebeit. S. & R. Backofen. 2005. MARNA: Multiple I '^'i*'k - f - 3 IS I I I I 1 S i^ Si I 3 I 2 | ! ! § | | || ||g|| ||3 3 lilll 33** 55 ^s^ia ^ 3 * , ^555 1 ip iiii l 1 11 11 223 23 ?S ddd s d ■* a d ■* , i, - <*!,<-!. a, ( k„ng (1992) and (1913) and Skvortzov and Wang (1955) described S. •■..-.. •. . ■■ V, , bescens Tuicz., subspecies patula tl> >hU, M < -'an- lia> - ir» 1, ! « i .J [.hi., mtloi. - n. and l< i •-. China (Chang, 1992) Swmf , u -rul.Ml s ; -, .' /« .i /Wens subsp. mzu< ellow anthers. (X. L. Chen) M. C. Chang, Taxonomist w taxa based on a and potaninii (C. K. Schneid.) P. S. Green & M. C. limited numb, caused quite a few Chang. Seringa mneri consists of two \arieties, . Schneider (1905, : ' V] SSI Bot. Gar potaninii, and S.giral, as new from populations. niiiiM. ,lm i.ii. - Fo- .Minol..T, i .u,l'>U>tk- . ill., i []), jumcipal component anahsj^ : objective of the present study is to exan th and width of leaf revise the 5. pubescent complex. between anth. at contribute mostly nd Methods A total of 14 populations were sampled across stly to the first two China (Table 1). In addition, a large number of , used by previous Peninsula and Inn ming, Hebei, Beijing, Shanxi, Sha. nan, Hubei, SEE 0F LEAF BLADES China were HIB, HNWP, IFP, K, NWFC, P, PE, SDFS, SDNU, our observatio ied continuously in larger in popu and LIA (4.4-8.2 X s,i u„ , , , , , p ,t I ,i,t. in ...I k t.. Ft \ xi i u~ ,u hn. . !,. s «^ihp considerably ensure ca. 10 indhiduals in each group. The among populations (Fig. 1C). s Because the indumentum is a very variable which have b densely hispidulous forms (Table 5 face of the leaves is kOK <,JN "111 and 1H hi, li i ■■■:■■ ..■■■. i i ::, ■ . 03180 i i /. Y. Chen anxi Prov.: Mt. Hengsha^ us /. Y. Chen 03208, PE anxi Prov.: Taiyuan, Mt. forest /. Y Chen 04102, PE verge, 1300 m andong Prov.: Mt. Taishan. sunns o,< „ -In. / 01207 01210. lY.Ch 93158,0316( PCA. The po IM, and HUA have The leaf venation from glabrous to meyeri and S. meyeri var. spontanea (Chang & Chen, HEN the\ ai- u >.'h "hl-ious „,!,, , >[> lm. ,n " Chang, 199S) Haung rutualh the lemammg p, t>< laU ., - lahle 2) Similaih, the (Pig 2U Smulaih theie aie hoth palmat, m • ■ La, .< I. i J.oi,, . aiil.i —• tih..mllieJ.,,t fu, < om.lm, i - li ...lis KOI!. D \N, and L1A form a loose a .(3 :-. ; r a third, h die I ill. i / « i ig M KOR DAN LIA JIN WUL BAI HE IUA QIN ZHU HUO TIA KON MEY Population 0.25 0.20 0.15 0.10 0.05 l DAN LIA JIN WUL BAI HEN SHM TAI OTH HUA QIN ZHU HUO TIA KON MEY Population "fll 2 fifi SH™Ki:::ffi F~5 KOR DAN LIA JIN WUL BAI HEN SHM TAI OTH HUA QIN ZHU HUO TIA KON MEY Population » (CTS), expressed as [(diai 5^ii— iffit^i— tot ]DI0K0D04X<]^0[>0( # D ' ♦ ► o o X O <3 □n OP- ♦♦ *i ♦ 1700 1800 1900 2000 210< ►Shape of infloresce) rachises ■Indumentum H * a ing (CTS greater than 0.12), and throats usualh 1.6- ind The second lations WUL, BAI, - M,n.lHn\ «!,[,'. I ^ i ind rachises 4-.n Zed and glabrous and leaves generally Hebei Prov.: 1831, P. J. Kirilov s.n. (holotype, hi p. . ere mem Leaves 1.2-10 X 0.7- :apsules 6-20 X 2-6 n drs; petioles 0.3-1.5 cm. Laishui, K. M. Lion 2 PE _ 7 "" /'/ ' ,' " L»]t J M I lull" A X 1 "nil hvu I Mi 1 (holotype, IFP!). ' phylla Die Leaves 3-6 X 1.7-4.2 cm, io shan, Lao yu he, microphylla [. L. Chen) M. C. t 1 ». 1 ['c ,1 ., M 1 < 1 » 1 ' - 1 1 Pl .stributed in Nei -ophylla var. gfaAn HU i\ . . .1- 1. 1 V> 1 ii, 111 .la v Discussion. Sp-ing ;scribed by 7 Schneider (1913) a: ated to differ more or less palmat. base. But our liaJ-.tlr-.l' •ix.'JuiMI - iwfev,,,. M |.,|) [k ,Ll ".J I !'," I'd'. V„c," »„„; /„ ta-s^ differentiate the ^v/o^xTwe propose thaTs BulL Bot Res " Harbin 9(3): 4L 1989 ' TYPE: C1 oZogy LE!, photo PE!). i. : idb ; . I i.l , V ( ( :. 10: 34. 199 u HIT . In 26 May 1911, E. m C. K. Schnek 1., PI. Wilson. (; Tpe, K!). Mt. Ta ui\r Tlu.-, j ,1 in Mt Jmil.iv i>HMi d.i t is not reliable to establish th« rf the leaf blades and in th« 1/1 11 1 Re p> m corolla throat. 21100 (PE). H t, I two pec chose G. Giraldi 1644 B. Z. Guo 1408 (PE); I (1905) described „S. cliehiana a- new anc specimens, G. Girald I and 742 (with f.iiln u-h i f* m , V -. I.rliltl U « ii ,ph i ,],„!,„ | | lm > > , , y (Pb) H«>,im^i / specimen as the lectotype here. . „, ,, ,i.snl i S s | 1I1M ,1 t n,„,'J >c . (PE) Liujiiu, i / / > (V- '" '' !' r '"' " i' »» "U- >l'.m . ; j (h „ H , n2n7 (lt20) ( p E) S/!mAn ,„„ (ill 1.1 < I . .1 ')(), I tj.h'i 1.1 111 ' ^ -JiMHllMU Ml» , A f , / (OTH) are clo-e to those in S pubescent subsp J j. ;; to. u i -i J i J r -i *■ " '<• '" Hi < i » is S ]., -I i dl '. 1jii.it s ( IE vi . S If 11 li , ] / H / ' i ' \iili *" _•/>■>< 4 )<»)., R ,„ m l i in , , < "> " >' ' ^ « » « <»- ?^> '*"" «PF> " I* hi fn h [in ■- hi iki I L.t -m'- WKE s lo< • ' } k ' •" l '' b 7 s "'««^l pE )> PI i 'II , n ! i J . , i , J. Jap. Bot. 14: 6? rea. Kyong- i.n. Louis. , TI not seen). ili'l kanv-m o I.m I'M T \„l m 11V4 Diels. F L 1901 U.- H.u . * l> ' e n.jj-l l„l,' a 18: 428. : ribed from a . 1910. Species et forma. culthated plant that «as stated to ha\e small leaf blades (7X5 en; fhe name was " . 1911. Illustriertes Ha v ^„ iv Fisher, Jena. (1989). The size of It ill li' -, i«i <»i . 'n rt,„ ( , , „, H „ „ , tl t ^ 1,1,, I 5. debelderorum are scarcely different from A . 1913. Syringa. 297-301, in Planta. A SYNOPSIS OF SOUTH Robert William Cruden AMERICAN ECHEANDIA (ANTHERICACEAE) 1 Eight of the 81 re • ., in South America species occur in \. i. /a.-]...UK.ri«a % t|, ,1 ..,< .i] ,1 [ui- ( . t! ii L»')< , i„ ( , , (11 -M. ' IHu i, 1 .:- t.,^-i . ' .; • : v Anthericaceae. Fifty-nine species occur in Mexico, unique traits that the) shared, e.g., scaled (i.e., ..." • endemic to Central America, and two species occur dts (Cruden, 1999). In both s r H , 1, ki.OM. ,ml « -ul.»c ns ^ ',>,,/„ I ,„ •!., (} l„ lh» ( , „'|p„ lu. ! Ml ' (I! D- I.I H | u I H !v I \l>> >hl! M .i \ » s»'i I IK (i- \< i\ I .. , mx.ciat. the information, f. Rodriguez (HUT); L. R. A 1. Lltj...,,!,!,., «lhll». /„ ''Will .ill" I I.W-^.II Kl ■,/-,- ,),.. ,-s , itllOl ,»lgK I., I.^-th I,. I, Mil genus (Cmden, in prep.), as well as the Old World nutant. Likewise, the anthers of E. weberbaueri ier Gawl. (Poelln.) Cmden are nonversatile, but are held in Echeandia and its New World relatives are line with the In the strongh ', .1 p«-ss,l,K 1,1,,,,] «!„),, M,\.,„oh J93" l»l 1T„ J,,.. .niiasttuo ]!,.,„,- .f ■ ... . :■ j.js ,i, r.dA ,i„, l<>{7l-,!i' L, d.luj«.». L„- a i- iIhIbw) and luzs f «o K Bi , .'.Jlt-i' ntnwesol t!„ ., ,, ",„„„.,,, i,,. | „, ln l jU i al , X] , , - ,-: .- ' with no storage . 37); and the outbred. itlan.Dh ,.l t'l.- unpui.h-k,! ., nu- .jp . ; ,n.lr,l above the middle (Cruden, pers. obs.). Ejndlmism ■ . .' . r - . -L • .oils,. tL.,,1..', aj.i.,1 (j .u,i.s rkU ,. ,ilr fi.m 1 ^ , UO k.n (Fig 1) The three of dehiscence (Crud -30 km noith and noitheast of i */v "i * i ■^F-^ s • 4f Venezuela > ) * r ^v Colombia y£j Ecuador, •H" E. bolivarensis • • E. ciliata E. denticulata E. herrerae E. lehmannii \ J* ▲ E. leucantha ♦ E. pittieri + E. weberbaueri \V \ v^** 1r~"\ ' 400 Km mens from Volcai lar, nearby other on the Serrama de Los Pijiguaos near Los Hi t i I ,,' , i 11 lit Jl i J, mi i suit nJ< II i h. w,i ,'!<», be more widely d .ns erect, cernuous to patent, or nut; ts -„ Vnu. t 1800. TYPE: Echeamlia reflexa (Cav.) Rose, Contr. U.S. Natl. Herb. 10: 93. 1906 [ = , the bases broadly to nar ely ± globose, leaf bases S. apes beai t. I U V\»<\ - I ,,.„ bUi <..lb ulose Uiiomosome learing node, these reduced in size 1994, 1999). not work. Also, because of lap of variation in both floral and \egetathe traits a] ' 5.E. weberbaueri THE SUBGENERA (Cruden, 1994 hromosome numbers are known for an species. Finally, Til' ' »<> l ; --' i< i i ' t I V || c si, id i,~ m.l <>j , m .i.K i„.-ii „i species m the two :•■-■•■ .■•■.. and the anthers free or connate. The South \meiic an The fne Sonfli Yimu. ,m ,,„ . , in s„l ;,„(,. EihewiJm ha\e elliptical to hioadh ,clati\es (Cmden & McVaugh, 1989; Cruden, 1994). greater than 4.5 mm - i V I in .t tlf 35 s,c,^s , llkll , , | 1. > .cl |,la, m.l,i,n r , , ,/ „ Biol Unt - Vmei , Bot .'»• 374 1884, leplacement Pr k-. „ name for A. ciliatu longest basal i flowers (tepals [12-] :,als that are usually seen, US!). ess and the leaf margins ciliate, tents of all the specimens of E. ciliate, rarely dense^ 0orl(or2),if those of E. del a vs. [9-]ll-17 mm . ■■ ■-.- ^ .,...■ 'i i.nli -lio,t-. ,),•'!„. ]s (2 5-6[-7] tW sheet has ,.,1111. ! 1 » f IltRB MUS. PARIS" . : , nutn 1 i luli.i a.,,1 ».» .Li, I - 1 » tint , s A hi [Un,h |<\ ( uutei pels, comm.): •:;■-■.' ([l-]2-7[-9] mm vs. [3-]6-12 mm), and, if present, si les. The scales were shorter cauline leaves (8-21 mm vs. 16-35[- >th a 10X hand lens in good, . • ■ : . .:,-:■ ■ : ■„.-.■ : , ..:■:■■:■ V during or after dehiscence. Finally, E. ciliata is ' r enations. The scapes are \„„ n l..f„, H... I..)iu|wi, Hi h,.* <«ih .,,.» iU , j li(t ,,,,, ,,,,„, rtlre ,„, , ' ' ' ' ' ■ ■■.-. ■:.. :' Herbier Humboldt & Bonpland. AMER1QUE EQUA- Greuter, pers. comm.). BowIvmI ,,]l -t ,e>.n„ Fm.boldl et al., 1815) desrnbed the tlcmeis as K.mli .ho-, .U.U.. « .-.11- .,'nu.r. hiln, u. ili, lfl .t L^'Mnl M ?n.l U t.,.1",.' 1 1 i note-, (« Uieutei no 0657" and included this bnet passage a- a K>1 >" unknown The alisenct . ...... :. ■ ■ , was provided. Earl . , iraboldl and Schultes and Schultes f. (1829: 466) in then tieanueui Bl,11 l' ] " '"- " m ' l "^ "<" 1 r ^V <>< -> »,un|. t , ( t,a,<. .., I „',,/.<„, ,m,n. ' ■ ...u.l,,,,, to estabh-h apostle on;, /' ../..rf « ! .M.I. « • -I..41. ~i~ -t i was associated with the isotype at B-W. Even s. mately des y Kunth (in Humboldt et al., i.,1 . I If. m < s,l „ ,. lt /,, ,/, „,„„<. t M l> ' i I >7G . u ' n 1';.,., \ H.m. . 1. u lu s ,1 d ml .Mt the east ,j.le of the Andes collected, i.e., Suba and Co September 1802 (Sandwith, 1926; St Nunez & Petersen, 1970) and visited oi ■■ .. . Cajamarca and Magdalena to the southwest of Also, plar I Venezuela were [Humboldt et al, 18: ely to be weakh scabrescent with specimens listed below). toward the bas ; lower half, which is )s s„u.. „f ri, l M ,. ^th.ni.g ,.i Kn . ' I l.v nbed the scapes Kunth (in Humboldt et al., 1815) described the ase. This was true filaments of P. ci were most of of the isotypes and 14 of 24 specimens from 1 1<|.I„, ,f,i ,S ,»„*£>/ rV.IHlM.I b h.,„ H-, ,,- U..J1I III D.f.dl.., flJ ' :....'...■.' Colombia and Venezuela (22 plants from nine f ical times occurs in populations) bore nu (see below; see also .•■--.■■, in natural light. be relatively c egions with seasonal '.Mir/ 1 la «r lf ,1 aHuila! K.mll d>,>W> R,.lh 1 «S La- , l<)f,V filiate to ciliai • ,m W,, „ «. m »! <., (,|,ii.,. . ^ ,,, I 1ji.« - -,1 , ,1. i. r longei rnations The leal maigins „l >lu ,j »■ .rum, labels on the isoftpes aie intualh identical The rWnhei, in P. iu - - -i.l I 1 ■ .„. L. M im ■)<- ',-7 mi.. consideied to be lelatneh accmate (see abo\e). «eakh cun ;ral; ovary (2.5-)2 •'• type locality (e.g., 1: n, 1986a); (2) and E. ciliata are r< and without a .ollwhn ,I£ , „ ,.,, mil,, .„ I «U lu ,| '"' ' !l »'- ««< V ' ^™ ( " 1 " m1 '" res and cultivated fields. 2, r.-i.e.m.Iia .1. utu-iilara < t 1 ., ,j. nm n P« ■ "-H'.,. t urn, .N l.i *>., . Uh>. 2750 m, 15 Auj ins - The filaments of the (holotype, COLkisohpes, F!. US!) Figuie 1 *|). < im-ti- <>i L W, uh i. t,nli wsjl.U him, jiN (2: Colombia et Venezuela indigena. (seven gpecim e (thlee speclme ns) Storage areas of roots enlarged 0.5-1.5 cm from the Ir its of most of the Penman flat to falcate, entii reh ciliate or fe« specimei ,ore small, narrow, long-ciliate, enations to 0.15(-0.26) mm; cauline ti ;ins of most of the lea\es to 2(oi 3), if piesent, the lowest 1.3-3.4 P : to long-ciliate (79 i ■-.••'■ ,' Inanches; main a\i ith (3 or)4 to of E. Jenticu of E ciliata were or patent, with tepals shorter (9-13[-15] ram vs. 12.5- Ku . i.anl. Uj. i ml < liuu 18[-20] mm) and shorter (2-6 1926, F. L. Herrera 1012a (holotype, US!). [-6.5] mra vs. [5.5-]( urther, the Figure 1. iL,,. Ji,™! 'i.ii -U >U) ilh less than twice the length of the branlhedTmai] ll(to 13) flow -i* i.nMi-,1 ...I tho,. 4 ' .,1, I 1, ,i.>,< ...i ! £. ' ' 'VlfteT'LliWnce" eral; ovaiy 2-1.5(-5) mm; style 2-6 11 of 11 floweis Horn s ten plant,, one population, tli. -t.-ii-us ,.i. 1 In Mir mi tli. Etymology. The specific epithet describes the to or barely exceeding the anthers. g to narrowly oblong, 7-15 X 4^6 mm. and habitat. This species is widely gy. In flower December to April. mdia herrerae are 1,»< i7J/ 'f OL US) Ubate lUOkinIS oiBom.' K. " m ^ V _ '" , """ irl " l *'"< 1 '" ....■■ ^ ^ .,.-. '.;.: 1 I «> ^ •' 1> i l " ■!'" ' O ^ l" , u.ns 'i 1 t l lM ' l„n ,1, ., <„, Rodriguez 681 (U). mens) leaf n plants «eie easih e the Ik-ii.1iu ; > lint -» o V ,t ,; "T (...LI (,H The erect flowers and shorter (2-6[-6.5] mm [5.5-]6-ll mm long), geniculate shies distinguish 4. Frlu-iiiulia WmMimii .rja'.en Maiais & Reilh. Kew Bull. 32: 662. 1978. Basionym: Anthericum lehmannii Baker as "lehmanm," Bot. Jahrb. Syst. 8: 208. 1887. TYPE: Ecuador. Rare on alluvial [soil] near Malchingui, S slope of [Volcan] Mojanda, 2800 m, 28 Jan. 1881, F. C. Lehmann 429a (holotvpe, BM!, fragm. K!; isotype, G!). Figure 1. m long) that terrerae occurred at high 's. 2300-2700 m). Etymology. The spec scabrous, whereas the styles, E. herrerae had ] cm vs. 34-59[-68] cm ;hiscence. Further, E. ous oi Tuth a fe« small en. levations (2800-3824 m z[tI:vi^K^B. epithet honors Fortunato lia (1875-1945), well- !arAnt S omo° f Abad ny del ely in the region around , nberg 1195 (U); (Mt I Mai 107^ Bnmel 1,21 (MO) \ncahuasi NW ' ., . . , .. , t .„ I ,l,e tlnuh. . u«l, _T- SO km to the northeast oi flwl'P ir(u')"7u I." ' "-/(""(111 ''-'^ , ->' l » ,.»» Hi . 11,, | r ' i i ( »-« ' r i I ^ ( r , i it. ,,. .IIl « I ca _ 60 km east , )f HlK «™.'''.>W-'. I Jff „ t ,, n j'.,n Uan ,1270(.m lh» 1 M,hl.lc.l,.,-. l c Sl .| presence of the isotyp. Etymology. The lors Friedrich Carl Lehmann (1SS0- I'H) 1 .J , t JJ, , ltd „..M\ jn /-/> ,a„ w PI ,.,h „J /•,/, ,i ',«;„■/« .«„ ,.,< and other species now included in Echeandia. h 42 mm wide) basal ' ,, limih[ii ', ,„'/, , ^ , , n',, \__i_-_, ,'J IWt it I.I in ..i.m !s (l iiii. s^n, _, ,a,lh,i. Ijlm.-Ml- - Lol™«l.anrJ*,l.„.„, V. II,. » n l-ymn „„,_ t , ,, , , , , ,,,,„!„,,„ , wm Peru. Huancavelica: Tayacaja, valley of the Mantaro River below Surcubamba, 2300- 2400 m, 14 Mar. 1913, A. Weberbauer 6481 i u .n. „m,..i >.ii.i I ,sm,«, * , ;- ;-. .'. • . . rhizome, 1.5-3 cm 1 7 to 12(to 16), (13 of 14 plai le plant bore small, it to 2, if pitsent, the West lb-35 general, flat and wider than the falcate basal leaves of 5 ([2-]2.5-4 mm vs. 1-2.5 mm Ion i ,,i„ilo 1 1 3-J6-12 mm vs. [l-]2-7[-9] mm wide). the upper 2- or 3-flowered. Flowers yellow; tepals 1 >v.^ tlit Nurli ■ r l! -t ,,„ n- M2 .( I", of 15 plants). Als pical; ovary 2-2.5 mm; style ciliata were three to four times the length of the 8.5 mm. C ig, 10-11 X 6. th i imi tl 1 ,1 f ll, I, u - basal leaf Specimens of E uebeibauen weie easil !2o(iO-K;i2l in ,, 23(>U-2 ■ ' M«i p. .t,,,,,, ot !>,<», ot tl, Tl,,!,,,^ 1 ,' \l n.n first collection of this species. combinal (10-11 r, areas that are Unn «i,!,>jm.iH il. JO 1 i yellow-flowered, which is rare in the subgenus (Cmden, . 10 '35'N, 66°45'W a y, a little d 30U ! ,V< t of Los I • ,e in th lortcd i lora (A. Rich.) Ku nth. Phenology. In flo\ i by the :rs, relatively s : < mdc n"l900). The subgeneric epith, urn ^iln (i ( ... ,1001-K,.,",,. distil ih Lih.„ ., » <- 1,1 ■ 1 il -£,/,, / / / ,n "> ; ■ -uania de species m subgenus /Wsco«™ mil, ti~- , .Lili, w »h, ip „ !). Figure 1. subgenus Msa i or rare in subgenus Echeamlia. Finalh, £ bolivaremis occurs belo« 80(- in, a- an euhr , t tin mli. 1 2"i spe, i. - m which has a temperate climate. La Guacamaya, Groger & Berg 1064 (MO). ^^ ^ ^ ^^ (lg4()) The design; ,e was necessan I,-... I.Mf. ,n _•:-, 1.JL.) IM'l n.^.nju.piotologucoi Otto (1840. mlu- ' IT. i-l. 5-.. 1 Fevdlei 1549 (neohpc, desigmt ] n Jn dlust.al., n- - 1 ' The most like loritz's specimen, if Storage areas of i irged 3-6 cm one were mac iC, or possibly W cauline leaves 1 t< pers. coram.), or W (B. ^ allnofei, pels. usualh reflexed :ar, (2.5-)4- made a collection of this species. If he did, it is 4.5-5.5 mm, rarely globose, ca. 5 X 5 mm. suggested as a implicates (Stafleu & (rtT-th ' ' ^ "^ ^~ ^- rarely to 1500 m. Etymology. The specific epithet describes the & Ortega 821). n- ■ vh j- l ,h ■ fri, ♦ lel rluL.lH ^ |)( .lUHMlulllVill, llll. tl. . ■>,)- . , ib ^ , „ lV »,, _ /., ,,\<, ,1\M ,„, f w« 4, mu ,J J ft«7 ideitified In t«, bilt H.1U1-, iHi'i 1,, „ , ,\l , /,, v • 1 ,\JU) V, ,ui n, , A, ,/, , i "/;-, of the roots that de i the rhizome. (F) ' The scaled filament , that develop - ■■• '■■ ••-. develop 1-2 cm fi Purdie (s.n.) Boquete towaid DaMcl, in sa\anna neai locb included here with reservation. Nomenclature. The Fendler specimen was select- Storage areas of roots 1-1.5 cm, enlarged 1-2 cm ed to be the neohpe because it is a complete from the rhizome. Basal lea\es 5 to 11, (18-) specimen that inc ,, 29-41 cm X (4-)ll-20 mm, narrowly lanceolate. 2-3 mm. Capsules ..5-7.5 X ca orld Liliaceae. Nord. J. I leucantha. Additional specimens examined. COLOMBIA. Valle del *- the Flowering Plants. i\kU i EJ 775-788 Mc tz h Di 1 M / < I , u ^1 . n Raveii^'p' WsT'Ne iliaceae. II. Phytologia . 1987. Dm™ ko and Dior a, two new genera of 7. E. leucantha Klotzsch m P. Miller Jot. 92: 185-193. Ku ) il H l'!<(. 'hi (( V], "VJi-.i-- J Pm n Appendix 2. Index to exsiccatae. Ho .1,1 l. UJ rtmeian in enezue a. u. i.e. i oim. pp. - . - _ :■",.•• >W,<, ,i ( ' DM »|, f l,ii,,, „,,., .', ,, i. ' ill I..,,. Jj07 1 1/ , 1'U'l {M McDamel : 387-394. Appendix 1. List of accer THREE NEW SPECIES AND A NOMENCLATURAL SYNOPSIS OF URERA (URTICACEAE) FROM MESOAMERICA 1 1 ', > i. - / , ( i .1 h hi's, ., s, ] >< l> -i mi „ li > t ITU N li .1 I -r ii '„ .m.l mil, liuf Kd'i iik-1 i, .|ii „U> m n,,.,.],,, ,nl dl'CM. • ■ : ...... ^ , :. Africa, Aushala-ia. and the Pacitu l,Li,,k> !m ,-. li. i, e fh ,iN ,., il Jm*1 , ? . »i,tl, \.n. ...... W-i ' Mesoumrricaimpioto'.hilh^uloiJ ]>>> Hat .It I {..oil based his i or that he did see the hpt rolle. (ion >t U pn.,1 to 1975, and that VI Hot vliiuimi . "I J7"~ mi J.«'im.i VI. Hort. Schoenbr. 3: t. 386. 1798. EVITYVE: Urera alceifolia (Poii Local names. Bringa mosa (Panama: C. White- foord & A. Eddy 249 (Guatemala: J. A. Steyermark 38770 Rooij [1975: 304], P!). handler 11202, F). chichicaste 34 ' 1817 ' as "tiliaefoliar TYPE: Colombia, ruwimgui iili ndnacioi i ( ^lantuey zi<><><), un, t (l| iip)i ^ (i ,^ (lf ]{( 11.17s vsi.l J • i NY), cow itch (Belize: 1856. Basionym: Urtica (i A,. f a. -.»...*. ,i. s a, ,h,n ir.„- lia because although de Rooij (1975: 306) cites . n.it'i, .1 rt'M.I ...uHbetiacedeithei at MlM. Esser. pels, comm.) - "154 BM Ni.ai.mua P C Stox//ei at W). Based on the type illustration, il is not possible E. Sandoval & R. Chinchilla 501. M< )|. chichicaste (Liebm.) Wedd. on sterile characters alone. Materia] Chinchilla 1182. MO I. chichicaste rojo picapica (El examined that had been determined a. I corallina Salvador: 0. 17, -.-. ISF225), MO), pan (including the holotvpe) In Weddel and as U. caliente (Honduras C Nelson et al 3955, BM), ype i> >,i; led because the (Costa Rica: J. A. Echerem, I 268, type illustration, although accurate, is not sufficient to Rica / 1 Echeverria C. 268. F). pan caliente (El this species. (Mexico: A. Mende: G. 8945, BM). and distribution. Cloud fo shade 'i'SuJ7\S2iZ^XuZ2^ Z , 1, *Ue p f|1 . -Umawh p. .Imiflr I,,.,... Ii. .1 I,. ]»..^ or o U.se 01 unbundled at base for 2-20 mm; (2) for U. verucosa, lea\es bullate, staminate 7-98 mm. the naked eye, veins visible for 2/3 of sent in the axils of " !!., *l-Hi mm, bearing 160 to s of 10 to 35, pedic. ,1 P.,1 j '„.!,] „ « „' "t ,, , t nill.^s.,-.],^ lt„ ! ,,-1 stem, the peduncle branched imera, ,1 liidm & M. Pereira 3955 (BM, MO). MEXICO. to b . 1 nbi iik u ]„<> lT5\I. MO. PM\. H.Z) PERU , ., , ,. ,, . . , ., ., i- i i e, the pedicels whe, 3. Urera fenestrata A. K. Monro & Al. Rodr., : Figure 1A-E. Stlgma pem ax shrub, vine, or small slen. ike, ca. 1.5 X 1 mm, red-pink, rching 2-10 m, stems not red-viol when fresh. • ■■.. ■ '. )wing to the fact tlv,( tl,- t ff,f, Lis U-en ' >8 times in several localities in Costa Rica and ....-equal to 5 mm diam. hollow, but Q u 1 JM i I'M M i I 1 L . u. 111 7 , i 4-parted. this species in th« ; of these a ' Adilable at the Ii stituto Na< 10 lal de Biu'gn & R L Li, w «C >' ,J , 1. w. ■- na i FifC A. Alajuela: La Palma de San Estui 11 . , 11 1 >i ' I I f, JM '1 Ramon, A M. Brmes 736 (5544) (CR, F, NY); La Palma de IM> 41 Kullnl. (>!*]. ".In,. \T W Vi mi A.D \ lib ) > 1 ^ r 1 Ml LI 1 \r k I Ii I ! I , 1 !., ,1 . .I.n Hi ',ii.il. !. '.... 7 ',., . '.•i< itque ructu >,'.,..-,,>/ i/.'(, ilfl|,'M/,v,..ni/,i'„,,»,Hi ""' ,' ^ Y!' 1 . i'Ti ,'",', '-""hs i U. .,.„ , M .l.' f l. /? < ')/(>«. 'SUV',, i i ML, ' ' " '", \ ,'* lanceolate, for scent; petioles 6-90 4. Ur.ra "Lil.iiiiMida V. W . Sl.-iniii.. \rl.i Bot. X ca. 0.5 mi pubescent toward Mex. 71: 22. I ico. Veracruz: leaf base, th i mm, strongly ap- 1EXU not seen). chartaceous to ixial surface sparsely G. 9054, BM). straight; the . m, inflated, densely MS ,,),«! f ,M 1..1I l.)l-,t ' u I f.h< -fl> v/.,.„ I"l"'ll""" t'>'.M<..- , ,sl, I o scattered and parallel to the * ems; priman * ems 3. y and occasionally quarternary v 3 the naked eye, the lateral primary veins I hanoiJu. m its 1,1 is i v . s f h ,,, 1 s f llt I, , j . , , ,n s llt ! , rh lV1 l liht serondan \ems < ^ ■ ■■-■■■ ' ■■..■■..■.. U 1 f 11 «s (l)lci I mhbnu ula !,!(>_, I il i i 1 , minutely forked; p ham.; achene cence 5 _ 40 „ flowers in a weakly 20 mm; stipule ape >de 0.375 mm to 2 \ per stem peduncle branched diam.; achene surface smooth. to base or unb , 2-12 mm (6-17 mm , *l. JW, ,= .-,•„, XI Ur,.,„ ', I W , 111( beanng 30 to 250 flowers (59 to 67 in fruit) in cssile to 0.25 X 1 50-1 75 mm mm .h.lH p ,< t . anthtsis tepals 4, ca. 2 mm; stamens and pistillode not seen. 5. Urera guanaca 1-0.6 X 0.25-0.6 mm, lateral ii ,-\ II.-., /,., .„;..,.,. .,.„.,,,. ,., I,.-,,, ',;':'■ u ',',' , l :i.' : ';')! ; ;' '; ",■""";" '!"'"'■ - ; ■;• ; - •;• / records for the spc ; from the 12 Bot. 24: iuatemala. Quetzal- n Local names. Chichicaste (Guatemala: P. C. records of this species have been collected. m ) In mistl.k l\ I, I, >nlns,.l v',»l -l.j ..U ,. .oltiM.l rj, . , ,„„! , , „ m- „1 l!,r • , . .iMityui^hnl iloM.il l<„f ,i;.ia.-.-Ju jiu-nlh Hill, Jl " -\\, I.ihvio: ' f ., // I /■ H' < . ( , <-.u* 1A t(>2">i 1 i imu;-li' 320 (GH, MEXU). pubescent, the hairs 0.25-1 mm, erect or weakly appressed, weakly curved or crooked, occasionally 7. Urera laciniaU d., Ann. Sci. !l-84 >< 5-10 n [Colombia] NouveJle Grenade. "Quindui" [Quin- & leP "- holl< ™ *'*' ' '" l0 l " !il ( il '™ 1 " Hl ""- •' «'"J l 1 j„n ud I , R(..i, [19" ()„| P '- 111 »< 1 ' » 1" !<- nii'il ! #00281783!). sparsely pub: nt; petioles 11-90 nil T J lu-l ,' > n \ ,, , , / i! ,1] , , lllM ,| , , H , j| ,„ j0 i j „ „ tl llt type, designated In i' k<> „ 1 1 >- r . ) J Pi I, ,t I I . r "'•-^j ' )-J 10 r 1 .1 ■ ■ . . ■ nioides, were publish , j. 1 ' ( < > d,n I -t il n *' inn ,. s i, i < ,),.,]] i li t. 1,1 .. - 985 : t \, 0.5(1) mm, appressed or erect, weakly curv< collected from rl n sea Wl to sionalh oblong, parallel to the \eins, occasiona ! ust, R„ tl . P,ic,.i. L..]oi.il.|,i m! Pun lo B..li\i.t |nui,u> K. quuil. ma v t.-m, , i-,,l,le lo il„- i.al -fl « es 0.25-0.5 mm. Si 12 per stem, pedi ubes^nf f ™ e m J^ :ence 19-56 mm, be '* " '^ ''» ' '' ' '"'"" '1 *1 *l (, l I^'V*J\ s I S „i,W .t l.'iiKuiJi i,iu, Inn Jim. i b.in. n, : '*-' N l '\ "'"« '" I ' ! " -' ' !l ,H « t. l ,„1 , ot ,1, h,t.„l„ lilHiin ,h fh„, , nieste del mismo 10 05'20 % ||,1 VYl";'/" T ljlr'2"! 84°24'20"W, 1000-1200 m, 21 N >* !•>;;,> 6 -i. „, m ..te tU , ls . , L >'. • I '" , H,,i .i J,-<, i'lol-mp IMT imi ¥ :ure 2A-F. "\l" f ' ] ' I / ' I ,1 "i 1 " , l""" hi" 1. t i LI 1 1 I! ill J i I I ii L L 1 I M ,' ' i-' J , "i" , ,"' M ' SCa ^ ml " Mistenos del Tenon... ./ I «,«>_- .S'ZJ (INB). Um ,. .1.- Lo- achene 123-13 l-l^„n, ^jnm.tucalh Lhll( , Rpfu _ 1(1 Nltm d de Vlda Silvestie Caf„. 'N. ,i,. KiV elliptic LeeJ shaped, uiU f ni < ih In H \ h i, up , ilf |u [( tf , /% y/) A ; ;/ jW |M J( , ( , . \.K,„ W.1.A' IF /,,••< 2-2L' ,Fl I'.v.L Z. i accojdmgtoIUCNr.II . >il< u iTUHJ 2«.>], \ v u >, Ulli ( . n . » . ' .,1 - i - 50 times in several localities across Central and South ^ ■■ ■ ■ ■ . is i a „ (1 Daha ri h?=f> R- T l„: r n ca„..„ „, (0.5 mm or less). Tl: :; '"''-• *■ ■/'<»"'<'- »■ W>? (F); resembles £< #kn [, "' ' ! >" (,(, <' 2, , 4 .XI *' i, ,, ,/ I T. D. Penningtor alone mriado/orerlookine Caribbean "lone Sei^Ne^a 18 28 ' N yS 10 ' W ' 35 ° m > 4 A P r - 1981 ' J - L d'.i.v.is i -I..A., li.i.H-nn .. H,,n .In. ins \\oili V\j. and onh photo- j,.1j, .,) ll h.W-p, o.Jd be cated (F, MO). The neotype ws sole. 1 it it includes go,„ ( I. it ,,i 1 t. .hi. nnlerial and was from the same country as the holotype. /--. names. Bilsimtezla (Mexico: 1 Mende: T. BM). chichica-te (Guatemala: P. C. Stan; I : M. co: I*/. Heath & A. Long M - ->2, ,10) ;aki snlsimtez laa (Mexico: A. Mende: T. 6238, BM), tzotzniz zul >inilez.4. Mende: T. 9066. : i .asco Vera '1 -.l> .drt. ,.,. Tfv .do, (2001) indicates U. tu, mala, Honduras, El iolni.., Bivil, and Paiagi flowers 4-parted. -1 cuj >',,!■•„>, -wnid RHI'l' M..UU .:,'.,jN% H^ i676 (K, MOL). £ H. 5/6, (k, (OK.M.l 4 Pi H u no I . n < U. general. Pp. xiii-xiv in G. Davidse, M. S. Sousa & A. 0. 7V» y for 5343 (K). de Rooij, M. J. M. 1975. Vrera. Pp. 301-309 m J. I 10. Urera verrucosa (Liebm.) V. W. Steinm., Acta ^"J," J"'' id. '~ ' l "" rS) ' F1 ° ra ° f Bot. Mex. 71 m: Urtica j.„ |','|' \ | „.',„ .'.',' ,, \i atura l Hisi verrucosa Liebm., Kongel. Danske Vidensk. CII. ■ ti...i- l>>, , ,,. Depart Selsk. Skr., Natu ;er. 5, 2: \L .-.„■■,,.- N.u History,! 295. 1851. LVera caracasajia var. tomentosa ' ''V,',',', ' ' " l - /lx,ta, " /( - )llen -l eg. superfl. TYPE: Costa Rica. Cartago: Irasri, :MK\ 'in P.' R. Crane' & S. Bl Oeraterf 24284 (holotype, C!). <"»'• ^stemalk-s. and F,,-.-il Local names. Chic 'ierber 333, «B. Oxford Science Publicatic BM). GaudTchaud-Beaupre. C. 1830. coffee farms, Pinus-Quercus-Liqui.laml,,,,- l,,iv,|. from Plnskiemie' 6 . par m! H" 500-2800 m. Mexico (Chiapas, Veracruz;, l^uaLe- Pp. 4yu-49< Pillel-ame, Pans. 8. U. lianoidt ■ \> ' ,t, 14 '_' ai->< ^ , < ,i ' ,'; L.-.27 I Li!' 2879 (9); Gentry, A. 2043 Gilly, C.L& Hernandez X., E. 320 (5); Gomez L 5535 ( i, C. 2327 (8); G 2 (9), 7267 (4), 9066 (9); ,' \ .r- il. 'J.-'.Oi ,520 (9), 6928 (9), (it j, 20 [5i *,r<>iL> , .' ] . ,.', . , OJO w , ^ j w , «™ , „. ,™ (1 / ,". i'»™pl / _ J (10) 2578(10), , ■ S M /fa : ! »/. 29264 (8), 3535 (2), 39£ 92 (3). \12 (7), 27740 (10); Opler R. 1279 (10). Oufc, ./. A. 770 (3): Tvnupp. .S. 70Z6 (8). 7 J35 (<);: , " ' _' 'm'.i 2>T, Km -, G. L. ei ,3782 (10), 4742 (10), 5t ' , ,lh ,') "r ("I „- 111 h I ' » 1/ /' Hg/ri, C. s.n. (1). A REVIEW OF THE GENUS Amy PooP DISTICTELLA (BIGNONIACEAE) 12 genus of 18 species in the tribe Bignonieae that can observed in this study had bifoliolate leaves, with a ' , » • , .M.TH.ndU !,, - r.,i„h„Ltt ' . "• ■ ", .'•■ . , : compound leases will, i\w ^ iwi„.l k tKt m.i lihr ! h Id- M..,l liu. u,< onspir uous pseudostipules that into a tendril. The exceptions are D. mo 4. H. Gentry and D. «„! Sj,. „,„i'),„ III . ,_,„,< 1 e ,, s,, f ,j ,i,,) , Us sensu Bmeau (Bureau, 1864). glaziovii (Burt . H. Gentry). In their »-.,< s . ,. , (JcqiHi 1= « ' , 11 j A I, |M,s ,, B wp, 'i, ii .- tuwnensn (= Dis- and three. oi " , 'M n.lulr i ft ; > t , /;V - ^ ..') 1 )<,_' tt f \>, / // n (' in (= Arrabidaea arthrerion), B. elongata Vahl f., and Ami he subtribe was PitheiotUvnim , i i , "I , ;,i tf.l.d -V . ;. ,,< l.m tV tin uis „>i<1 l.i-n hl-ts .1 Dnluhs ami mam Kraenzl. = Distict K. Schum.; species nf Distictella (such as I>. arenaria, D. Sprague and Sa ublished the .■:■■■■ iu,k< !■«. .IditinnaliecnmbmatinnsmDzs^RfW • L> c.mhi.l „,m pitsmr lb. fo Pieties nf i),s- published five new species: i3 pH, / »'r -■ / ^ . 1 1 , » li in he! study. Distictella fits intn her "group 4," which IN.,1 ,fli I'M?,., = /Wr '„,,.,„. „„,'„, S,„|*j t l, .s m ^ ill, !.•»,, IU.I Z) 1, , „ ,\ Mlh\ L"!OI, lius th W .l',|l I t>l..f f •' ''«],,,! 1^.11,, h.i,< i.l i |„1 < • a.lhiii, i^ipu.hll, ih .1 • InliTji Hi '.tn co, . ])f - fig. 4). Urbi uh described and illustrated the wsa var. translucida between these t«o closeh lelated (as D. bioadua^ana), but «ith bulges on the reticulation extending into the areoles (Urban, >) outlined the genenc diffeiences 1916b: tab. 21, fig. 3). Sandwith (1965: 412) ransferred to Distictis post-Bureau Gentry and T 1 to multi-trif •a in the group are and Disd rith (1965) and Gei ously suggested as closely re] i, ^i.otliei ..t-nus suggested as (,Mitn, 197 ku, h • poJien 1 ies, and ha\e a different ftpe of cambial Jcto and Distictella have non-echinate capsules jtfrazum has e, lunate capsules (except for P. m (Veil.) A. Pool) that m» I , n,l n ill rH huh i (hnacteirzedhv each ' Generic Placement Gentry (19741 flowers of Distictella as falling int, n-type or xylocopid Glaziovia, and .1? 'tis was not the calyx), am it the base), with an ft tl« jsul K il f tl t « ,j| ! a „ ,| ] sfl til ..,.!.< ilm. int. ,r at all flattened. Houever, in othe. ,,,(. rN //,,, z N t Sll tL tl ,- ad 1. ,1 < .1. >.,t ^, al Il>e x> I k,,; „l, U .,n does fit the descrip ;teniinae. The "converted frc .ate visitors." : = PltlK.'l'rlMmtll , ■ ■ I ■ I,.- tli^l'l. .Il"« .. Review of Distictella (Bignoniaceae) .st of the species ha\e terminal (or ........ . ..,•■•.... , •' u 4,-l.ihon (. .iiil]i bus. (v.ra.los. or pampas): ,„ ttl ,i ,„,ti i iti . t n < I ! <<"' Me lli, i. si i) |, , ,1 in forest. Distictella , • ij, KnfMi l*ii/il I > , m mi " « «" i liVl--o»- ni j) , ,kwc, Q60-1400 m, laevis, D. magnoliij ot 950 m). • •■ . ^. . : .;-. seed wings. ulata, D. can Iso southeast Vene- overlap with D. mansoana and D. ra Distictella campinac and D. laeris (Fig. 1A) are botl Amazonas, Brazil, where il is known from similar non-inundated forests, apparently not associated with to inundated non-white sand areas. Both D. lohman- tricha (also north-Atlantic Brazil; Fig. 2A), D. elon- niae (Fig. 2A) and D. reticulata (Fig. 1A) are found in gala (also Brazil), D. obovata (Fig. 1A), D. paiken and D. cremersii (al Para, Brazil; ■ ■' rsii All specimens examined by Gentry and as part of , ,n ill. l'V.f >88), D. elongata of associated with sch nd quartzite of the Orapu-Bonid ist, from: AAU, B CM, F, FTG, G, GH, oro GOET, K, L, J S, TEX, U, UPS, US, stictella racemosa i *n from seasonally inundated fore '^ fleered Jom'thi! Y.tTbaTIoteL'ite the dTstnlm- i»l u r >^h,,, tricha and D. racemosa var. tram ,.i.l rlu li 1^ t<- Mi ml nd L -k, t,- rnrpmnsn var rnrpmns,, and D rarpmn*,, var mansoana (DC.) Urb. (lectori, pe, designated In 1965: 412). late, truncal, minute denticules, usualh «ith glandular field itely pubescent abtne c.lindric base, intemall illy densely minutely pubescent (rarely nearl lis a id D. magnoliifolia have found above 400 i 100-700 m elevati on and £>. monophylla ra 100-1500 m eleva tion. ensis (Fig. 2A) is the on! absent, < •< lAst, t*J>, k-,l, m from the Choco and Volume 96, Number 2 Pool 2009 Review of Distictella (Bignoniaceae) stammod. mseited H ipp.fMi,!., K ,1 ,m. Jm I i i. ,nl,m. m , , , . .1 r.mt -. If .I0.......1I..I I. ..u M.11 1^.11 < 01 mngs flell to puoih denuw/r. ,1 l-.i. ,r,l i, Annals of the Missouri Botanical Garden 1. Di.li.tolld iren-o-u I II ',.i,rn M. i, IV . ,']ti. . n 01 amending liirhomes, tiichnn,,, rt hu. York Bot. Ga YPE: Vene- tan, or ferruginous, 0.1-0.7 mm; pseudostipules ing with higher order ven jaxialh, both surfaces d Review of Distictella (Bignoniaceae) tscus ton Distutdhio mi n i -I unihn U '" i m,.,!K l.niLH tiu.i|..-tjnlps). capsule much tlnckei (12- , wiiiacus j-_ cm Viabital from I), chocoemis, which is endemic to the wet ceous, tube 3.2-4.2 cm with cylindric base 9-12 mm Am; long and mouth 14-15 mm wide, externally and ^ internalh pubescent with minute trichomes abtne 7 j". ns and st; in-.e.t.nl "MO mm from base of corolla tube. sin, .» %-J " uiiM ,Ti 9 mtL,H-ol>i. Cp-id, .bU.,- not curved, 7.5-11 X 2.5-3 cm, ca. 0.3 cm diam.. 2. Disli.l.lL, <:ini|>iii:u- A. s.n base acute and I irly Brasil. Sci. 7: 120. „,„,. ,,fi„,i 100 to 700 m Figuie2\ bunchlets, h Ml, t, ll„,ti, u ,i , with the midrib and extending m d nenh -tiaicht I,., a n -i i 1 » I d, ) mi mm, is, It. ill ll.t.,,1 ms -omefuus -h h lv som tmies Mth scatteied «l.nd ,1 , , 1 i .t . i , * »\ D >, < Os ' ,,• .- I " glandular fields at base and sometimes at apex or *< "#>& is endemic to dwl- - a omas or pampas with :- ,„„ ,,, j,d l| ltn i.urdcd .,,« s „'•,., d> -rnbed as season Jh colorless trichomes hghtb apples, i bud. 1 u I, Ms *.' W (MO) 1.5 to 2 pairs; cc idibular from externally densely j lute trichomes H wy., W m, 12 Jan. 1979, A. above glabrous base, internalh puberulent and #. &,„„ tf9 (holotype, COL le t'l f Inn hhts I i I [ 1 n , [ , .t , Ihpti. ,(0-)U-21 I.: 3-17-11 i ...i.M.bron.i.,.,)!!-. £ I iti df'.sl, l.j .mi sul.oii,, ... , ,{a d). umsin.1 , 11 1,1 • , ,• , ,• • i < -r ovary ca. 3 X 1.3-1.5 mm, , lliptic, not curved, ca. 5 X 2.2 cm, r a 4 cm diam . base >, i t< chomes, glands sc .Ite.ed ..d.li m • ! i ! ' . id ui Jm Review of Distictella (Bignoniaceae) Ipjpn ,,,!!,,!, l )10V n «ith pubescence like thai ..[ like tims,- ..t J^.mchlets, or sometimes branchlets, bracteoles absent or not seen; calyx (7-) 0.3 mm; leaflets elliptic, 10.3-16 X : li« 1, 2 «,. _. i ,m . toil * lu'l, orbicular. Fruit unknown. and then roun, 1 with one side acute Discussion Distictella chocoensis is most similar "' thicL Seeds 14 " 16 stipe (see discussion of D. a lOT to SurfaCe ' cjsle.ii OJumbia, «here it is usually found in white Sun. un. 1 i i. . H ltl de Marajo, Para, sand thickets, shrubln islands in sa\ annas, and Brazil. It is don; the elevation 59) also described ^MarchandApnl.Flowenng "B L fZ\MO™ Lllpapel/Buena¥en - ^^ Fioweii ^ ^^^^^ ^ ^.^ ^_ mrai is not ruit and vegetative Tampoc, le long des Criques pres de la ( nqut t,an-lu,i,lu the Ja.nl - i i lis n nil., m.l ill i t MO!; isotype, CAY not seen). f mits an d set mora var. rarcmosa. ner (8-16 X 3.: .. .111.1 'ihr ^.'.1- ,1.1- M^-is t « branchlets, br not seen; calyx 8- 5. DisticteUa cun ith, Kew Bull. curved, 6-6.7 X 2.5-2.7 cm, ca. 0.4^0.6 cm diam., "Para," s.d., s. coll., s.n. (holotype, P not seen, trie; minent, wall 1.5- 9.196.16!, G fragm. not seen). tnchomes like tho or as « ith numerous large ...... ■:,:■■ leaflets narrow h sp sng, or rarely Figure 1A. oblong or elliptic, 4.7-8(11) X 1.8-3.3(4) cm, veins, each lateral v !5°(60°) angle with the nudiib and cumng to* aid the apex, DxsLUsswn The only collection information on the ,j]^.l AkmIK i. I oll.ei win, iuuhmm..] t„ ILt o, ilh..wt,in P.i. Jl J m . ,li..n, ,1 111 h i lin iK I pl.k i u II um'i lTir-1 I uirilnrtht Pin ontheftpespermi 1 1 n « t I -■: ... ;":.. ... ■ ;-■■■ •'.■•. emarginated or api< ,ny. However, while Review of Distictella (Bignoniaceae) Orbigny did travel in this genera] vicinity, I found no j,. h „A. ,j| ., „,, m.i.,..^. „ddMalh and laised ■• : : : .,- : : ■•/ , . ..- ; ,, -,;. ..... e 1.9 mm, dense to scattered on main veins and ' . • ..■■'.■' Sandwith (1962), identified specimens of D. cumpinue without scatteitd .Liios ,1 mil -mfate nith dense, also studied. The most striking difference between D. margin flat, lias, , , n „< „,. m., ,i leaflets, which die iniMU,! v-nl v., mo,. |,„n I p, ■ !.,... f Mtl. j„. In^-mx ( >i ,-i,li 90°) in £>. compmoe 'uncle 2-4.5 mm wide at base, from the seed bod. n, I) #",;.■, ?> f<^'< / •>,>,!- {_| , ]<»-lJ ,„-,, s,ib<<,u, „ lls , pubescent with B. Ih > t , sr campinas. stamens and staminode, lobes 10-17 X 15-25 mm; til ca. 4.1 cm, ovary Prov^es^Eco^trEfRef^r^SOOmdSE^fak 4 " 5 X 2 " 2 ' 5 mm ' stipe Ca ' X mm ' stigma slllmlate ' nous with min les, somewhat warty, 6. Distictella dasytricha Sandwith, Kew Bull. e- lent, wall 1-1.5 mm 1953: 476. 19. mm, wings poorh .' n I ita it! pnl sc eh!, I 1 i l I. m H t , h hi- " 20 mm, with pubescence like that of branchlets; Discussion. Gentn identified numerous sterile 9.9 cm, subcoria.- or 5(6 or 7) Peru, Bolhia, and Venezuela) as Distictella aff. pairs of lateral vein a, and included D. JasUricha in his ■■ : . •,.....,•..■■ collections differ from the ki «eakh ciming 01 ascending; tiichomes tan, 0.1- very loose network. Jso tend to be X 3.5-8.2 cm bcoriaceous, 3 to 5 V nezu, a] - clasUricha. Distictella dasytricha is similar to D. reticulata in its connecting imdub „,|Ii1pIm hh, to lcssei extent ■•■■■ n idnband .. similar length to ichomes to 0.1mm long and Review of Distictella (Bignoniaceae) slighth Epical (.olle.tjwifi.rn, \uu- ..la-, R nl, «>J im . W m -..am in.ll. ..Imi^.I Phenology. Flowering January, February, Jul ind December; fruiting June. ,.t tl k . ,|,._ iri .e : holotyi Je RR , Orkimii i 80344) and - .wing th, ' . 938 ■ ideSoE fleyde 597 (MO). rj',/.,.'/i"l, .'„)'.",. The general di with) A. H. Gen New York R. !9: 274. 1978. I Distictella monophlla ; var. laevis :: Mem. New York Rot. G ard. 9: 362. 195 ■ ' ■ • ' ■ ' ' • otl.,.m.hlM-. |»-N..]..l.s.,l,vj, t lo !<-_' mm. ,.nl.M- and anastomosing with the higher order venation to . warty, and the seed 13-20 X 32- to 7 pairs of la ; at a 45° angle with „ jii.1 \enat..!, I. ii i th In... (].- nub.L.il.,,1 j !v,^ >i jid.hjl. <-AU sf..ir,i ! ■, :::!:;::; , d 1/3 d »t T , *',!!,: ,™r\,f™ ..Uli ul J..-1, aa.i .(email; lE p5s 1-2 mm, stipe 0.5 ate. Capsule F. da Silt I; isotype, INPA not oblong, not curved, 5-6.3 X ca. 2.5 cm, ca. 0.3 cm seen ). Figure 3A-C. thick, base rounded. s compressed, 1.x. ol „K r m, 1 't.i .mil „ lu , ,) u 1 . 11, .1 „- ....'■;:■ H to M.b.M.lu.fK ..l^J. ,111- , Jlr , I ,„,i„- -I n u . r ,e ,..i 1W,- ,,.,..-,,,. „m D'/.<< J'-. / iw j. L«m, un, ,| Ll -,h „ 1„m. it tn.l. P! ^Jut, ,)■ I. .•.,,,!„„. „ Review of Distictella (Bignoniaceae) '• mal?m lohnumnme 4 H <*..". U'lTu, v i- «! .miiuiiM I 10 X 10-11 n Lucia Lohmann, who collected the type specin li, l].( 1 me h, i. . < 2ii]/. l,,'m,ji tube, anthers ca. 1 mm I0112 lon^i iilaments >2- 33 ru. i„ , ,Jn , , „,„ , , its utli th mi ml 29 mm, shorter fila ,taminode ca. not at all evi< and seeds with the 4 cm, ca. 1.5 cm diam., ba and forests. s, .11 n ., | ( |, s , I ..„ it! I .If.',.. Ml MM '. .. Hoi. t us li 4 F„^,tnl JW |Wi K. li la parked). 10. Distictei (Kunth) Sandwith, Phenology. Flowering November and December; fruiting July. 39415!). ^ ' : led, drying dull black o 1-1.5 mm. Leaves etiole 25-55 mm, wi densely lepidote; pe i'eX - ems r: , aclaxialh or midrib o immersed, all raised abaxially, ite or elliptic, often 1 •roadly so, 8-24 X adaxial surface mtho ut tneliomes or base of midrib Review of Distictella (Bignoniaceae) with minute appiessed tnc homes, abundanth lepi- "Distils mugnolinefolia Bui , Bi glioma r;ignolui- fields at apex and in th i\ Js .f t! kiln Km- -m IimI H ] J, t' . , 1., So k . l( , I .1, h ! vihtl, abundanth lepidote, i Koto of the type. . : cence a racemose ] ith rachis 12- (1953). ■ ^ .;■■ :•:■ or bioun pubescent vith minute ienugmous, oi xaiious public .. t . .,.s .<,. n.r L<-7 throat, infundibular ,e, curved 1/3 capsule of B. h treated here as D. stamens and stan -10 mm from sterile, are of Both varieties of D. i l.ngeilila- '«"™ ow fllffel mml ^ . ' ^dujoha m haMiig limij ^_ u - _,_ - lh , ^j nl ()l iS 7 k.,s. K ,.. I.. n„ .Uo.lh 'ith "..' ^iV X'lhh ,,!iTw tJ< j i"',^ /' « -o > .1 '« .'diti.i fiunO »i „,.//, ^idoiil. «.,J1 1-2 mm thick. Seeds ca. 12-15 X 12- m "f lM J u! 50 mm, wings poor n seed body, ,v,i.,-!i ditlti- ti in /"/ n the abaxial surface in the axils of the and in the bas . Both varieties of D. racemosa are . D. raeemosa var. from Colombia (\ aui s , ,„l ( 7 luumu. fn., .1 ( A.li,.'., Mitli 1 Jk 1 ( ., . a .! J - «,-, , - \ai tianducida in nas), and Venezuela n non-inundated forest. between 75 and 150(,00) m elevation Figuie IF! 1 >7H in 1 . , „» >,,-.,„/, „', , ( , L t ,/, , ,'v f Andersson lb 3121) pertain to £». magnoliifolia , 2426, 2496; A. ff. ■ '■ ■ .-■■<• ' Phenology. Flowering February, April, May, Jul Septembti, Ncnem' i, ..i.il 1 and December. 107474). elliptic, 5-17 X 3-12 cm, chartaceous, 3 to 6 pairs The specimens cited in the Flora of Panama rvitiating at a 20°- , ,■•:. . ■ abarial surface and the midrib, lateral \eins, and c ral \eins immersed • ■'.'■ ■■■■■■ is stenle excluding paits of the \ahes of thiee old p homes 0.1-0.6 mm, , liml, ...<• I»;.,jdh „bl..nt. .'biiu^iHK ! p.-K.U-. usualJv iwth )iel,k «,l : -,l„i;.l- in ',■.■:■■■■ be unique in the genus Distictella. erect t( > ascending, fer Nov. Regni Veg. 14: 310. 1916. Basiony Bignonia mansoana DC. in A. DC, Prodr. B. Verb, Rev. Hort. 40: 154. 1868, as "m<= soana." TYPE: Brazil. Near Cuiaba, s.d., A Sil oblong, curved, 9.5-16 X 2.6^.7 cm, 1.2-2.5 err at nodes, dnmg dull 'uc n hi.*] o, >n- w*i<.][\ rt ]i- - "Mm« .( i it o-jcii, -u <>| tuse and apiculate chomes erect to ascending; trichomes dull «hite, bro«n, den^ Lite white, ferrugi- |,H 1 1--. nil M'l b . hie tl , ,i 1.U1 h M i ,i, r, s 1, h, !, , , .ml . u- Review of Distictella (Bignoniaceae) 87). been collected in central Brazil (Minas Gera i.l [> r. ,U . I ,11 have been found in Brazil in Bahia, Para, Am with Bolivia in Mad)t A Pi. .rU ne h.lh.,, .sfn,. Mi i,.,l ,p.r... t , ^,U LJ mm st miens lUnb.li 4/, ,., X" \,7-'.",' (Mu.P.iio: .i».l -t ipi.ik. '< i,,-~itf -I ! ,-i'. m" t| ,» k.s, . f m..H,. rces.ra. (BM). shorter filamei ninode 2-3 mm; disc M.ul.M, t«,e,e. .Ir.au, d„f] h,,v n ,.i hi ,,-k. m,|,„, 10-15 x 23-32 mm, wings well d . ornery I-, I. .H 1. << ,1m,s^ r , 1, -, ■ i„ >. ,4. tt. honi- seed body, reddish brown with tips ^eikmish tan. .sually tightly apposed (laieh as, . hdu.,1 ,,,- i 1]( m l , a.ous, halme at least at h ' ,u - ,1, II x ml, no,,,., ,- , .1, su, ,].,,! , nt fontiastmo m color to „ , Him, Im oni- \ . >u ,< h an < ,i |,.it .1 1, -Hi,! •igure 1A. closed network, midrii ly, and October. lo fiat and loosely fpen-reSichie and ihe margin flat. 11,, u a .,.■ .),!), n„! * h„bi.aN I lint, • lit ,...„./>,/ „ similai lo tht leaflets of the ! .bovata. However, the rs, peduncle 1-2 mm wide at base, 1 i.sl.L. fb.'s. ofsi u. I„a. .MiLs,,!,.,,,!-,, Review of Distictella (Bignoniaceae) 13. Distictella o m. New York ca. 2 X 4 mm; pistil ca. 3.7 cm, ovary ca. 4 X 947/A9!, digitized image!, LIS i,o- -cei,, digiliz. .1 -L i>u„t-,k all \t«,- I" 1 ' ii.nt, wl .il.M.Jh .,i,l i ..- .1 ,! nialh, j.LimjI suggested that l> ,,',7,/ff- milt lll !\„„ l( l J. t/. Ii-.us - (J ttu ll'l ' ., ». 1J vary not only in leaflet number and in habit but also ,n.u II ,]],,!„,/'.//„-, 1m„ , iWu.tn iu> j.n.i.ii b u, -.,! , „ t u.h x 1,1, and capsule coloi (blown m D »i»of' 7,-1 ii< lands scattered in basal area areas in Colombia (Sandwith, ca. Distictella cu s without trichomes drying black, pubes, i „t Mill nullum, , U- lli. -, ,i < ,<*> i ,. ,i 4i.ni !l n I, nam i- ,1 n. lb ik, !, ... hl-ts 1„mpJ . rf l -il.iinf - i ..Ks.'-lO n,l i ^n, ill, tl-t ll.l.oil and nil n 1,1) I i i.. 1,1 k, In, It, , , - 11 , 1, t n, 1! s ap, ,f the kail, ts ,f Dntu'tlla ihovata and from base, (4-)6.5-7 cm, chartaceous, tube (3.5-) I, hese specimens are i„ l,n .l| ( ., Uacieol.-s 2- > - Luis Raul Vasquez [cabaru, /. A. ca. 1 mm, losl calyx 10-13 X 10- ith minute, ferrugi- "" n0US ° r w! ' l6S and lepid0te ' glandular fie] i white with yellow 14. DisticteUa parkeri (DC.) Sprague & Sandwith, 1/7 distance fioin I. >- f ', 3- ' r tl i s„I,< f i , , ,»us tnl.< ■ lit-', v em ,.. t I. -Imnif.l. I i- . 2,1,11,, ,[ .nn-io.1. ". }-~ _ ...m -..,11, l.<.llon, ,.,,,,,1,-J wT |, m , a h, ,„. u ., „.,[,!„ , , , glands overall or glands numerous but !,..m, !,l,-K 1 lie, ..w,^. oh,,ai..,.i -ll.pn. -, bW„ .11 J- 1 , m. il i. k N . .', 1 -20 i -0_ n „■ «i ,g- ,orlydemaicatedf..mi s ..I I ^ >e\,U Ulu ui, <,,Hi is hghtei bio«n, membianous lnalme. the ^em, ■ i. Vui./i^Li. -iid m,,,„ Cia/J It i, mdi-'.ulwdto t men described as veins immersed ad lateral veins on white sane ween sea level and raised abaxially, tertiary leins flat to immersed, 500 m. Figure IB. Review of Distictella (Bignoniaceae) .iis,-) .,,!»»!, c1 Diitichs giiianensis (Bmcau , Discussion. Distictella pauciflora is most similar -i.pi.iJK ,-l.u. ll.».rj-l -,11ns m h.,,,,,* l,.ei..l.i ', ,.-,.,, t.,1 u l,iM ,,i,]l. ,lhb ,n(,i I one specimen of I described as slightly t iche 947/A11!, digitiz, 10 — =d, densely pubescent nilii 3S , with few scattered glands, midrib inconspic- i-,,,1, n II ,i n^j.1 nil ■ 2 - } ' ' ' /'(''' 1 l.,.,a Id, r- i ii .u.l - --.0 i uu v 1H1 , . 1, , u> el ' " 101 "- ' ( ,l f " l)t " ,1 " 1 ' "" ,u ! ' late, 4-12.5 X 2-8. lS ing May . Review of Distictella (Bignoniaceae) or apiculate. Inflorescence a racemose M ,nrl, ... in H ^JW-o,,.,, ,.,1,1 ,|, 1.,-v,. I. ,U ° ' 'W . '' ' '' ' l!i * ' ' ' ' \ '' \x\ I l'-l 2 lii , f Imp, m- hmIi, I ,li n „it! \i>, „,„« ■■■ ■... . • , .. • . - ■ .■■..,.- without t.ii J <>! Seeds 15-28 X 24-85 mm, wi: seen, and the she 880400 was trcrn annotated In Schui fore selected thos here as the lectoty t barcoded BR Unf 880502 was apparei by Bureau or are Schumann and is bet -osyntype. The indi Martins collection (b also does not tain Gentry (1973, 19' d Burger and sepaiate species, emphasizing the elated teitian . 1. l,k Hi. 1. flets .1 ' , /< ,' ( ,n , n jm, i<>" i »'• *> i 1- distinct from those of D. race magnoliifolia is further separated from D. r tened). Leaflf in ilus ,,u.. .llo ,a*m. ; ... ,. ■ ....,, /-,. ■ , ..1.,., I..,, ,n. ii.„„, - «„ ,h a^ls ,, ,h, ^ LJ -'■' l ''"" '^nolutohco • In,, i-e- Review of Distictella (Bignoniaceae) Grosso), Peru (Loielo,. ( <.|„ijil).>i ,< ,i.|„ t, V t .|«p«, I., lh< I , tl. U vl.i. h ai. ..tt n spatuUe and lack rasMz'] and La Paz). It is a] 6-6.7 X 2.5-2.7 cm) than those of £>. „|.^ ji..' n, ilir- Im t ,l U,,.,l win avik ,.,< lidrib (ca. 45°). 27 -']'"'.. 'i?.'rt 1> '"'•!, m.I . '.Wml"™ i,lil " 1%,rl K " 1 F| ''' ""-•'''""- i vl "> rli:l l««-i«= tli. d. uning isobpe (RB) and illustnti. Urb. var. translucida A. Pool, var. nov. TYPE: Colombia. Antioquia: autopista Medellm-Bo- gota, vereda La Josefina, camino hacia El Pital, Mpio. de San Luis, 800 m, 29 Nov. 1983 (fr.), 5. 2» ,./ .,>.>!,„ i, h,, ,1„ ,„, ,,. ,.K , ,m,.] kh .'0-1- „_d, ih tl. -nul . However, 1 Ids 2 'or 2.5 pairs. ..- i h Pelt.. Am loorly to well del, eaflet bases. • mi. I U 1 k' mined in the iresent -4tid ' ' ' ,,„.;, , .,1 Mm ,|„,,.- I,*, I. I!,, I, ,,,!,, Si i - l> t 1. 1 ) % h ! n 1 ' , id flu I I I J- t 1 Ji i . u 1 . . i [l«iu-S ol Mutm .If not l..-en ^sociated mth D. cremerui; it is povudf ^^ /(j .,,. | MU| Slm . p,,,,,,..^, .,„ l .. u „. „,„.„. ,_, tli .1 -mil' .-i th. II... *.i.i<- m.,[eiial .it. d W . , \ m |,„. ^, „| Leiu, de Humo,/. A Review of Distictella (Bignoniaceae) J) \. o I'HC I) h:h ^> s *: ,„>< ..i«!< .il ,>.tll ,. I ) i.m 'hi k Sctds ra ]5 X /. C/iugus J6iW (holonpe, INPA not seen; 35-50 mm, wii ted from seed 2 mm. Leaves bifol eleNation «as not indicated on the labels; most lepidote, without glands, abaxial surface with a v ,rders of Nenation. Imll «,tliil\,jll lul m j. i. i n I ml i . t U diffeis from D. base rounded, apex diets, which have cuneate to i-1 i, I. lai till i- .' j ins , (i II . rt 'nt. mfundibulai i!V "mil mf" mil]' 10 ^ 1 ' 7ut ' ' '-" ''' " ''' l " ? "~ '"''' 11 m ^lil'.to. Ill"" ure Cited 4 X 2 mm Bailloi i, H. E. 1891 [1888]. Histo ire des Plantes. Big ,sule elliptic, b -- i, Vol. 10: 1-78. L. Hachet. e. Pp. 1026-1053 tham & J. D. Hooker (ed 2(2). Lovell Reeve & C ipressed or sometimes one Lon :andolle, A. P. de. 184 L 42-248 in A. 127-153. 1-20. ' DFID, Man . 1978a. Bignoniaeeae. /„ B. Magnire (editor,. The Sampai... A. .1. de. 1435. Novas especies de Bign Gi r.,,11., MijililancU — Pari X. Mem. New Ann. Aead. Bia.-il. Sd. 7: 111-127. 25(1 > 1-H<* . 1982. Bigi Gomes, J. C, Jr. pi. l^t. Review of Distictella (Bignoniaceae) 22: 403^20. (17a). Aparecido ..-. ■•.'•-■ ■■..' Kingdom. Sclmltes, R. E. 19,0 Dp planti- tovi nu- . 'i.n.lc v, < (V «*//„/> > I • -> j ,. 'i ,, (1 i>) Schumann. K. 1894 [1894-1895]. Bignoniacpap. Pp. 189- (11), 10132 (17a). Berry, P. E. 1382 (10), 7426 (10), 1426-a P. E. et al. 6026 30: t. 2933. 1932: 81-93. ir.ll. ii i i ilj h P r ^| i n \ < . , \ I ( I I ( .. Hort. 40: 152-154. r '' .''-I 9 , < ' .»' r l 'J. 'll , V' LIST OF S.PECIES j chocoemis A. H. Gpntry t -,'./.. -vurh,.,'! - D,- „,<■!' j eforagata (Vahl) Urb. L". .;,,,/•/' \. H. Gentiy 7a ■■ «»> •• J race™«a (Bureau & K. Schum.) Ur i racemosa var. tramlucida A. Pool ic 9, , /,' (17b), 76930 (17b). Gillespie, L. J. 902 (14). Gottsberger, 1. S. & R. C Uevdon^a 172 (ID r < • /' ^ ' s„ 'sPFl-,220, ill). , , sr-F'71," II, ^,m»^ -' /■„„., / ' „,/ //'«.o,17,,, ' ', ) I , (ITh, Pues. M J P ei (17a). ■,■. a *L Andersson 1194, (17:. ■. // eft, 6678 (11), 70659 Review of Distictella (Bignoniaceae) A REVISION OF MALAGASY Zachary s. Ro gi GNIDIA (THYMELAEACEAE, THYMELAEOIDEAE) 1 Gnidia L. (ca. 140 ] U , s H ,1) , 2«M [V- u j Ul? 1 ear lit- as in tast as Arabia, India, laeoideae, and is almost completely restricted to h significant regional .^,-l/sll. (-1.1 ,«■ . I,.... I. lu «! j- tVu wn.v ■ MMu.iln! \| Pie. ... ,., 1 P i, v ,lli t ,„ . ,umJ I'.'l. J'«*n 720CK genus was la,t deal, < I. > Mua ,-, „ lv Imidn > /. ,„ ,1947) Most segirgates weie i. Iiu.d to ■-.. :■■■■■■ that define the groups (see also Peterson, 1959; f] the first half of the -LUi.'^ ili.il lh« (.rim. i, pau if al classification (Domke, ]«;n 1 m.ln il"".'. m plrdetic, but additional genes and much broader tl mores was probably molecular data. species of Gr scar (neither genus I ml.,, i f, date, placed aigutd stiongh to suiommize hi w n „ «ii>, ^ - : -. : . ■ ■ . ■ (Heads, 1990), A dri, and two molecular ph Bank et al. (2002). is teimimlm £>.«;«,,, [\n' , I ", ,H, 0! I" ,K s u H> s on, -auk., nl , , L M 1. ^ shghth exseited stamens, the sepals notir.-ibh ,..„,- Ihe \l„rjii s.*, ,^ D cotimfulia L., was Lasiosiphon (1838), Arthrosolen C. A. Mey. (1843), Plnl.ih , Is ■.-,,- I «, nik, 19 Ml i L f, ,'.,.' w < iU |L. 'I ( ,<>,,•,„, ,j';)l, substantial moiphological diffeie.i- can still be separated < Le could be observed and sampl. androecial characters ■nbach & Van der B k et al, 200: publ. data), tl ie The s ; in the northwestern and far west sland were unable to be reach How those t^nZfZs and ,d Xt, ) a ecent field observation xonomy of the Malaga! rf elevatior s- the Gaz *zr t rztjl"7.z.!Z'' ^--'oStZere 2 is Localitie. is data inc: i (Schatz & Lescot, 2009). Full '■'//-iM ^ill.ldl JO!*., I. cji,-, j.,,1, , ,i,f om ^ ys\)' allv SeCtl °" ^ Pr ° V1 6 m PPe " K (Leandri) Z. S. Roge, ' TERS AND VaRIATI0N IN bed. habit of a few species (e Materials and Methods MA, MO, NY, P, TAN, TEF, US, and •iffrutescens Leandri Specimens and pickled material were colk s, at 1 m tall, have a single h .s 11 1 llul l(j 1 -| 1 uli 1 M,.l[Hi,> l 1 »i -b ,. Wln-m w, UM. ill 1 n , The same blanching pattern occms in t«o closeh between lea\ ranch (e.g., Gnidia and described in d< »66c). bracts), inch, species, G. dan- : - !..-■' „i. I ,,l,\]].t,i^ l.m<:,u ;^.ir„rrh- ,., ( U,,.',, i.^ % , i .. .M-ini. ./raruHHi Leandri (Fig. 5A) and G. neglecta v; the length to width (Fig. 11 A) differ markedh In their opposite to (1™) ratio of are taxonomicalh ■■: : ,: .-..; ■ • uous leaf scars. 15:1 in G. gnidioides (Fig. 7A) and G. linearis d cences (Fig. 5A). For most species, leaves are Z. S. Rogers. of G - neglecta and G. perrieri are often glai least after drying. Leaves are shortly peti ^ases, howe\er, the trichomes are faint and impel- humbertii (Fig. 9A), and on the abaxial leaf surface of ,ericeous indument remains so dense on both leaf humbertii are generally matted and ca. 0.3 mm long, ambomlrombensis (Fig. 1A) and G whereas those of G. ambomhombeiuis, G. ra:akama- lalana, and G. tlaphnifolia are straight and n length. famih characteristic, but Leaf bases are usualh cuneate to attenuate and somewhat revolute, the lat lecurrent along the petiole, but the bases of Gnidia more frequently on wider leaves and especially nea kmgmana (Fig. 3 ... 11B) are at the base of the blade. east shghth coidate. ^ ithin a species, leaf apices Most MaL i exhibit some forr an from obtuse to acute and the tips are usualh of brochidodromous \enation. The \enation patter consists of one, or sometimes two, submarginal loops UNtLORfcsCLNCfcs interconnected with ; ; twork near the margin (e.g., G. danguyana, Fig. 3A), or of strongly Species wit ate phyllotaxy (e.g., arcuate secondaries meeting the margin in the upper GniJia Jecanana) consistently ha\e temiinal inflo- 1/4 to 1/2 of the bli olia, Fig. 4B), lescences, wheieas species with spnal plnllotaM or by a pattern only. e rune axillan or pseudotemiinal inflorescences, the by a midrib and two or four longitudinal plicate veins latter condition being the most pronounced «hen ;n visible. lid), the inflo xtra-axillary with the duncle being illimeters above its irt G, TAN), and Humbert 12844 (part H, TAN). There is substantial \ariation in the inflorescence found on the leaves. Leandri (1931a, b, 1947, 1950) structure of Malagasy Gnidia. Nine of the 14 species considered peduncle length to be an important inflorescences (e.g., G. linearis, Fig. 10A, C). The possessing involucrate inflorescences. His observa- Jangmana, Fig. 3A); (5) mam -flowered, bracteate, be classified according to the length of their composite-like he . 7A). peduncles as follows: (1) completer* absent (e.g., G. Peduncles are usually erect and sparseh pubescent ambonJrombensis), or nearly so (e.g., G. gnidioides); with the same kin:: iment as that (2) short, no more than 3 mm long (e.g., G. hibber- !«.],., -, -n ^ aie Js. a- inn,. II. « .. .-)>< I,...,] „ l.|. ntk-K ^u».,l 1 ered and usualh consist of fhe, or rareh sk, (e.g., G. daphmfo- imbricate bracts, but the imolucres of on specimen labels. leaves in shape an, in texture and Flowers are sessile to short pedicellate. The ■■ ..:■ , . ■• ■ : . :.■•.. :■ -t the' inneimost bracts will be i\ mm 1, , li tl (F . J Vl ihtt lie not ahu^s • ■■> ' I .11 .i t^ mut- ha>t iallcix oil Tl.. 1 ,11-i U-,, >,i\- |J.,« U in.l Jk.Uj line,/, l,i a ur,,,! it i.M. • n 1 .i ..li «- .1 >lil- tl, , ill. l.v tht hu v 11 be des^ibtd as the ■■■ ■ ■ ■ ■ ■ ■ as the fruit de with both articulated fL0WERS and unarticuh it remains inside the ■>■■■...■. d out (Fig. 11C). For species with inte: ice of the hypanthia in G. ambondrombensis and inguyana is the most obviously pubescent of any ies in Madagascar, but the indument itself usually omes are sometimes impossible to see when appressed or sub: g., G. hum- (see review in Heinig, 1951). Petaloid scales are desiccation) and soLudieastem p. puhtioiisf II mt * .lii i hi in 'i . the 1.5-2.5:1 range. for any species. When fresh, the scales of some m the abaxial surface is ridge becomes darker after drying. earned faithei In I 1 , v > id <)\o .1- ,.< 1 <,U M.i.-ii In Uv M 1 c ., s ,„ Jt . th Beaumont et al. d that long the lower who: rved as being about ^ -... .,.;.■ ■ ig, ca. 1.5-3 X 1.5-3 mm, treatment. The eshy, and usually ca. hmts laigei than the petaloid scales 0.1-0.3 mm tall. I , r , ., h« s „ 7 ,m de\elops), a lateial -hie, and a capitate ,\, V nj Hi II nu n< % \w b ,. L •> - f en species the ape oillie !,\ „ < meied l,w subhuinid monMn. 1i, -. In, ,d) as Jim us,ed in (eg, Giuha danga, 1,1 Fi- M I.. «.nti ■. « I t. « si , - ,s„ tl. istent in fruit. Clearly, some species are restricted to higher FRUITS Car ' FWe SP6C ■■ . . SEEDS diverse substr I, laterite, limestone, : essentialh matching H i. lot the h ml 1 l.e -eed , „ ,t ,- ,stM> ipl l .j th« ,,ln.].l. >..i,.s .„ cmstaceous, thin, and can be either black or dark a on label data and ire minute. G. razakamala soil on granite rock. :>■> pheuoKt mirn M.la^asN species, by IUCN (2001) criteria. Gnidia negleau and C Endangered (I nana and G. perrieri a Gnidia L., Sp. PL 1: 358. 1753. TYPE: Gnidia ■'havoha"), a Malagasy ous bark of 1M.I .») a ,mnU J.f , lu.u.lKial. Had. and s. Id t. tourists (M. Made! 3 . Tlrunelaeaceae aie kno«n In ha\oa oi one of pis dioica Capuron (Rogers, 2005), three species crata Baker. 4226). The leaves of :e cal « lobes 4 scales usually s cal (in Madagascar) to Mrict rjulzTi bensis Boiteau, Bull. Trimestriel Acad. Malgache, V ,' >.l, i.t 1 1 i , U, . , i.| . , i Ui Mil .!.. iu .( t .nil n i, i i ■ ■ . compared to adjacent leaves), 8.3-9 X 3.3-3.8 mm, are still selectheh cut down for local use. This i I ,,tt, ,1.1.1 w,i ■'„] ... *. .< iEN .- '.1,'iiu.oIKNOO.. half abaxially; i the species is kno«n from a single unprotected ^-4 mm, j eave ^ thdt aje rmele j , m 1)((th Mutaces j,, , U u— anther, oblong, 1.2-1.5 X 0.4-0. mm. subses.ile; anr) lts nmoh sll) !ji M r2 -'l 5 , 1.7-2.5 mm) calyx v . ipitLl. n„.. (• Ml inn. (all I .- brous, fleshy, apex i inuses shallow I tin t -d 1 mm hi h |j J I .( til ii pit \\ "* l u'l |'>1 l'V •>> Jilg. Nat. Pflanzen- bojerianus Decne., Vo ui' , '. ,k U n L (*-n - n ,U n . .1 1> !"'" jo, T^iTTm'' k'T[2]n population gro». on scattered patches of soil on a large ^ «/>« % ' «"« Ba.ll ^H.st Pin- M ,Ja t as, a, • ,, i [ ul collected in Shmbs to L£ ray-brown, usually category. Gnidia bojenana is seKj.ub s .-t „n(,!uo tl H ..Ms 5 , ..I.s.imI^.U .,1h-m. ,p. ..-,l t ,l in .. ■ , : tiirl.(.i.i.s CJ-Oln.m. h.M! Discussion. Gnidia t lobes 5, spreading, obovate, 1.9- t!,. '.,[>> 14, , I, n' licl, >!* mvMi- , ' «< ^i. | U dging from ; . ■ ■ or subsessile; subg.noecial disk cupul lost on dried material, and that the < , ,<> ' M . « .t BM and K ait regarded as isolectotypes. . ■ . ■ •rl i.l- h-e.l ^..it it- Mh 11 jn. i.l « I, 0.7-1 X 0.4-0.C ipe ca. 0.1 mm), "^ [Pierre] 8874. gascar 5871); havoa ■I 118, 197). midrib depressed adas :urrence (E00) of the lighter green than bl km 2 and the A00 is 1600 km 2 IMithi.uii 7 r -<) l...h. ..it.^.l.t. r ^ ir,1, "" J densely pubescent a s later in the Flore tie Madagascar et ties Comores pubescent, apex rounded; petaloid -cale- absent; (Leandri, 1950). One sheet of each sjntjpe is ,ndition than Louvel 197 (P). 4 km NE of Mahabo village, Rogers et al. 76 (B, BOL, K, L, Mada»as< ai it ,. ,1 J-. ' .ill. i ... ' i ... !■ ' . Nat, ser. 2, 1: 436. MO, P, TEF). Amkivcmbe ( "daphnaefoi ifolia L. f. var. t 19 S glabra L. f., Supp] Dessenia ^ : " daphnifolia (L. f.) Rat, Fl. Tellur. 4: 106. 1838, J(1 Rogers & Spencer. W-SM!). lius :, P 00380375!; : .-, • L, .i.lu Bull Ml, "Ul H.,l f J .t 2, 3: 151. 1931. TY1 P!l idtii Scott-Elliot, J. Linr tl-e f, i \ ii i h I ill In pub ,..-,11 Inn, ' etheadaxial an- plateau link the raised and or ally found on rocky >!]• ,>v,it~ ' .h M i i .1 2 Int. i- -< li- 1 im.liiuka (Cloiscl 115); lu\oa (Run- n-i »n ell.v ,». m, . .., , duds (0 2-)0S- .■.,.' » ■■■' 0.1-0 4 ram till h . li In i i tl i Jh' ,ui, llq.-ui.l ... 1 ^ OOmm, sparse tnehomes near apex. '' M, - lll,M,f " ,t ' 't-.lmtl.. hr i.tm. . i-.,...]ii. -,[211', /_•/,/_',, , ip\ , ;«/*i,pi 7Li_\liM (, h ■■ ■■: ■/tf.snandi ? *^n» iespi.in.-K ,| , ni | n i.»r, hwlhlt [,,-', , Malagas. ftn./Va, inc. s the lectotype. Strangely, Leandri di, phnifolia L. f. ^ W ' in his second revi his yoS Whatever the case, L< ,,-n n . Wo ^illa S . L>x-li . r/rd ■ '.».' ,M<) ./ ] A\ If i-lM.mluu ,. > |Um,i , 1.4-1.9 X 1.4-1.9 ir .-•.'•... pTc 1.1 X 0.4-0.5 mm. senile" or ten shorth stipit Nat., ser. 2, 1: 436. Madagascar. Toliara: Fort-Dauphin, 3 July 1926, R. Decary 4332 (holotype, P 00373426!; isotypes, P!, TAN!). Figure 5. ith 1-: «>id-ellipsoid or ellipsoi . X 1.3- .: .11- ■ > -*H~~ astern Madagascar near F from sea level to 950 mek 1 7, Th.- a .: be : rdc ■posite to decussa, 3 shoots, appressec p : a :ular, (4.1-)8-15(- 1-2.5:1, both su i.i»i-i,'iil,„«liwt 1 ! 5-11 n ilaxiall status of Near Threatened (NT). Gnidia subcordata Meisn., an African species in the bcordata (Meisn.) Gilg), differs from G. decaryana by ■,.'■■ ,l,s .1 i.de low, i ,s, ,!K U ..II t ,',.. , 10 -l-J urn ■ llU h kJ ,i hst.i.ru.nLU. m1 ll'.nll.. I it ni < nl u. II. J « 1 1» Ih -I !<•,.,. In.r Is) ben n.e.nl.i ,ii..i w | )td , i. ,-k s,t!.'r>i.!- , ..rh rJ.-w , i,i ' " -paiseh pubemlous .daxially, .h..-it.d .1 P I;,iL -! -t- .(.->] ■„-.,!, ih bir[] ^ ~ ,r, lt p '„, -,,'nh >' F(, ^~ - ( i.,t)« i i.H 1 i.., 1 .j.,.i M )llll|t!llll . u , H-r.ui.i , , . 'I, , , ejt UltlU {(., P [2] TAN) slij.il> J, I,.] o..r lii^fid 1 spaced trichomes at the apex. 6. Gnidia gilbertae Diake, Bull. Mens. Soc. Linn. Distribution and habitat. Gnitlia gilbertae is Paris 2: 1218 ir« llir- ,!.-,e acute or s - 0.1 n ' G: d • fri 27 (111): 46. 1934. Basionym: Dais gi Baker, J. Linn. Soc, Bot. 20 feaferi Gilg, Nat Frl.vi/t-i [1930]. TYPE: Madagascar. Antan sh, usually dark red afte • ratios (4.5-)6-15:l, ii J;;'; [Ruzafindrakato] 3055, Reserves \aturelles [Rah* [R lo- to WO- ^ 8. Gnidia hibbertioideh (S. Mooie) Z. S. Kogeis, S. Moore, J. Bot. 58: 189. 1920. TYPE: Madagascar, s.d., /. Thompson & J. Forbes s.n. Ml). Figure 8. within the head an< us and thick, leaves. Examined si o Shrub?; young branches densely pubescent Rolnns (19-5) m th. Fl , / ir „ ■ I i t u Ir and Ih's „uj I lh s >h< . . Ji^btl noted that the thiee s|, n-, „< Is!, In, I 'ml ,uiJi p,,(. i<- I < it) , a dense uniform indument of ca. trichomes; hypanthium 1.7-1.9 cm, articulate; cadu- Later Domke (19; lion for the trichomes near articulation internally; persistenl the lectotype to y u PP er half ' with an irregularly lacerate or sinuate Phenology. The type collection is in flower, but covered with prominent leaf scar denseh pubescent, ± exfoliating or in the protologue (Moore, 1920). spnalh ananged, closeh spaced , rarely h - IUCN Red List category. Gnidia hibbertioides was nodes to 1.5 mm, persistent oi ily at t he tips c 1800s and is lepiesented In a single 1i.l1m.jiui ■htly obovate,' or line 3 mm, l:w ratios ca. 4-8:1, silver-^ s^5) a tr ta ^zr^ ej,„x iF\ ot L\\) 1) - ..,-. . h,'s.,^ been conducted. abaxialh; \enation obscured . Inflorescences terminal, erect Discussion. Gnidia hibbertioides is distinguished to 1.5 mm, densely nvolucra urn (vs. rarely to 1.5 cm Ion- broadly ovate, 6-0 X 2-4 mm, l:w rati as ca. 2:1 erect, persistent, densely pubescei pubescent on upper half abaxialh. e. cik-HIs >!,., ,. ■. Lit Mi- 1,^,., fl, rtll 'lis K.I.J1.1H-IM1K -«* .IJI.Illl. Jll »> iiu.sl „i'm ,,. t n,s ,i r.l hi,^ (, /„ ,„ |H)ll „ ol , I,,,,, FI. ; feature of the Scales is the id ! 5(-l) mr : are rounded, emarginate, or lobed at the apex. 1 i . ll 11 I 1 I | external] Typification. Gnidia hibbertioides was based on a trichomes 0.5-1 mm, glabrous ir •' "■:■:. i.Mi'kutti .1 l.^-l', itru m lU " UI " |J «i<"^'^ [u bt scent abaxialh, apex emai '•'■;'■•■ V ! hillh.,1,,, ll -.x, ) j. hill, ill (P II JO"" i° I slid J. '.1 . h...l t ll.Uul.l. • : .• ■■ not arrive until 18i L ca. 1.4 X 0.5 mm, stipitate pubescent, ap, lubescent, trichomes to 1 mm; sty! rous or with a few Bull. Soc. Bot. France 76: 1030. 1020 [1030]. 1000m, 19 '),t 19J' ,/ If ml - " .,!.,. 1 !i ll. il i.l lu . |„ i,ti .« m - ,„il„ ., height of lower who) Luji.lliMs 1 -^ jccur from the Mai Isalo to Cap recorded in tw d areas (Cap Sainte 5ainte Marie and as far west as La Table (a mesa-like Marie, Isalo). Se\eral large populations, olnioush calcareous limestone (Rogers, pers. obs.). preliminary IU f Least Concern (LC). Phenology. The spec Discussion. Gnidia s the only Malagasy pedicels ca. 0.5 mm, densely (K, P); Mangorovato, , trichomes 3-1 mm; hypanthium 7.1- 10. Gnidialineari irs, comb. nov. denseh pube glabrous internally; \ ,• rilllM ""< ]')](» l')_") |l w ll] '„ . 7,./;.' llRl.l'l- ,.|-M. til ,« [I n '-'"mi TW M.-uua. i Jv.li..,/lK,iuM.t,.. ......mum -i i^. -.i.-.din., 4.1,. .u- .d AialU ,!, Vll „„,', ,,,, „ ,,,,,. 'I jlalM,,,,!,'^- s.mm.sj., U ,J *■,»!■<. lot />,.„, -' ilr.t.h|, I « lt .|„, P' ,.„ M „. ..itl.e,-,., 1/2 to 3/4 exserted, lower whorl 0.5-1 mm G!, TAN!). bel, ic, 0.8-1 X ca. 'I", J,'/T„\\ L !! U „ ! ) ! ! !!'„'" a.,' C f'l-> i,n, t,ll. .Ln.., L , .1 L mm. Fruits ellipsoid, I 1 Mi! 1>1 I- 1 11 -J mil nil ••inn-N .<,»M«t...l .flnnatir zone of ki:, .lism'.iiiK.ul. ! Dtlui ]...;. ill ui. ii y pa a SE to SW r ° Und ' elles [Rakotoniania] 2777; Reserves Naturelles 5 here, and if the decaryi Discussion. Leandri (1930a) used two collections L^osiphon linearis and L. decan decani \ar. erectus. Later, Leandri (19 .on at the species lank as L. ,<> 10 iImwi- Ihp.ti nrin!.., - m L . ti.iml, rarial iniloiescences, and ] i breaks down iuoI . t. ...i < th-i n.jLlm,., .1 I , si . ■ . ..■■■■■..■ : . -,,■. . paper factories ope ere cited in the protologue of central plateau (e n decani \ar. erectus (Leandri, 1930a): narantsoa, Ambala\ Waterlot s.n. (May II. tois „ I „„!,,. int.. ft .... l'u .... ,i ii .Id }„>. , Li,,, it . >,,,i slL. mdii 1947): A. to better understand the observed variation. Grandidier s.n. (Nov. 1868-Jan. 1869) (P), Humbert name L. decani, along with two othei \aneties (L. ( AG), Perrier de la size and shape (eg I ,.!„<,' ,11,,. ,n jn t u I , t < ' P, , ' , ,1> Ins li , t K xarieh littoralis; lea\es longer, but not broader in and leaves i he description and 'in; in, renal im. flore ■: ba ; |2j k. mu. i '-.\ upper who . 0.7-0.8 X ca. .es; style ca. 2.7 mm, glabrous; town, Sogers et al. 911 (BM, BR, CAS, G, GH, K, MO, P, stigma ca 2 7 aim I.. Jo I..,.. iliml TAN, US, WAG); Tsin a, Fruits not seen. .... ,■,":•"■... MO J* PRE T \ T.l ■ \ ■ i,"t»-l,,„ l'V) I l " H ' 1 '! "1 '"' ^ ^'" ' '. '.'' "^ (a B«*^i« 8553 (P). /{/GV figrf .^ „ eg/ff f(( was time since it was h ite, persistent on ol be less than 10 km al petioles ca. l(-2) mm, spa />,...,....,..,, \,- : ., 1,111, I,. il„, ,„•» I., ,.,, " Ils stl ' ' ! ° ," lL '"",'"'!"' V"i"t'"i ! " ""' "* ^''" N l1l ~" 1 l]t ° ,|lx li - ,t,t -' i.'' , 1 , ...;., , ;.. , ;.. , .: ,: :::. , ."i '■.'.■ ..'l"'."...'-'.'." :;^.i^../;:..": 1 , :. , . , .. , ':. , ..' , .;; , - I;!;;:!:;: 1 :;:;;: 1 ;;;;;';;,;!::;; 1 ; 1 ;:;:;; 1 ::;;; .'-f Jiuuitc'l t- IiishW, , 1a .ui and rV l-1 ii .,1- - >h -pi. i - ni .r i -en. hit - rlit ue-> -tic i Gmifia raeg/ecta superficially resembles the soutl east African specie? letimes treate hypanthium. Etymology. The Jabrous; style 4- been originally misi isly as Gnidia decaryana. 19 r m- a , 1 a riw No election data ,ii<- t. uU.leo. *p.<)m-,>. k.^.lt,,, i.UNlvO! /;/ .,..,„ > r.-/-- ™> lyP- .iK.1.11 ,.,!.! .....L.n,,.. I.i.i.l ■, U >vPi_ 1. mi. , ! .■ ,i!, ^aiseh lenticel- edg, s or in open sunlit patches within it e: 11. vitl. ;, 1. 1.2-1 ■1 X 0.1-( 3 12. Gnidia occidet Phenology. The species flowers and fruits in May, Discussion. At the time of the original description, October, and November. Leandri (1947: collections of Gnidia ocudentahs, «luch «eie infoimalh categonzed as riging to the normal form (Periier de la Bathie 1276), or "probablement des formes de la IUCN Red List category. Gnidia occidentalis has m ier de la Bathie 998, This species is pioMsionalh assigned a c onsen ation descnbed in the protologue as 5-merous. Strangeh, status of Least Concern (LC) based on IUCN (2001) Leandri (194; tice that two of his criteria. syntypes (Decary 8181 and Perrier de la Bathie 998) and bracts closeh matched those of the similarh G [2]!, I ). Figure 13. t thi _T< , ,<| ft, e, de la Rathe Hii9, diffei „ . t ' . . n , , , 4:1 l:w ratios), and its 12.5-16 mm (vs. 6.5-12R5] 2 ,anthla - p vate, 4.9-10.8 X 2- "" -"''"' ( """ ' l ibmnlMiM, .,.-,) ill am . ifloi k, !, ■vith 2(4) \eins di\erging bracts with sparser indument. longitudinally ir base. Flow. L-Gnidiaoccidenlalis Kegel was never validated late, In eM.n.ih, .11,. s,,„i . , ,.l, ,ent m Wi half As .id... ,1 h t ml . (1'4- 1 r.), ..iM.l ' "- v,l "» t 1 . , i.in.s.i -05(-07lmm c-ilvx Perrier de la Bathie 1276 (P) and 8549 (P). The sheet t0 suborlnculai. ',-») (> ■ ( l.,*i • lu.,1 . t ...11. .w, t'l to V Reserve' #8)Tsoakla! rL™"L Naturelle^Rakotovao^m ellipsoid, 1.1-1.5 X 0.5-0.6 mm, sessile, glabrous, (P). sometimes wit] a 0.3 mm near apex; Basion m- L , ■ u>h ■< no- T.ailu ""' ■ !,u ' fl1 " ' ' Notul. Syst. (Paris) TYPE: L5 mm , Serous. l'^ 4 H.,i.iil,„ i i u.Nitm... 2.100- ,|..,k ftl,. \n,l.i.i t jt.., m.-.i f , ., J». liogci^. \ wag mlgtakemy species Phalena octandra (L.) Baill. attributed to M aeus (1782: 225) and Dais rhamnifolia Baill., Hist. Phys. Madagascar 35(5) « a " apparently use d as the original material in the [Atlas 3], pi. 318. 1895. TYPE: "Mad > n oi G - capitata L. f., an African species Sci. Nat. Bot., ser. 2, 20: 51. 1843]. G. capitata in Rogers and Spencer (2006). Gnidia da, engnned «ith "Madagascar" and the species name ^"~y e ^defth^tpJ^T^ & ' 1 „ (; j V 9 i ^ Baker, J. G. 1883. Conti aumont, A. J., T. J. Edwards & F. R. Smith. 2001a. Leaf 1 L. & J. B. P. Beyers given on page 255 of ■ of Lamarck's V Ennclopedie Methot parently, La- Don marck never actually published the Dais name, and u Decne. was treated ies Engler, H. G. A. 1921. Th> In Meisner (1857: 599) and Leandri (1930a: 1042; H. G Engler ters to determine the affinity of this taxon within the Gastaldo, P. 1969. Adumbratio Florae Thymelaeaceae. 343. 1890, as "Lasiosiphon* rhamnif alius." TYPE: Madagascar. Vonizongo distr., s.d., Baron 1895. Histoire Physi, i. Histoire Physique. Nat, I[,i l\ -001 ill' M , ',,!,„ '<,',, 1 i ' VI!') [9,. 5097 (10); 5170 (10); 5294 (10): 0); 987 (10); 1064 (5); 1214 (6); 24575 54 (7); 5770 (4); 6191 (4). Basse s.n. [18 May- Louvel 118 (3); 197 (3) L,. k .• 4105 ,i 422f, .4, ., |" |ul\ ]9»lliloi -p [II |,, f l"}l]ll()i •!.',', .1 < 9n~ Ml' Manjikahen 171 (10). McPherson ,,,,-, l.Til DuUisl.o, i'-= 1 l<* 4 (10, IWlilil M.Wlu.,- IV'llO' \I.-m-1 i' ; 1945Y (Y). L i.|- i , , C < i I h'" 4 I -el ,"< ,101 851] (7) 85411ns (10) 8542 (7) IS. ii -I h im. ..mm s ,i .1) - i, Ui -, , I, -, , , |>, 110 !])'». l.l. J.'.liM, ',H",Ui 517', i,' 5275(1.1. .K", IOi ^514 (10). 31ol7(l i-l 10 1 1(». .T->-t. .!<>. 2 4643 [=Boiteau 4643] (1) 5051 121 1 t . . I t.L 1 I 10, illi .25 ( lili -2>> 1 1.» 5' lh,)5l t .kl, • S . 1 1'., .."- . I, : ( [Till. i..iii3Jf.9(Ji ,P, i.i IT..,, ilr lt,-il i>r -hi mi. mi l<» i'. I -. I...I7 I.Jl ,10, Niii„„li4l'iJi!, in.. . ,, i _* i 1. 1. i'ir,|,:, = Gnidia L. L l!'-.'t|,; ^im.il.'ilti , (T, !!„). nh = Giudi i d ij.Lufe.li i 1 < Rogeis Arthrosolm L \ Air-, - > . timdii, L. •?-, , '•/■!, J jf „. In = Umdin a-ipbnitolKi I i daphnifoliaLf. Leandii k.ic-au, [L. lu .11) Z S R ( ,»eis /"*••'" L.vui(lii - <.niilia daphnifolia L. f. www.mbgpress.org CONTENTS Biogeography and Phyiogeny of Cardamine (Brassicaceae) Tor Carlsen, Walter Bleeker, Herbert Hurka, Reidw Brochmann 215 A laxonomi, Revision ol ill, Syriuga ,mbescens Complex |( >1, a, < a. ) ntril)ute ating outsidi [he ! ; den >rinted in the back of the last issue of each volume and are also available online at www. ml Editorial Co C. Hollowell " Editor, otanical Garden Beth Parada ig Editor, Allison M. Brock e Editor, otanical Garden Tammy Charron \ssistant, Missouri Botani Cirri Moran Press Coordinator, Roy E. Cereau Latin Editor, : . ' . nte, oldblatt Missouri Botani IcPherson Missouri Botanical Garden e Taylor Missouri Botanical Garden Henk van der Weiff For subscripts Garden, % Allen M, keting & Management, P.O. Box 1897, Lawreru KS 66044-8897. Subscription price for 2009 $175 per volume U.S., $185 Canada & Mexk $210 all other countries. Four issues per volun Tl,c journal Novo,, is included ir : Tmf, Annals of tin-: Missoum Rotanicm G\i:i>ln 26-6493) is published quarterly by the Missouri Botanical Garden, 2345 'lower Grove Avenue, Si. Louis, MO 63110. Periodicals post- age paid at St. Louis, MO and additional mail- Lress changes to Annus oi mi Missoi hi Rotanicm Garden, % Allen Marketing & Management. P.O. Box 1897, Lawrence, KS 66044-8897. stract/Clobal Health databas, , in » 'OH, Research Alert®, and Sri Search®. The full-text of Annals of the Missouri Botanical Garden is available online though BioOne™ (http:// www.bioone.org). © Missouri Botanical Garden Press 2009 The mission of the Missouri Botanical Garden is to discov © This paper meets the requirements of ANSI/NISO Z39.48- 1 992 (Permanence of Paper). Volume 96 Annals ^j||k Number 3 of the ^"5C 2009 Missouri Botanical Garden BIODIVERSITY AND Peter M ller J rg CONSERVATION IN THE ANDES: INTRODUCTION 12 varied region in a single day? the area. Missouri Botanical sity and Con- a diversity of , that developed in undemanding of the critical questions— « hat species *egetation came into existence onh lecenth (3.5 . .. 1.M...J thai i.., ilrh n. .uulu-. nil Mil - mHishl i il. -,[ it f I. »U.ms «,i _-,.,0-'(>i.f, m rii./l linl, Lm\ . m iml'iKi collaboiatois and space or for differen, law, :'..-.■■. Valleys, rivers, and ridges— aie the. all equalh Luge data set of collection data, important as bar on, isolation, The Andes is one of the world's cradles of tion units, using g. in the past and «ill continue to hme in the future. m. 1.1,1 oi-im- Hoi.- i.. .-., THE ANDES: A GEOLOGICAL Man Graham 1 * OVERVIEW FROM A BIOLOGICAL PERSPECTIVE Hi .1.1^ ,1 at- .1 lli» iix.i-i ihxs.ojifhi. ,.']., ,t >\„{„, l it ,-n Ian, n-e dunng the in !>..]«. .a 0, m th. HiwtU - a Hi, M.M...1 Lu-m ,1 m. at.ante\ent The inllutnce of altitudmai gradients (8848 m) are higher than the Andes with Cerro o tionary processes is t. t ' wlh.Il ,111. „d, in ,] il.ll tut . .ntlul .!> Caryop eae, Empetrum L.) in until South America separated horn \f„ra ,t . , 17^^^^ ^^ ih^.iL!^ l\ . Md >Mli. hnti t t ( itB ulUn. ci . ) - )t) t _ i(| iij)( lh ijjj,,,^ «'• -'<*" T1 - I'- 1 ' i! "'- ™ d l! " ,., UW v.„ ,, a „.,.,,„. e"t al, 1999) as fllt-o '<.! .MllMVu MtsV Z^T^.Z^TLl L l!i ""r 1 . "J/, ■.ill, habits of Mtocene to Quateman rf ^ ^ „ , ,,„ l( f< ^ jn , )[])M „, _„ ,„ ](l age, and Mercer e, al. (1975) found ev,dence of „ by 3.5 Ma. The principal tectonic rn Andes are generated by the Antarctic Plate subducting ■„ The central Andes is the longest segment of the i'x. Mi n ^ ill' \ut n.lWin pht ,,| li.tmt],,,, „, P , ,,i , 5200 km of the 9000- and paittrulaih betu- u "II »V '_' l\l . .ml 25 ,n.l m.U.a , , ... a.u.H , , ,~h downward. Relatne sea-Wl use at \anous times, T o as mam as se\en .•■•■.- Wi - li,.^.itii ( ,..l I.S to 20 ah, .,! 1. m, f , ..n.l Mi'n, . < ho . is «e)l [ „o v , <1,m- to' h- lowlands of Bolr. a Hi.. .< n «h ha, i„ ii, .,iu : ii.Hdl.iUn ta-iLmu/m!,- m, >i. ! ii, -d. ,. ; L, Passage took , ■■.:■;■ \anes «ith the di'^.n' b., r r b ,. v s t!„ ,, |, , ,„ , ,[, ,, ,\ .. ,h the ,piead of aias,] , d m I l , ,i t ((,,!.,„ ,1 l„„ Hn.lw ,h. I .. . ,n . A ,, si , t rl„ „ si 1999: chapter 7). In western South America ca. hsequently, there ] r - M.i, tL, < ,.n *, ,. ..n.,1].-. . 1, i ¥ < - ,„, ,-• fin i imiescale that leflect an ecosystem recognizable as desert. the driest int. 1500 years BP) the Amazon Basin and ,n the already ~ " bean Plate at ca. 12°N (Fig. 5). V focus of m . •• • l ■ '.>.,., I, in.. i> |" I .'I lln \lli|.l. - In ll U.I. ■- ..I.. I lo I"' .". < --, ( I,, ||,.„ until 21 thousan, around the glacial maximum as ca. 500 m northern coast as the Caribbean Mountain sWem belt™ the current 4850 m. Then a warmh iged to foim the Netheilands 7 m .1-1 ilCitlv i Pn „1„, i> ,Wi Itwk *' <3 ' ordillera Oriental and Middle Miocene, and delta deposits of the same age idcaiut oiigm ^ hil has brought vole mir ,.1 „ I k n.l , . n s ntoil. .1 n ( m.l M and it is estimated that the northwestern coast. One of these is the Cordillera Sierra de Perija has risen 11-16 mm/yr. since the on the immediately -na„vn,Utu Mgion In ill. L.i.e kin, i ilj. liui , Pi '..Iniali i between the upper Miocene, uplift of the Cordillera Oriental of the h of the river. Patton (quoted River and formed striking ... is that all 11 [of the 17 species] are reconstruction resulted in exf ch, which is one of ■■'■■••..:'■■■:. >l dim ill mi l.,i. u la I\ ll, M | L Jr ,,Im\ 1')'.. ill 'till,!, B t m include, the ^edtest late part of the Mi( \s, belieie that this dhersih present position in the Venezuelan Basin south of r. uentation and reas- ■■■■■■'■:■ ■ ■ • . •„■ .. •■■■: •■.:■ and tributaries in the Amazon and Orinoco basins 1= nargins of the basin, were the result oi lem Andes. ago. The suggestion is that the eient drasticalh 5 r Dryas ]'.... U n..l tlit Tittle Ice Age (1300-1850 CE). Basm has been the site of impoitant p ilpopnvnon- iD It, oeuts «.,„, i. v -iW,. M) hpat m the foim of »aim surfarp vw.tpis from the Andean vegetation dining cold intervals th.i v»i i -1.) »h-r< it \ hjh- ..nd le'utiis j« mi imilli ,utl suspected to have J When results of the Climate: Long-range Investi- rainforest. Later studi. h„ si, ullusi l„ < L , |, < on M i P1 n> ml T'l .]..«,.', ] i |. ' (UlMvF lepiesent d b the 1 sin s di n ,i - ur 1 il, » I , I M , „ , , , t i ] t i i. i t s m , 1 . -i •• .,1 • c t ,!. , i In l uZ'"'!,"-, \«' J 1 "',!!? ^tlTY" 1 , „' f 1 *.'!'.'' ',' " m r ' *" n (1976, 1981, 198: ray . L. B.. C Ciofi, D. Geist, J. nities and other at today a] and from Brazil, Colombia, and Vene ha I .,! ' n r'l i. f t, , ,,' , tamable development efforts that fact alone implie: . G. 0. Seltzei. K. R. Kelt-, & J. Southern. 1997b. Chong. G. Franz, P. Giese, H.-J. Gotze, V. A. Ramos,' M. Kingdom. & I Ai Pud I'M) W.HKiju belt-, and the mw Heme, K. & J. T. Heine. 1996. Late glacial l'i.<~ .V f u„ , 'I 1 W'»6 The changing course of the Dryas time. Arctic Alpine Res. 28: 496-501. lv \J . K 11 H <•, I I ]1 b t « B H , i„ Hiii II l"84 Gestational and climatic hi Soc. 129: 93-131. .h. 1.1.1 -.i i - .J> I ul-P • MicW P.K-...I f, l".'-U.i Ghosh, P., C. N. Garzio, f Sci . Rev . 44. 14^-183. 435: 865 - . laeoecol.' 109: 1-54. eand Jaramillo, C. A. & D. L. 1 li ll>"l \ T ej Killeen, T. J., E. Garci , B. lA^'ll 19< : Rodbell,D. T. >00. Rapid ice margin : 821-824, " u hi .1 t id P„ . *-" II ,\< , i 1 \y,, [o',»,'m] Nni'/ M K J Li I , i, A i M Bell. 2000. Timing S. H .1. 1. 1. n and CLIMATE IN THE DRY CENTRAL < Jay Quade, 3 ANDES OVER GEOLOGIC, Jull ° L BeU ' han P^hett, 3 MILLENNIAL, AND INTERANNUAL TIMESCALES 1 . ^ ...! 27 > and u.l.M.p.nvl timc.l.s TV! ,-tW t. -Im, t, nl, h-i. ] iU t.» .1 g.M (hull n 1 tl - 1 u«.^t, .illui.lun •.."•.'■ high enough to exclude much of the moisture .ditions as recently . details and timing in uplift and occur both in ; ighlands and across .'■',...:■ .... ,..■••'....': ' : ■ , ■■ ..:■:■.• uppermost few met, rface and can from relict b lat mam of these record the age of material suddenh exposed or 1 Ma (Placzek et al, constrain erosion rates (Lai, 1991). Cos iity. This movement faces in the \tacama lesult in some of the oldest suggest that penodic lainfall and flood eNents inferred from dated egion. Here, we foci 2009). Together, the. n the Altiplano am lake that is mon ep (Argollo & o,h, H tJ l'»'»J ( flu Bio L>.s>_, - Hi „, ,1, ',, r hlel .,' appa'ient in the sl J .",.„.!,.. - «e of two deep-lake and se SCl.ll Itlil.Ol- over the past 120 y there are mos ..]( flats m ine lakes. The Ouki lake c. rele was ca. and 19 U-Th dat< dee, ike e|, IHO.L.d' ' ero empties i nto the ohgosalme Lake Poopo 1685 m . >! .1.., ,!< ( Oljds, km 2 ), which is separated by a le, the Laka sill (3700 m), from the i (3656 m, 2530 km 2 ) and Salar de 12,100 km 2 ). In wet years these salt lak cycle; md berweei !4ani >0.5 ;. 3). The Tauca lake c i i 111 . If :ted and filled with shallow water dart, 2000). basins, multiple sites were studied 120 ka. Multiple 14 C and U-Th dates direct determination of lake level, replication of ori an0 (Fig. 4). ll.o! , ( id 1 lb PI,. /.k A. p. « 3760- -§3720- 1 Tauca Elevation (m CO CO CO CD CD -vl Q) 00 O O O O 1 1 1 Coipasa j : ■TTivrrrT'Vi ■ ■"■ 1 Sajsi o o < ### La A/Zna Air-- £/ W/no II 10 15 20 25 30 n Pluvial Event (CAPE). - I'.l .' <[] xl .1 |«s,t jii, ■ II ., ! i t >t In. s,,,l J^cU i.luu a Jtp.sjl , .1. ,tr n i .U^i -ire- ( v n,.->.tn»,i. i ^j.iliti" hu 1 ■.ju.ll, ,1^1-laV.l in*.. » lie. uUt 8 ka (Fig. 4). itral Andes. The CAPE is divided into t and the depths of t isa lake however, suggest that the fir on the Coipasa' id 10] -io. Irec let, 1989; Bensor : 1 . 2002) Thus ihc lake i. . cli mate j U c1 at the lev. In Hie a- ,,l f] ae P( sWcm. th« 3 P< 1/3 th. Tl us ' a al ;tii. il ' a a sill (the s . Thi :\l\ Ith, , Coipasa and Uyuni l..t th-L.L 1,11 Sh( felines would nc • ,Zllt phase of CAPE Toda., the sources, timing, and Mmalrilih of il LA,- T.w .. r.i I..,th .,i. ,is. the I,. .1 ]U . .. n „uu 10 Ma), the uplift of the Andes 2003); and (4) changes in sea sur face temperature rmation of the Altiplano plateau are critical gradients (e.g., Betancourt et al, 2( )00; Garreaud et in the fon nation of the Ind i, ain .- ■ n king zek et al, 2006b; Qu, ade et al, 2008). ma Desert uniquely long-lived and arid. and we rule out local January i] stead, ev insolation. The Tauca lake cycle rea ched a maximum between 16.4 and 14.1 ka, ca. 5ka after the indicating no direct link be y is current l y mo : ■■:■-■■■■'■■ "mate changes in aridity over the region are, (18-1 _, u b . .![. h.nn .., IVMI ..j.l ' " ' mtelac hou the « estern Andean flank and may play a greater role heh n„EM) m^j, ipit .i I in m lies n , 11,- ,,,,„, ,;, Ut , piii e (|f ( y> E (aftel j i ka) lhus wetland and roden to the operation of the uiiiili m mil si. iill>. hi poihijiis of the cenfi. Andes during CAPE first phase of CAPE coincides with i Literature Cited Hyperaridity in the core of the Atacama Dese lominates over a period greater than 10 Ma, i :ontrast to the western \ndean flank and tl ( Barnes T B T \ Ehl.i, > "" > i < .. P I I - 1U1 in." V i J ',)...( n K« '>! '.,-lwi , M. Vui 003. The climate of the ^eis ocene piecurs. e ea ea. ua ern. es. . ^^^ Q ^ p Molna ^ j c Libarkin & R j -i Lot LN1 . Cartajena & B. Mess :s. 51: 14^26. M. C. Hein, T.-L. Ku & S. Luo. 2001. A 106 ka ™ Dunai. T. J.. G. A. Gonzalez Lopez & .1. .In, / I ■ _' 05. 5 13 G and F U G in (he .-pin. - Volume 96, Number 3 Placzek et al. 2009 Climate in the Dry Central Andes I.1.U.-.H-. 1 .,r...-!n, 1 iichalski & A. M. Cows it M '.i>. H t , km- < I .1 JU.'f, 1 1, l,t,a et al , 2007b). ■.,.■•::■■;. ..•■'. Ou3) Because diffeient measuies of past Km >,< ml pi.^nt »,].,m m e M )lainmg the blindly as a "currency of biodiversity." regional conte ke a continent-wide measures reflects ecology such as contemporary tations of di> endemism in the 07148 : /.-.! Pi ik ^uKIDMAr -.Urt.-e M1 despie,,d hut lait specie - In ,1. 2< 2006, unpublished data). This f. 03). Only 60% of 1 I leiie-trial ei II. I t» tn liunmh, (ra.lv h l.Hx.n .Imli. m innu, 4 ,l] U sti ate, [ in Fig- analyzed t! < di,ei,itjf .iion ot l!,e ^H'lij) !•• jp ill" ,i, 11 Llus' -p»< . - rfie ' ..] \ iuJv j number of nodes to the root of the phylogeny. For a subtiopics, with peak concentiations in dynamic simple illustration, we dhided the' species into fl imp) and saAanna quartiles of branch lengths, «ith the first quartile h i in Ecuador; Loreto mosaics of Amapa, Bra that of the 2 ad South American HESULTS birds, which .est explained from VARIATION IN SPECIES RICHNESS IN SPACE AND TIME ecosystem pi. et al., 2006). The PA J... Ill published molecul '. ■ ■ nc o ancient pecies in the hydrol, v e , 6! Our n ext examples are from the larges ,,„,„, to the Eocene o (more than 1100 species), whi ; screamers, Anhi . Jae: . subject t o detailed phylogenetic studA (Ires Opistkoc broad-billed Sap eldsa et al., 2007a aenigmc , - cies represent lin : : " 07; Ohlson et al., 2008). The ear Tertiary) catchers e see them here as surviving pu-euiatiAe manakin , Vml.MK, 1,1, Km M -1 ,„J ..-iu>, n^ad ,,nhuv ,11.. I,. .„i, m ,h. . ,|. tin- nn ..up] .'txtensn, swamp throughout the humid tropical forest Monies, with the mcuisions in the \mazon Ba-m. The mole demed Andes. the grassland 'lata and Bio Negro Tuanmdaeis sti Fi_2Bl - 1 rh.l s .,i < I ' - i>m cloud foi est zone W. the Andes. The sin shows that the The specie for range-restricted The tropical Andes region stands out cleaih as ihe \ndes (c g , Mitteisfuld et a] , J99f ■, center of avian dr. Neogene. outlined areas characterized In separate sets of To illustrate h< i old South species but paid less attention to then stiongh of spent s (Fig -!V m >,i «>' *li. m >, mm., H,„ ..[„.t.,i nd h ,t- ,li I > t \» l=an < loud fon st is North Peru Low, anc ) [Peru] and southern La Paz and 123 on the slopes of the adjacent rain shadow basin. The species richness curve (full line in Fig. 5) of high-altitude woodland deeper into the highland. ridge. In the i species richness is Typically, the mo pecies inhabit lower. The kin liversity peak on the corresponding elev. ent slopes (in Wi slope seems hpical of elation giadients some cases replacin ven if there is (Rahbek, 1995), although there is some \ariation in different elation zones on the same slopes (see tl .ez, 1999; Kessler et Krabbe & Schulenbei more distantly al., 2001). related (review in G ldsa, 1999). The endemi je rarity per species, stippled line in Fig. 5) describes a different pattern, THE VARIATION IN AVIAN DIVERSITY ON AN ANDEAN with ^^ loW Species) at the base The local \ariation in biodhersift parameters ii act but is mostly within an Andean hot point is illustrated in Figure 5. depressed to below 3000 m as a consequence of Humid Andean slope 1000 m 2000 m 3000 m 4000 m 3000 rr 010 -,ml l.u.miu to > r 1\. nt, i . .1. .\\.,i JOI't.. "nr i ,. 1 I..] ,! 2003; Kreft & Jetz, 2007). In birds, ca. 3000 f; diversification rates continental species 03), far more over the same 2006). The contrast occur in northern South America (Fig. 1). This p ems of endemism. '....■■■ .,. Stotz et al., et al, 1998). The Dhont et al., 200 occurred in flow (Vuilleu rill, 1992). These v ....-■, .'. Neotropical Montane Forest Birds %: s ^*fi£r Oto 1000 meters j 1000 to 2000 mete > 2000 meters iver valleys, and hig sterns of endemism, m distinct conservation regions. evergreen for '00-1500 m (locally lower, especia ic slope of the north Materials and Methods Andes and in ds; Gentry, 1986) to the tree line, latitudes generally TAXON SAMPLING varies from 30 00-3500 m. Humid montane forest is irevioush publish .\ ,,!„)„ w n ( ,„ , M„. . „., ,,Ju. » s ;iim.<<.I I I t! I-, piOjt.t If .M. - 1 2 tai.l piotorols (^eir et al., 2008). Model ,,..1.1.-..! ,. ie,s. hm. lt ,i ;"» " v^r^'vi'T?' ,\ iTi 1 . rvi'ii'i leters were estimated .. ,J.U -..Vdplli. - -Ui-ilUi tain GTR-r distal viduals. (Table 1) Sampling V .thin >o, o->, j.ld. -,>i< i« s , , m..M -iltJ.) tl « " "i.'itli-n ,it,n, -t ,<]< s, i* ,-1 Costa Rica. Samples from Colombi... \ene/i^i .. wl ' ' species (Table 1). transformed into time esti ,!j.i UMI s.ul, .1 l„«l], 1 .. I» , M l.-Hl e.i. dl-j,, 1-,. 1 ,« o-s a Areas of highland night result in the formation of founder Neotropical birds In Cracraft (1985) for South p r fomiation. Again, (1998) and Stotz et al. (1996) for the entire b ition will occur on Neotropical region. The regions of endemism defined t jtli. , -i i oi . I*- .„ .> in .11 .1 - .]< ,s -J 1 », bunt ,1 .p. nnl^ ..i ,i „e flow, uhich may Neotropical Montane Forest Birds 1 t- >m < A Id. ,ik. m In a-, , , if iu.ll ran also be code.! Imt lesult in n-l-p ok, ■ ■ ■ graphic speci egion but was not equivalent if no st ed at the time equal weight to each column in the supermatrk coalescent times predate population splitting times by analysis have traditionally been j,. Awm- I m srp ,|, , tree for a common set . »s (supertree replicat . "l op: . . v fej ■■':■■-. ; § § S B- - r , f f "J - - | 1 g.2 J ►.11 Neotropical Montane Forest Birds (SV) UBIAIU9J ipJO N (gV) uBuopenoa iseg (SV) UBIAIU9J ipJO N eS eS fS fS !""!" p™:S. las, Santa Fe STRI EC-HRU-514 >. km W „{ ' 51 '■'.' ■' i % STRI PA-TAL-1014 character independence is not met. Furthe [SCAC]). This Neotropical Montane Forest Birds h),hh>,A ii, tboK lh..r i, .«]>.i yli i -nlialK ,i m i n, m'hir ,. « Lowland isthmuses in lower Central America. The Guatemalan r om the Talamancan , n JOi'O i, in, a- Ih. hi,' 2, 1 -"^ J> iPj t .d hail a sii iilai distubution of phJopt.utu.Jk iirf...i,,l. , ,[,„ dlll | ^li ,„ b.. ' ,1 »,7"-il 0", U ■■■■■ j. Jlwnf umtueclididctei- *' * ' ^ '* 3h, .. i. would f. cee .,...■..■ . I..*,,,-;,. „„,,! 1 l„»- >•«"' ''"'•'""" '»«''«"'/ ^— The Santa Marta, Mil, branch Parian, ami. i regi :nd is not i, ementetl here. ■->»'"'"• \.hI-~I-> L-wland the SAC and SCAC analyses except that Cinclus endemic specie* In - l,",l and subspecies (n = 55) , puhli-hrd pin I,,-,- >» GenBank. In the samples available, genetic netic anahsis of Cinclus iWlke, 2002) were not divergence betueen ||„ - „,l , M.iila and east Results (Fig ' 3C) ' Th * " " N a " lsolated GENETIC DISTANCE ANALYSIS Venezuela. It i ,i . . • lowland barriers ■■■ versus conspecific populations. distinct (Myioborus S. F. Baird; 6.2 n.i. ,u lu/lil....! 1.61... Mexican and Gua mazon and Orinoco basins. Neotropical Montane Forest Birds • i.l.Jleu, .1 th. \i,l - Im, i-jw] is helped to Noith of the 1-lhmi - .t h hnan, p< . m M. gus ophthalmicus she,..] <><< -li vm, t ,'ir,s this .1. 1 t '>/ ,• > ^ t «> •;•'//., '/,,.j/^ (li necessan. .■■■■■ tiation.in.,..!. 1 1 -(,'. <} and suggesting f 1.6-3 Ma (Fig. 2E). dilh.M.tut rll- ^--Idai. I r TVUk.u.t.,u-.n 11. f n) , ,|„ species had 124 dii • | >jh'M.m -.\ ]Jo...tthirm-t.a. 2o00; Hoo-biemstra «X \an uncovered in the reW.I-.- ' , 'l der Hammen, 2004). Neotropical Montane Forest Birds Mexican clades, alth ability was low Amorerec. last 1 million years Ji.lh. >U ,oj<,l.^, T.l.u.M j .1 1 K.--2- -,„ n, ,ii |.t,<>.m LWiLTM'.. i,«st.,l will,!,, II,. -vi Hi, Im. ,-. ,:M.,.bti ». s e\eie glacutions of the \ndes (Bennett. 1990; TopologA in those p Us t tn- *-> I' and - ,l fu, F , ,J .. .„,n ! ,1 i ,emstn cV un der c (S( • . ■ • • im IJ . .1. .. Jl. n n lill r„ i, \{ m h, l.hii I >\ lute .-, Discussion (Figs. 2, 3), but none of them exhibited a distinct interchange burst during this time as obsened between the i, and west Andes. i*' -It l.s- I, ..,t,l dii-, i, f . 9 ,H i mil. -it-ilv b. ti,. >i Pieist m,i, ,s ] ml. nh, , In hhnl ,,-h.i ,11, ,h. J."!,-..,! an! in !il,»in lunri, I Mann-'tt hitne, [ ha\e been hindered by competition with the species- , n 'tn the ralaiiLiica and Dan. , ai.l ..t.n.l mi m an, I |U u„. limit,.,,, hij, pi »,.„,„,, ^ Cranioleuca complex Chlorospingus compli h stable areas ma) help maint Hemispingus comple> Henicorhina leucophrys \S -E! nphus pusillus Lampornis amethyi Hemispingus complex A Ochthoeca cinnamomeiver, lodytes complex ^1 \lmost 50% (16 of U) of populations fiom ad,acent acioss liver valley barriers, followed by isolation by endemism regions • ,ys date to the those barriers. However, a large proportion (1 1 of 36) of r^i ~~ 11-8^ branch length (substitutions per site) divergence events a ross river valleys. These results the late Pliocene and Pleistocene glacial cycles a role played by arid Neotropical Montane Forest Birds The role of dispersal and Variance the foimation of patterns of endemis lowland to highland regions (i.e., Weir, 2006; Brumfield & I owever, Ribas et al. rften been Eastern Coidilleia of the Andes. Although such a ersal from vica a role in the origin Lowland Barriers 12 8 3 6 o 4 2 At i River Valley Barriers 10-1 2.0 2 4 6 8 10 12 14 16 genetic distance (%) v isoU.-l <-■] .lie ,!.!-• ol .1 barrier, dispersal across the barrier could prodn. <• I nr.,jn.i..ir>K ll.e - oW... I,., .atme does not L< ]• si 10 ,nil]i,u-^i i.i f.n ].i .nl IVma.iddi. i^-i ", million wmi-i.,< ol. mh.-K^M,- ,i nl 1 x> li i Li I It. iii'mI t >i..i, ii lu M - 1 xsts of diveigence dates !,,,.. „hk1 Non-forested Montane Barriers with form such a pattern (Fig. 31, J). The Rio Marafi 2 4 6 8 10 12 14 1' genetic distance (%) , su.^lli r,ln in " .trli \m i, . .1. Mm , (i ,il,< in <;/ i , ,, ,.l, ».,.) ...il (- ( ) ',, I '. t, (V( \( ) ( j tl i 1 I , . ', 1 , , s ,1 , ,, ,1 1 1 1, ,1K 'lH Hit. I ities ujth luj-liimd i i n t t>. n. ih n >,» . , fl , ,, i J ,n ,r in. i- , . n- fn I ..I . '.■... detected for the Rio Apurimac. species conce hese results suggest R.-lmonship^ is, Mnldk \merica and Mexico are that mam currenth defined hjflila.i.l ^pou.- contain not well resolved for deeper nodes, but do strongly multiple, unrecognized species. As the biologic .1 1^1 ri 1 les . it! n In '1 . I !,„, 1 rli. >« i li Vlu ,k ... < 1 .ssitication Commit i each other. tee (Remsen et al., 2007) for South American birds ir.lt null, . .!. ... uislin. lb cl< uLn.lbain.isha\e k\el taxa as tl l,.i irat ,< ,.^u >tz et al., 1996; (k 2005,. FAi-,t„- ( 1 . n> .ge is louei iDorkon & (,enti 1 *>1 li. I n ti.,.ii..^ . ;. it h~i* tl.it ii,...r , uLsn.^vl mom^ne k>,< st ,,,,-n. boundaries do not adequately represent all evolul arily distinct lineages. Under the biologic spt Neotropical Montane Forest Birds gen. 16: 11-21. remote 100: 629-640. j'h 1 i i 1 (1 ._ .' -'-.-I 1], 1 i, F, i _] Ll-1, ink-on 1 s Collin-, T M C lomi, HI I n'. I- \\ J n, 'i I i 1 \ ,1 1 1 - M M f -d,ede 2002 112: 156-170. Dhont, D., G. Ba< ke & "> U ,o„ . H. in, K 1 )":' 1 i. H, ,'.i,i »1 . i J . i. Kwi ih JERS imagery. TpitomH.li ,i.s Vi 2->w^|_ Dm»l< ( I 1 Lo,,t !f l i,d., i r I, -1..1.1, ZO.6 Neoiio,.Kdl plant dnnsrU la. . Bronx. Annals of the Missouri Botanical Garden 124: 537-551. Mayr, E. & L. L. •iLgdom." & J. Diamond 2001 The Buds .., M t , tl , . i ^ (n , ( jN | ', 11)r( ewe, R., C. Navarro & C. J. Villa-De Leo Millei. M. J.. E. Beimingham & R. E. Ricklefs L>fi'i.m i \ - 1 l-i. n 1 Hi i 1 ii.,. i ' - > rt Mu, I) Rohwer, R. C. 1. Ukbuin p\l Benneu K "l' . t. 't ,\ [" P 7 , 'm m_l, I ,d , V ,t i, , >, .l\-is „ lu ,, r l„te legions and habitats. Various another dimension that ranks these organisms as by the Han aid entomologist E. 0. Wilson, pla} a degrees of latitude, from the coastal ranges and montane ranges in northwestern Argentina. This collections housed in the Berlin herbarium «ere arched backbone ..I .- ,. , • ' • : ■ . ■':■..,,:, Peru to noitrmest \igentina) at 5 S to 27°S. The ti e.g., plant stature, and slightly less than 9% of the t.,t .1 I >nl -vi* » . , ,h. s„l. f es desoibed at that time In ditfe. ui mill -s Churchill (2000). the increasing nication was limited or long dela. coixespondence and The Myth of Tropical Moss Diversity publications d. »h its.,, ul I t i h-tn! iuli( the. existed, and in addition, theie «eie .anous sorum, was published in 1801; in time this would be ti need for revisionary includes the first mosses collected in tropical naming for the tropical Andes: Frahm (1991) The second e.ent of that .ear occurred in the tropical An, viously published es, ■ ■ , etofore accepted \ .. L06 Edwin Bartram. The -[. hh <>t rh. . t rl.ois w ,. .1 tl, nsh« ml i i-i "lan studies Embedd. d within ■■■■■:' ■■:.-■•■ many collections sent to Miiller from South America, 1936; Pursell, 1973). A complete summan for the n.nl l-h.l. Mull. 'Iss M,t! r J,,i lit. et al (2000). All of these checklists incorporated the ■eriod in which , d i the 2! M,ll.-.. V ilium iVuti.-n. o,dk,r,i \ iLim IJimlirm- dubious species reports for the tropical Andes, above. Excl, s are all species from checklists to anah/- m i j i i lie -nenb/ations included in the analws. The initial sped-, li-t of ih- Nr..ll..n. s I,,. J I r. u lit.,, n.u, f till se ,,, lint. Irs \ultJ.. serve as examples of inflated species numbers: A second ed from this initial . .. ' .,- ,nUi. 11.. In'l-.lm ihi, an .1 sis aie the Cordillera de la Costa 1. «rjt!i.i. tolomb,,.. .„ ih, -onlr, tl„ b.mndan is defined to , 1, , , • s srch as the Neotropics. The anahsis of moss dhersift and Moss dhe :ides is estimated endemism of the t tail error, but at 1376 species, 327 genera, and 69 families -. ;, :... estimate than to err on the sid species estimated diversity. preUoush (Churchill et al, 1995), whose figures I I) tl. ia .).. in. ti I m. ,1 .1 Ih i. | i i! Ami. i, lh> Wl ,i. Is I-citl, ., | ,l L ! L siinnl .,( HuM i |[, ,d m .n.tu* nt Misuul.t,,,, 1 ,. c.nll ...n! it i! , ii al Wes is 929 Garden Tropicos system; (3) a compiled world (Appendix 1), many of which were recognized in the Venezuela i>. 68i <; X V f ColombiaV w^ ? v -/ /v 883 \ h \ \ Ecuador,' /r^\^ ( 796 > I % f\ SSr iji^ ^~^~^? \V %Peru \884 ' j Tropical Andes fly Jr NW Argentina / / 122 significant increase in newh described species of I ;dw.) Cardot. Table 1. The 10 m ost drverse m oss families and genera for the tropical Andes. Family No. of species Genus No. of species The 10 most diverse ; tropical Ejndlmism Table 1. Just these red endemic if it is only known ;.■':■■'. ■ . :.- ■. . : (see families discus :ss certain for Andes is 428 among 137 genera are represented by 55 of the 159 Andes (Tabk ospecific with the 3-2150 Ecuador, 'Zm (Sel"phyll a ce mIk <* W * U.it,uJiella Bioth. (Pottiaceae) is pnmai «e, ,«,... lh irj„.ll„ [1, .1 n „ pJIh) J to. ill. Peru to northwest, Lit has been recorded from a sin co ANDREAEACEAE forest near the sul nd also from ,ost Chile. \ndieaeaceae is almost exclusneh found in ■• '• ■■.-. " ■■...,., Twenty of 69 families of the tropical Andes the author, <: ly be endemic. Bartramiaceae is the fifth largest famih for the \MBLYSTOGI\CE\E ^ ldeS ' * lth ' * luch ^ endemlc - Nearly all of the genera of the Bartramiaceae are the high montane to paramo and humid puna. p le exception, mostly Ecologically, a m up a major found in montane forest as an epiphyte. Breutelia ...... ,.■.-. ...... Ul .ii-1..^ \y..tir .iiil smui ^...ti. f-oi-i,. jmlmlL , | lrl< tlu -U^ .- ..nkn-vi Crato^urorc (Sull.) Sp 1 species of Breutefio. (Schimp.) Limpr., &' BRACHYTHECIACEAE only known from tl eae is the tenth largest family, steinia 0. bid. found „>, „„ 1^ in -\u c is ot tl,- v,M .".si. ,,,. < s ,r ai. .1 «,th th n., The fimilv, as nm fiianwnml ,., mi , Ijn , I j , ,,l t /,;Di, m < / t ,„' n Bioth i email, to i lI( >i ' ,, M Mei, 1 /77«/i^u Hampe 1/rfeo; n u. 2«J, \li us en, xuiall, o, , ui ,-, Hl '' " " " l < {li D '' '"'■ • ' ' ^ the thud largest -„. „|,> „„,..,, . M ll.wh geneia aie qaphUir f „„,,„'/ '/^ /, -,.'« P/,;fJnpL/n™ M Vlei-cir'hpi, 7k irich\cl\l Andes are Aligrimm and Coscino- dontella R. S. Williams. The genus requiring ceitamh be endeniH U, tU \vA, -. ilv -tal-,. „i J2 «" unknown . contain a si § f endemic species: 0,tl »-i >,< , Mtl. 11 i I'*-!. U< ,«]!,! HYPNACEAE . -a and ,.,-j, , ar ^ idespiead and of the southern South Amencan taxa In ( alahiese Hie genus re- , ,.m- u.'-'kI .Vi. "this iu< -I h', u.l.'f ] Tl f!,t.ln .-t'. .n. 1 n, t I. '. > < ml ■ • -eil „ til i B.ul > examined ft pes ai ons from the A. Jaeger. Cm il. is one of very few described from the central Andes, particularly in Mil an W. R. Buck. Bolivia, but fewer tl .ely be recog- NECKERACEAE '' * the species in 1 in open mid to high •■■■ . . ■ ■ . ■ still requires a careful revisionary study. Neckeraceae villa sprucei is only The Pottiaceae is t rse family for endemics (Appendix the genera are common in the wet and dry paramo and >_,W ; hii.i. 1,111 ,.!mii,- th \.i.lt- ,-.,!<>, V V, M. .. i!u,i bait „1 th.-c -!>„, - lfl.UK -1 .■< .ll< -U1J. ,11 M .Jiih II. -,< ..jd.ltIK TO -.-U.jl.Vj l« /'/„..,/«. -I,™,. ,/....,;.,>■■"„ .IU , »,.„„„,/'.[[. ... homa-inj , ■ -i , h a ll, ,i*«, fr*,,,,,,. ,/,-„. and Timotimius V, R Buck, also from a single but this m ting artifact. The , Ecuador. total), thus the ) is 1137 species. At SORAPILLACEAE least a numbe e more restricted in ; ented by two species. Agai «cies are narrowly species, 5. sprucei ,nic Streptotrichum < : : lotric! ,: .: endemics (56) than Colombia (49), follow The patterns of diversity can be viewed at different valleys. levels. The si observed pattern of Vi^entma (Table 4) » l,m, ,.a «s Hi m.-l .!)«.. ,. 1 ,l„ ( i - d.w^ih .,„ , , The Andean portion »i i 1. . hump I 1 ' 'e |i mk *- osnii loi HmVd Mites, is ntaih 4 5 turns Lugti ^ -i • ! may be due to the ( rea of the Andes is classic contrary exam ECOREGION DIVERSITY i''lii,i1: ( Iilu lull m A. l'"^: -ha- H .1 , the \atious ecoiegions (essei„i„U , qi A >„ .-.t.tion birds, trees), but in some cases this appears to be ,j mor e diverse than rontiast m moss dneisrh between the two I ,m ,t s]1 , . ,,_,„! ,| „„ ,( n m , P . * band of vegetation, including both forested and open e .fir-si..n n, r h< ^mii u t > .»,!i i tiopi. A Vndean stilted a giadual .n - >„ je > ,, P ,km Ml ,t montane foiest, «nh ■■•..■... two such zones. Oi occurred in the past decade. There are now t ,.s lt K,l I h, i „OUt 1 (,Kst , ,|, t11 ]llM ll| .1 .1 . t > Mluo ■ . .1, /J* ^ N A I H^. X_ "TV 6 - TS^W 7. "XT/i \ -/ 1^ I v a V 100 .: . T..r „ , i-ill ) liti aceae 2/9 (3) [2] ■ -. rsia D. Erpodium ■'hj I [I i •iopsis Mull. Hal. 3 [1] dmia Brid. 5 [2] H,l 1. w, .ptopogon iTaUoi) tt.Kon ex Meteoriaceae 6/8 [9] )L.,ip ,,- 3 [1] Hornsch. 1 ne 1/5 (1) [16] Mmaceae 2/2 [1] Mnium Hedw. 1, Plagio, 1[1] .5 9/17 (3) [6] Myriniaceae 1/1 [2] .l». (1 |1| ".,.,» a .1-. i M Fl-wh 1 il). AV. u .'.j ,. Mull Racopilum P. Beauv. 2 [2] Octoblepharaceae 1/6 Regmatodon Brid. 1 [1] ; 2/51 (33) [37] 33 (22) [21] Phyllogoniaceae 1/3 [5] [1] n Brid. 3 [5] idiaceae 1/1 L] (\ < Mm 1 1)1 hi - ! i locnvhur , b Button ex Bic th. 1 R. llurk K.I (<.![')] 6/10 (2) [2] 4 [1]. Eu ANDEAN SPECIATION AND Lena Struwe, inar Heiberg,* and VICARIANCE IN NEOTROPICAL Jason R - Grant " MACROCARPAEA (GENTIANACEAE-HELIEAE) 1 . -v p, i . i ( . 1L (a, . nir r-p « v> . vi ip H' > .1 T]Rr . >vi < >i ( . - » ip HUCP HFO Hll UN »B E\PV HUM IBsn IE K L LD LINN TOJ\ I > I -• \1 \ , V u I! ' lln \1 'i v[i:lt wp, w m< ji min mo mol M'-b mu m\ ^ \ > <>m t nr p<>\r ^i pr- ,< QAP. QCA, QCNE, QPLS, QUSF R KP. t \ » -I K' -1 L sp '-PF 1E\ U U< Ll( \U. LIDBC. IIPCB ' F 08901-8551, U.S.A. .Ici I'HiJ-yvriMii ■■..■ have spread north and sou stud, of its «ood anatomy .In.i. « . '!.!n lime-, thiough lepeated glacntion- Both heibaceou- ance^o,- |l ,ikji ,-t J M ,.,., _0t»5> V i>h allopatnc and sMiipatnc speciation piocesses could 105 species currently recognized, Macrocarpaea is by have resulted in such large species dhersin. f, i the Helieae, while Generally speaking, the the. era in this tribe are either Sranl il patric events. 2007, 2008; I 2003). Fieldwork by i.s I ... , (I <„ ,u.-,t ] >r, „<' > m I „„., 1 , „/,„„ , „ J ,^ 1( , , ., it ,-.1, (Griseb.) Gilg. 1. Continental patterns: What is the relationship palh in the s from Venezuela, m. rf.i.i.it , 11,., ituO -i,us o\eilapping (sym- Magimlifoliae Ewan, 50% of its species). The genus is lutii. i -i,k. k,,a lux. -,11ml .t m temperate South Neotropics at (30- m ele\ation. Andean, spe, r,, and dispersal. .. -,. i I' , , , M.ln. ih U 1 1 rid ,., ,1 . II t .1. ) > I tl - large (up to 7 cm long te, yellow, to kind of study. pollen exine iracter 9 identifies chloioplast data as well as nuclear ITS markers oi nme m rs and anahzed al., 2002). ping was doi ates in PAUP* v.4 areas, ITS and NTS -NTS]). These (Rambaut, 2002). (GPS) coordinates were included in the anahsis. chaiacteis «eie col I ].,. ,11 „ < 1, Wl s,, ,„ i,.«JS <> _ i!>Pl R< II n. <- < An pollen and seed sui tit & Struwe, produced. In n 1 <\ ,st, tl i s , , o il m» , s htm the SLEM anahsis «ings, and chaiactei 3 is coded based on whethei Foi the D genus distribution rically winged). ( haiactei 3 ,- (lit weight .t ai J Ion. .1 , I, ,,,„, 11,. , i ,,l,hcd aieas wete V individual seed in B, Pantepui of the Guayana 0.1 mg in Macroca, lan 0.1 mg in Shield; C, G the Caribbean; D, ccidental in ' Venezuela; H, amba region in Peru; J, Bolivia itral in southern Peru; K, Amazon Bas bia, Peru, and tion. Venezuela. A map o .western South Areas were delim ial, historical, and geological feati . - «,i > . Jh.i- » > < ' >ilv J '" t!i... c-l. II, Niwu 1 tiJ statistical e\pla . . , ■■■ ■ ■> to western Panama (Sjapp. Hon JM»„ 1 h, t!„,x Ml i — »tt Lvrt I ,M„ , -<,! V, .Lmo-hI V\ .,L. V k i> .nrntam langes The CoidilU , i .1 M'.al.l, !■>■ <■' h „t », P „ , 1 2002). The two sou grid; scale: 1 km), geological Abancay Deflection, another shear zone 6; unit: °C X 10; format: grid; (Clapperton, 1993). scale: 1 km), n (BI012; unit: mm; For each node in the phylogenetic tree, extracted > -.logic data aie ^ tra Mh,i f !I!'v, a J. R rant & Si dm ><* i/«/rrs,,,./7i'; ltd ■ N'Y) mbia, /. R. G i/ t J3J8 . I : /; ' l III / i. 'v d ' ff. C ^ r f il>. Jumu.-nr-, , m« o ',7 in, I. ,-U.l ■, ,, ,tnih s.mpatur (shghth o\ei lapping), or symp for US and W nucleotide I,. ^ NTS TV im L Lifpiu^f a,nili,ni lin . '^, ,1p.1 IOH ».«--,( lK-LhJ, .1 spit^l -listiibution of each species. [CI] = 65, letention index [RI] = 74, Tig J *j n -t hk«-l n, n k.si s, ,, „!, ,. .MfM.i.i c,.,n Mil, n m i ) |( ii itlnc Jh M!C | , |. ,{,s* | )ril 1 np, n. - r » m.i.I (,',//,•, -Upe pollen (clades II and III) and swiipatnc (= paitialh d l],.,,.fi< ,, „.. 1 J! ..II.,., .,1 U tin in 1/ „,, „ ,I L „ rl. opt.m, .1. , „ I m optun ,J i< „ i M --ia.,1 -1 ,(»«.<- „f si. 1, a, tli I. pins, 1/ neWznue 1 ithin Maaocwptiea. theie a.e 17 dispeisal d species are placed the southern Figure 4. 123456789 111111000 00-011011 00-011100 00-011100 00-012100 111001000 111001000 00-001000 00-001000 111001000 111001000 111001000 110100000 111001000 ?????1000 111001000 00-001000 111001000 00-001000 ?????0000 00-001000 00-001000 00-001000 00-001000 111001000 111000000 111001000 110100000 111001000 111001000 111001000 111001000 111001000 00-001000 110101000 111001000 00-001000 00-001000 111001000 ??-??0000 00-001000 111001000 ??-??0000 00-001000 110100000 111001000 00-001000 00-001000 111001000 111000000 110100000 00-001000 111001000 00-001000 123456789 111001000 00-001000 110100000 00-001000 00-001000 111001000 110100000 ?????1000 mg/seed (0), > 0.1 m, (1). The selected optimization (Fig. 4) shows an ances- and the Amotap ;one in northern l.r ht-t ,i..,raif wi.t lies and also imohe one dispeisal to and one horn Vicariance and dispersal between tht Cordillera Oriental (G) to Cordillera Central (E). • ! ,-d.k ,<.url»,aruthward (two both to I d from J to I. Northward or and Colombia (E and are only found in two 01000000001 "^ ° ' ' ' ^ -i"-Je- 01000000000 ^'"P^tiiL 10 md j„i , ...sat 00000001111 indicate 00010000000 than bo 00000000010 Allopatric lerminal species ] 00000000010 nll) "iMl oooooooooio F oooooooiooo ] our 00000000010 '' ' ' '' " } ' l f' i "'' J ""' oooooiooooo ill.),'. :.:i. v .'-,/.,(..> .i k (,),,. 00000001000 '., -,...„ j v. p ml <,l „is,m,]|. , 00000001000 andM. 00000000010 ',.,; " '", '. \ 01000000000 ™ nerenf 00000010000 < 00000000100 00000000100 00000000100 00000010000 00100000000 00001001000 00000000100 00000000100 OOOOOIOOOOO OOOOOOOIOOO 00000000100 00010000000 00000000100 00000000100 OOOOOOOIOOO ion (i = 0.34; P = 0.0037). ; = 0.51; P < 0.001). i Macrocai-paea (nodes 6 to 61 \s. the Fig. 2). Clades II a les 1 to 5; Fig. 3; see Table 4 for impact tepuis, and Mesoa values). Allopatric nodes had an average (Fig. 3). SEEVA ar I — Symbotanthus ■ — Tachia guianensis [TooL_| — T. grandiflora so"" — T. occidentalis T. occidentalis Chorisepalum psychotrioides . — Macrocarpaea obtusifolia I — i . — M. glaziovii »*-"< — m. rubra I -M. gondoloides - M. luteynii , — M. valerioi • — M. papulosa - M. macrophylla sect. Tabacifoliae laudialis pachyphylla M. densiflora M. pinetorum • — M. domingensis A — M. thamnoides M. arborescens M. cochabambensis • — M. maguirei """' — M. pachystyla - M. bangiana ■ — M. subsessilis I — M. stenop - M. chthonotropa - M. opulenta : I + J, clade VI). Seven of nine variables show the whole G and in a detailed .' species from a clade that dispeised ;,.,.. tins ,u< >F l-'F' * «» <1 -i- ■> i* tl .*- d.-iin, t ,j. mces m si^ oi the m 1 , 1 1 » ~ ( r il i [ ''i ' I * 1 i U J .: .. . islands, and even 1, f historical sea incursions (Clapperton, 1993). l.i iK He i < i„n «, i, lit. Mi. nl i,i n ,1 , . Mano wpaui md Helieae, but South American continent, e\ohed in situ, and A an area being more dispersed within it. ancestral, has also been found in other groups that we can assume th.tt !' i r.. t n c dispersal from the Andes, ^>]j>[ irllnsn nl u tin- ln|» th. -f-i, l'ii 11 1\ illmi t.»>,u 1 I In i-'iiilii-in p,i t i jtli I he mam noithein : . .. ■ " .'. •. ■'"-.. ■-■,-■ theory (Stmwe et al, 2002; Frasier et al, 2008). u Ghosh et al. (2006) The ilemation of Mesoameiicai, ^.cms ,!', , .ml'* n.-Wi 1 .< 1 i ,2'«.'C, Ihe \ndean forests of ccnpaea aunculata ^ea\ei & J. R. Giant, 1/. tl arpaea occurs have ,,., ...J 1/ ,„1 i u< M.u.11 . fiom I,.. I- >il,M'-ll -i/M't, I i i.t Mm- ene to lower Pliocene ■ " .■I, Jll..' „i.l I/z/c •*,,.., . .1 -in »li. I) .[nul. . tl. \,. ^ .and I.na.Vi, north of Costa Rica, n Based on our DIVA anahsis, the ancestral inhabited yet. distribution within the Andes includes the Amo- of the Greater Antilles: M. dommgensis Vrh. &ELmcai most speiies-m I, a., . , u. th, \r \. «.]><]. k.Ui thanuwules { Gn, e h)<,d. .Tin.. .} j.I ..♦ nwli .1 d •« ntaJ . i ' ■»] mini , „ 1 t, m.l.. F. m.l Hi 1 oi.lill n in this stuch Then ! .<.=.. . .y .pi. > • Uk.iJ.ijv a.. 0- until .1 ( o! >,.!... a., I < ....hli. ,a ,le M.n.l. n rS„ ,,!,,]« „.. ,n.l ...t.-in.,! un.il tl„ ll„l„ ,,, ; t >„.plK ih , Hili,. I.. 1 - .Is, !,a.. in.' 1 i lit i m m il !!!!!!!! : !!!!!!!S ■ ■ ■ Islsllll iiliiiiiiiiiiiilii ■ sii-l-li lllli'liillis'lliil!' lls'MI iiiliiiiililliilili iiiiliii IllllSlli'llliisllls 333333S3 ^^^^^^^^^^ SSEES<<< 1 |^^^^| |^^^|^^| | | | >H S18SKSSS3 — Symbolanthus jasonii - Tachia guianensis • T. grandiflora - T. occidentalis - Chorisepalum psychotrioides AT- Macrocarpaea obtusifolia A, SE Brazil ' ^Silfls" M. glaziovii A, SE Brazil ^ SJ — M. rubra A, SE Brazil - M. gondoloides F, N Ecuador, Cor. Occi. ■ M. luteynii F, W Colombia, Cor. Occi. — " M. valerioi D, Costa Rica ■ M. papulosa G, W Venezuela, Merida ■ '/. macrophylla EF, N Col. & E Panama, Cor. Occi & Central - M. auriculata D, Costa Rica ■ M. subcaudata D, Costa Rica & W Panama " M. glabra G, C Colombia, Cor. Oriental M. neblinae B, S Venezuela, Sierra de la Neblina M. gattaca F, N Ecuador, Cor. Occi. M. nicotianifolia G, C Colombia, Cor. Oriental M. gaudialis E, N Ecuador & S Colombia, Cor. Central ~~ ■ M. pachyphylla E, W Colombia, Cor. Central ■ M. densiflora E, W Colombia, Cor. Central - M. plnetorum C, E Cuba ■ M. domingensis C, Hispaniola " M. thamnoides C, Jamaica ■ M. arborescens H, S Ecuador ■ M. cochabambensis J, W Bolivia ■ M. revoluta I, C Peru OT7 1, C Peru s I, C Peru " M. robin-fosteri I, C Peru ■ M. jactans H, S Ecuador & NW Peru ■ M. tahuantinsuyuana I, C Peru ■ M. cinchonifolia J, W Bolivia ■ M. normae J, S Peru a I, C Peru - M. luna-gentiana H, S Ecuador — ■ M. zophoflora H, NW Peru M. fortisiana J, S Peru " M. maguirei J, S Peru ■ M. pachystyla I, C Peru ■ M. bangiana J, W Bolivia M. subsessilis H, S Ecuador M. stenophylla H, NW Peru ~3S" M. chthonotropa H, NW Peru M. opulenta H, S Ecuador - /'/■ H, NW Peru M. pajonalis I, C Peru M. noctiluca H, S Ecuador & NW Peru M. apparata H, S Ecuador M. e//xH, S Ecuador M. bubops H, S Ecuador & NW Peru M. harlingii H, S Ecuador & NW Peru M. innarrabilis H, S Ecuador & NW Peru M. kuelap H, NW Peru M. micrantha H, S Ecuador & NW Peru M. angelliae H, S Ecuador & NW Peru M. pringleana EH, Ec. & S Col., Cor. Centr. M. sodiroana F, N Ec. & S Col., Cor. Occi. M.jensii H, S Ecuador - Symbolanthus jasonii H ■ Tachia guianensis B T. grandiflora BK -T. occidentalis UK - Chorisepalum psychotrioides B e (3 spp.) A —Macrocarpaea glabra G — M. neblinae B — M. gattaca F -M. gaudialis clade (3 spp.) E -M. nicotianifolia G -M. gondoloides+M. luteynii F —M. valerioiD -M. papulosa G — M. auriculata+M. subcaudata D — M. macrophylla EF ^Caribbean Macrocarpaea clade (3 spp.) C —M. arborescens H — M. cochabambensis J — M revoluta I Geographic A Southeastern Brazil B Pantepui of the Guayana Shield C Greater Antilles of the Caribbean D Mesoamerica (Costa Rica & Panama) E Cord. Central, Colombia & Cord. Oriental in Ecuador F Cordillera Occidental in Colombia & Ecuador, & Choco G Cordillera Oriental in Colombia and Merida in Venezuela H Amotape-Huancabamba zone in S Ecuador & N Peru I Andes in C Peru J Andes in Bolivia & S Peru K Amazon Basin of Brazil, Colombia, Peru & Venezuela Vn-].en.|(2lN,2 JOOlHi.^MllutMieHi^iKmumlM .1 «li,„ isolation Both scenanos suppou thf ■,< d ( u hffeient altitudinal •l'U'hll.,,1.0 1,1^, ill V.1.1. ..,.1 HI Uf. .Ui j t „, , , ,1 ,, I,, J J >-.> more data need to be collected and analyzed (Youi et al., 2002). One major difficulty is the lack of up-t date revisionary, phylogenetic, and georeferenc, only represent a small percentage of Neotropic biodiversity, and there is a dire need for mo ! , ■■..' : rva d 'i I P ' A ( .1 , k k ) i il i _( Oi. K uluilut >l l%S-l')7,i til.,1 „t y 1')... fOjWl'v 1N\ME)vi 2. Published by L i i i bi L i 11 J 4 Press, Cambridge. erret, M., A. Chanter, 127-134. F DF FG Node "W D DF DEF DG Node 39 • !<1 V ,, , ,-',f ,..,„ ». H Node 49 :■ ooc/iys; a: i \or r i ,,,,,-v ...w/. r i JHIJ u T 1. , ,,„'-!<«, 1 U DETERMINANTS AND Trisha Distler,* Peter M. . PREDICTION OF BROAD-SCALE Alan Graham,* Gerrit Davidse," PLANT RICHNESS ACROSS THE WESTERN NEOTROPICS 1 Patterns of spatial \ mation m the numbei of noL.'.h llv sp. - u s ,mm-> pi .. -p ltj .,l 1, were discussed by 8th and 19th The SE, bi s that gradients of .■•■■.,■... ■ ■ 1994; Rohde, 1992; Palmer, 1994; Rosenzweig, on extinction ^ i uhl, t)"„y evolutional % lates ]. n in a o-ii.J HusiM tl J:,(." |...i.> I J J \57( Ihe SH posits i work in this units increases richness, again through \arious 720081 ! s the Western Neotropics among kiS„1at - (li.Mii, „], d, I, -,1 rJ,A 1- (Fig 2 V, B). patterns of species r of relative richness due to ■ ... . ,...: ... Inoad-scale patterns ,1 p!,m1 ■,,>., j,^ n, hi. ,, k< i. diM.U, ,1- ,1 Jj *a,cuhi plants occmung in that the SE, SH, and RE in det* .ression model, «e patterns of vascuL ss across the derived predictions from one or more hypotheses ,,i - M , i> L'^i.. 1 i £ . I=i i,,ir I'll ]<;.,'> Tn ! - .[, 1 m JeK li. ,,), ^.i I d northern Argentina, We mapped these data on 1500 2000 2500 values of regression, f uir i. nN, i), im .I I.n imtmn, s.ntr 1' ,V> I A,h> } 7 t < il. i I n ,1 • , ian, « n« we selected data sources to reflect the IdClim. Principal toi >1 >„o,lels 1 ami J (F i„r» is ,V I unit. 200,,, V<) ' ' « ^ ■■ ■■■',.- ....■.:■• ' . ' ,ii.„ii.in.u.„l'Pi„i» l . 1 iuir n.r.,,) m.-tLt'ih i«.. 1 j..i..n..^ lata ,,n 5 » 5 mm lesolution (Global Soil Data and annual precij p:// Task). based on hypothesized effects on plant iichness of was the Inte second-generation Jetz, 2007), climate ale patterns of woody plant $gg335¥?§888fe88S jJY"7"" n L|"[j||ii| --— — jjj 500 1000 2000 3000 inual precipitation (mm) | m>[TTTr[w^ i ■ Jul resampled to 10 X .1 richness across the western Neotropics (Fig. 2A) • a, ..,! .i.Mt, '..«.. .unl f,i '.,. ati. i in *..-. - il „sp,.~ .,.i.,l,l. rl..t ,s ., t counted for by a • ,. ■ ■ :■ ■ the Neotropics (G ntral Mexico .il \meiira). luenced by otogenic actmh. and In l,.t, ,j,..u..l ,- ; i— 1.», ,.„„!,], thai - -Urn I, ,[ ih „ Uk„i • Models and % LoglO area LoglO area 3.5: 0.01 (2 X lO" 3 )" 0.15 (0.13) -0.01 (3 X 10- 3 )» s the Western Neotropics Model andvanables Pred Re81e nons!atl!l iCient ' Resressi ;;^™ l efficient ' K™™ Is Model 3 SH, R 1 = .oglO area ty , : 1.2 5 32) 1.2 ;■: ( /" 5:!:::;:::;::::::;:: \ ' 0.011 (2 X 10- 3 ) c 0.298 (0.1 l) b Model lad ho. Mi m, il , r;::aZ otrans, ' irati< \ A + _ oc, R 1 = 0.258 (0.300) oc, R 1 = 0.254 (0.300) and variables pred «, Re g ™coe ffi cie„ t , + 16.19 (2.04) c 174.42 (34.07)= models those spatial e: the spatial autocorrelat ix id a spatial and nonspatia 1 reg controversial (Dorman p. -U.Im] predict To predict relative nt nchness i .cross the substa, only sparingly collectec [.we- L. -dill. |,...|l. from the regression rr ..: ■' SU] r data— that is, models t respecthe predictions. to 3 individual hypothesis, differing by > 41 > - .■ .. l« itioi : [( M ' ant a. : e 1). The higher performance of ad hoc models 1 explanatory \anables in om sample of the western However, we used -s (Fig. 3). the IGM2 as a id, therefore, at least Results a single hyp the complementary NONSPWRL MODELS " atUre ° f the SE ^ SH ' All models tl epresenting the RE e.i [li.il ihe -U h.ni u.o,, vitli|.i..| 1 .ti.,ii^...m!,r,lluli. I-E n».i llie HI 1 a. !. .»t ' I l„.ut th.'u m;-ji [-• MHlu'ls* M(hH~, > • ,■! tin-.- models representing the SE representing the RE did best overall, with lower AICc did so (Table 1). The AICc values for SH models 1 v els combining terms . • ■•-■ other models (Table 1). Thus, the relative perfor- led in this study, richness across the study area. substantial Foi exmij.l. , i^h.ial « lwi« to -jyiih. .i.t.,-y. .„..„. .k Hi. i "1 tl. .r «-i. i,. -.U deer, is. : a: tu d I- i < .led that N ( ential \menca compaied RE Melded statistically significant regression coeffi- tl. m tl.- sF \U ll.uv n, ,dek ,~ u( 'enUa; tl,. Ml *, H f „ lK , rj]l |, , i. _»"„,„! ,1 . „ US., .pKs ,«-i. n. t odels2and3, the RE. Nonetheless, a spatial model including mean annual tempe e = .326; Table 2). This model mealed similar ing a .mgle hMiothesi Ml . , ,d, k I t ,n,l 2 , ,(.! , n, d , 1(l , is j,,,],, %J ( , n t, -,1 \mwa .ompaied to ■'•-•■ ■ , results from nonspa uggesting a Fig . 5D _ F ). E.fh ,i tLr far s,m'uI mo ' M..M i,i^.i..i;. •■ ■ ( : ' ) ; (21.6 s the Western Neotropics the western Neotrop s in Peru (Fig. 6C), ilK in- Im I, -, .,.,h I. .,-l, tin lattei model (lid not include era Central, the Andes, and the richness trough in lowland Amazonia relathe to S nalh 1-.....I hmd >., w. ». . , ,• ^t.ino-In.oJ.l . i ui tl„ i^ik..^. uinal.l- (R 1 = 258- 4, which predicted hi !6; Tables 1, 2). -0.121 -0.382 1.000 ge 0.172 -0.061 0.261 -0.563 -0.118 -0.375 0.951 1.000 a -0.095 0.058 0.129 -0.083 -0.013 0.060 0.055 0.033 0.545 1.000 1 0.073 0.020 0.220 -0.444 0.031 -0.261 0.838 0.799 0.161 0.112 1.000 i 0.166 0.149 -0.] Our results suppoitid most a pnon pitdu in s( [n n mi mm 1 n < i ll 1 | i Imli I | oti i -| 1 >u i based on previous studies about ik ' hi,q, , < ' j ] „ir ,,,,,( *q, u i, ,nm il |x nptldii ui uUi ^. •. ■.-. . -i*)' Ii.iim 2iMl' luom^t-el ,] . 2)()' l^J.Ut J rtii.l ^.itlim-, implm. unit \ m n t water deficit and annual me , 2009). This result .U!J i.nt jnlt.-i-h.-i. ,i V ' ,c .egions (Table 2: Fig 5) ,k b,mb i.uv ,i bam D 1 '', 11k- iin.l]..- n- -,rl. i 0i< il, .1 oj I al J)')-, bi 1 I ti al >005 Moser et al., .■ . . R 1 v In.--, mdi. ah I that .\i I <„aloi- \.,n ibl - ,v]>.<"- t )j, ft J.Litio,, of 'hr , -.pU^fon \ariables. , : representing the SH, ,. uaiv ■ j„ vnJ . iJ...., J- n-if, ..„„ ,|,U ,, , ,, M mll! o t |,, -[I Specific alh, the SH ,»! ,hl it i.um lepi. entniatht SE mo,t ,cl I, c u Tib t ll, ,t , t , hi 1 hffuulr I caw f, Ltr fM..l,Ud ll M ,1 lt h, ,, lUib k A (,n - 3-(l »a, hnJiu? t Finally, \ariables representing the RE sometimes al. Haskell, 2003). Central America than for other regions, a higher ] >rholm et al, 2005), .,.•■..••:■ ■ •. and eneig^ a^ailabibl, im >ih ,1 . „ poll ,,i- { )i. .h.„j in. ohhll. i a! MjH,K ■] ,1, .|.1M. ll «Hlii,», l.„-.II. H. 1 ., -,, t ,1.1 LM ■imlLnlcsh o 1 I? § | | " 50 100 150 iw .lull u.isnti, ll-. -II trh .5, ! w' JIK)', k, it \ I »/ _*(»)-, v tfl *1 ]- uu.i tU le ;pe :s ,* . .,, ul, nori, , 1 .,ti p.. 10, -t, U is, lu.uluilm w, 1 , ll! .. I s in Another important difference is that between li ting the SE and SH. ■ an rarefaction as an attempt to correct I e of the SE and SH. In tropical and ■.:■■■■.■ . • hill. )1 (1. 1111H..1 I. Mill,,..l Ur.itll,, hu'* \ It/ _'0(7i Ll [,i >h h t .h.l p,«, t,,l le to imagine monthly p< liration (Tables 1, 2; . ...■ : .- '.. ji. j iium - .t I. th rtoofh and noiraooch life 2Q<> I ih . u ,i ,. .i.l » l t,.,n His „ 1 I1„ t. ,t ! for the SE would still be valid for this area. ■■■.-.. \~ ■ America, consistent ,ed on our sample are more n- II l\,.M- l<>>, P.lhJs 2001 .l„l,..l . Li. i<-.< •>! th< li M2 suggesting that the ki it.vjtt .007) Identirungthcuiid.iluug i ~. ■, in, t ,,.<-. < f ll, 1 Mntl , i 1 I , h. u t .. < icl ti lo.hn.M.. . l,,,,,™,!,-**,^ v,Mil lrtl ln oi an-nal j.i, < j|unh..n 1 .i ,li. U^uUi nJo.aH, other regions ma) be due to limited southward The foregc een the maps of movement of Laurasian clades that are moie mipoitant piedicted plant lirhu, -, (I i^ 2< , , , ' components of montane than lowland Ne, ive importance of * t.ru. ,n.i ,-,.,, i Ini nil ' Miiilii.n.iiniiu, Ihjs.u k^si'i. is^U'is I JHM and anothei lecent mapping of annual precipitation for minimum monthly potential effort (Kreft & Jetz, 2007: fig 3b), but contrasting with etd|.c>'i, 1 i..pi.i'j..ii Jt hi^mm-s .he amount of annual : ■•■-:■ , , (Fig. 7E) because, as pointed out previousl} to the IGM2 the relationship between minimi piration increases, tl IRS of annual of the logarithm of elevation range relative to ,„ tl .„ ■ >. . Lt w , T r J I , .s.yletV J., P ,. lg combinations *™ W " Barthlott, W., J. Mutke, D. ] ;y 60: 611-617. •ill. .,, lt , „1 .h II-N2 ,f.. 7F) I'.- (.. Ii„. "»■""'. 'TIM \i.P'„ : Annals of the Missouri Botanical Garden Foitin. M. J. & M. I Analysis for 1107-1116. "! iai' _, !.,-;•» ,,: . . . . . • -. d un pi ' i P. 200: 1i. dia on I • 1 ;;;. r>o:-<,H;;. Moser, D., S. Dullinger, T F ,. L- h H ^ .Uf LI '- Pint I M I m> , 1 I T i „ , 1 M r i «' L M * J "!' ] * H P '" '' " '" , '' " ' "" ' H Nelson, B. W., C. A. Ferreira, M. F. 3105-3117. . U.S.A. 104: 13384- 13389. 600. Press, Cambridge. " ' H. P. Jebb J.tz. S ., C. liahbrk & ti. K. C.,:\v.; II. 200-1 The cnm_i.l f -:ice studies in Thailand. J. Bio», <.;,i' HO. I";u200. i.t ill. 1 ■, L . .. 1 ^ i - (H Tn I 1. 1 ,1 H , h ANDEAN LAND USE AND Kenneth r. Y ouni BIODIVERSITY: HUMANIZED LANDSCAPES IN A TIME OF CHANGE •i ..dine plants and ammak lan.K. q« . .,,)i. .11 U>„ n jn ibis , ont.-M u, m4m X. ..j. and on the living Ian face tens to several hundreds of important to evaluate. For example, it matters greath ij ( h nir-i, 2005). This article (At,,, « k.Uii-toK .1 I,,', .m.ljonn, ,H,t ] ,rj i„ J ui.k .,].. - ,.- ,i , ku u.u s< ,k' rt .lhvh, h ..,Uu^. , :. ■;...:....-, • t. mi.-u,ik. . \.,aUJ.r\ .,i.l a - 1 t ,.,i...li<nt!i. .l^.H.i'^.d.^h.v^ > < .. ■ < . ijj e<«-J.i>\ J-iiiiKi> PL.il V ^f. ,11 d h.lm ]-..]i n Hi, , >Mdi ,dd i, I ill I t m , .1 , , i-hdia™ .. .U.M1MH..I <2 i" f. S7J -,,. u-,l,0L" 000 ,e..-> which affect the de ? and amount o s such, there is a temporal scale t er.iJ _'. .tuns .,1 ,<„ik ki.i-U r.l .W-.h — J,„ n, ,„ple, as measured in ,. at. r!-i„,'pli.'].. ■ .«, ., 1 ..l'u-,!s li/wna , Lit, d ,,!..,< .l„i : were crossed and at that feeds Dilleha. et al. also showed culthation of squash ■ .. . . . . . ■ ; , . . . , . .land,. onti.lL.1,,^11 lt .i..l,.i.u, land covei, accompanied l.wli. d>n,e,h, ,1km. tu.l.l (Brush, 2000; Young, 2002). Andes (Denev j et al. (2003) found were based on cons i ty et al. (Chepstow-Lusty & • ,,. ." .-, views originating in tl nee of ancient agroforestn . tud th. ag ■ .: : were deforested for timber and agriculture se\eral c 2001, 2003; Young, ili.it/, , 20! •-, T,,ili, fJ ~, fti, \,,d< m.Kl.f'un ,,.„ !,..!,«,. back to land use dm e comdois, can be tied conceptualh to theones and imbrine et al., data, in this particular case, on economics and on -i< Intinib ,n ! lh. l..„, ^ lo I mot i i ', ijmj.kI M,M i. -, . a i, l.ril . lb., I t„ 11 «,,)0-7(.r>0 x ,is BT', th .,„,.,. al |j. 1. .In,, mg pkmled and fallowed tiel.h - landscapes of south ... [ 11 ,1 1,1,11. t i i- i .1, ' u s, lt „i u iSi-t Mi uc tm al features seveial millennia ago People "aimed m Nn.lli .',.1 I U mtensne localh a/ound houses and c Men America in the Early Ho? *tain peaks. ,-m.oi 'e I mi ail r, ,~, l,,.c ipu j, Km. bid I U -i ,.«^ "hen hate small. uell-Jispeise.l seeds, tan al, 2003). Ho«e\ei, theie is an oln/ous human u imprint on the abund.ue. >* d „,Jiii „ \«1 >n lx > oi /it, ,, ,d K, .In, ,20„1 Joi , period ca. 3500 years BP when charcoal d/amaticalh fam ledijob ,v < , < , ,o 1 mil* ^ .*hc <'m_<-.. -,,..K-,s , .l.i «ill u-mv, Jjitsin („.,, ,*.(,„„ -,U v .,. luil lo-t , nsi.li^l.U aaop^ densm, thus or the population . ' m, t , pu u ,>,-,» .1] ' . u- I- urn.. uitl . h, ■ .1 . ,^ ill-i-.M.i' .m. (I. M .. a (Fi. '» ,N..i- tinl 1. . ome smallei > ,!>U.m e A 2>>()2 Staizomski oveiall, but in the i > tlu Vndes, the^ fen , v i.J.l i. d ,o ! (Ui v .1 t< n unit species to occupy ! . -0(1- . - ..iMtt, „J U, nn'.i.nve .,, ,„,, - efjert- of land use in while those dependent on humid microsites (Killeen et the dominant pait of the Ln.K j ■■■■'■ ,,•....... other very humid hal bly embedded s (Fig. 3). :■■■.■ Me 1(l . „nl,i h, . ...liu.L',1 m ,. hliw to h„ _„„ lp - hunting Still othets m.n clmntic conditions rounds et J cO't L ,,111 tl. , < ,< ,.1, .1 in , hnFigr.it -> implies that different •..-on I- Liu.] d..,n -Li.te.u-.,,. roo'-.M,, p-.n- ih..t ..ie p.edicled Lo baled by habilal fragmentation. sensitive, or fo imited in distribution likely Lo be found in more siles and/or in siles p, d nalural land cover .-.:■.'..• :' expand Lheir dislrib; pecies will be use of heav 5 -nt Look such as Lion of conservation % (e.g., Odion & Sarr, 2007), while Ltd- f>. in. til ■ ( „','!i 1 lH.u.iii. i- i- I u. .Jlgiass pi tints of 1 Park. The presence of a conservation- stand of 5-m circun nondii Harms Nat: : , ■ • reevaluate ., -hilt..,. I ,U .siln . .»,1 „lll ,! .,_ ,1 [, , r i. ,K I, * Kort periods. . M. 2 I , Fortin, H. L. Beyer, D. 1 idscape Seabloom, E. W., A. P. Dobson & D. M. Stoms. 2002. , i I. , turn fll, .] '.'. r- ,1 Rand, T. A., J. M. Tylianakis & T. Tscharnti " P ° nSe t0 Redford, K. H. 1992 Reusch, T. B. H. & T. M. T. Sykes & I. C. 1 ' j,l -,i ll> » ]ll! J I] .-I UV . .. r U i I h ^ In, In. ^ -t«i iUu. - H m i, :. Ecol. Applic. 15: 493-506. Bioduns ,i Cons umill. ess. Geogr. 58: 436-447. > il. ., ' HI Cairns & H P Po,sm»hai , . 855-863. APPLICATION OF SCIENCE TO C nd Gustavo Kattai PROTECTED AREA MANAGEMENT: OVERCOMING THE BARRIERS 1 foimsofljle let, the Inoken lines of conmmmcali, n To m >i t,m snenre into the transfer this know! ; and conser- decision-making process. Here we will focus on what : Murcia & Kattan Application of Science to Protected Area Management ■■■■■■.. ■ . . . :■■■.-:.■ ■...-.-■■■..■ by external factors, itional component of ;.■,.,-. : ... . ■■■■:■ similar for the social sciences. gap dynamic in sufficient detail, and across ( is and spatial and IlNtOHMUlOlN NLLDS ()t MUKGLRs ,!,.<■ -till .|jph. .>!■..!, ,1 ,ln s, mmiI ui, n.i -ju, ul ■<"- i-t. 50 ^ 3 & 0sterta g> 20 °2; •■■;. : . .... demised using localh pioduced mioimation, but few tem, ,n . „V. ,i , V, li ., i,~ , V -. . t -j.-ii .1 ,» L H „„ One option is to ext. mal stages? What are the harriers to Many conservation problems require long-temi or th. .-..:' ■:.:. ■ a long-term researd riple, Brazil, a Funds can come as giants horn pin ate donois 01 Factors That Affect the Generation of win be adv£ l0sals that address Applied Research cutting-edge i: frontiers of science aie moie successful; however, Colciencias, the ■■■.■■■..■■:. and applied (Fig. 1). On the scientist's side, the make it hard for scientists to sustain long-term ,.lie.. tin, re,,aKn i, puLlj-l,. .1 .,,• ih vancement, and th, nding cycle. papers. On the mar t A more recent source of funding that has the decision- aie- the ]», . .!! i ^ 1 1 „[ ' i (.<< -„. 1) ih, [Ktirl .ppfitu.LU 111. « ul.lil.h of a peinutfiee m.,1 s ,[ h\h nl ., >1, u tl,.s ,, „' m „le.l n u., ili^iecl In- - "»"...! ii..,J«m-iiti.is. .«««:. s ■,„, ..fUitional levels ary, multicultural « iibbons et al., 1994). In this s a in. U ninltidisriphnan appioich thai th it iniuiii/Hl <|i l U. n i-l >' the i< suits and the tific community at their own forums (Ruth et al, a »06; Aarssen et al., jm "'IijI t ikIii = uli, m .Ji^IiiikI ii.iRs.Fklu 20' >i, Olden 2007). While • ...iHHis.ionng «..iHi..jii scientists on specific topics. n<- ' -:■- : ' e research. Murcia & Kattan Application of Science to Protected Area Management r<. „.e.<.rl. anessiWe lo scienlifi, hteutuie piocW.I m In,! ,..<■-]« -m. - manageis. Howe\ei, this is often an opp, 81 from the gra^ activity and not part of a concerted effort to i I onh four articles «ere in xation practitioner- ition with the Dutch, Frenc leijaard & Scheil, scientist ends at the end of the talk. 2007). In time iing should make the mam of these issues theie m.n l>t siij(i<»nt ..-.u- !i ^,ts„ii«»j ,k H I.. ^Ip.-t 11 lo I Ik- I.hjI ron.lih.-n-, u.jun, tun. t-.i a 1,1k. Jit!-, p-.re ,tnll, In „h, T,,,, „„ t h I,-,, IrU pli-s,, .1 j |„t, ( n ,,.,1 , L,. !„,„s ,. jv\ -I , >»>u , , , ,-> .- tli .In., i I ,uo,m | -mil .< I l„t,t. I.eai 1,1 ' m< ,th L.ih i - u- ,-a, Ik I li.trtl •, n. tK t.l, . I, I mm. tl, n. M.m.l f mi i , th s in .... .ate m then «oi , 1 , mmnm.h K, >d,l.. - s,,ot ifi. makers, and at the s £ ning and action. NGOs to address specific conservation or jment issues. reach. Scientists oft* t that they are which they need information. not sufficiently reac nips to ensure 6. Universj graduate students policy makers, perS differences in to find, read, proces> e literature to apply it to specific circumstances. Even in countries In 2000, the Colombian National Parks Unit plans (Pullin et al., 2004). process was -ial sciences). This We propose that there is a need for a science s phases: a diagnostic The Need for a I lh ,V tl " ^nr'zihefih 1 ::. Imxi? Is „ the sibility to do sue] 1; ;o roups a Murcia & Kattan Application of Science to Protected Area Management ■ ■ . , .... . :-..... ■.. analyzed. Of these e As part of this activity, 40 Colombian park h.uiti. i ti ih s inl - hlWVI'p ih,.t ,,,,.11 uit.i.suf ,tll. im, !.!•).. . Ih< i mmoi nop- (mo- Murcia & Kattan Application of Science to Protected Area Management militants) and m» ' , 1. so, ul ,!.,,! , oliti, _,i a.e, ,;-. called the Eje Cafetero, land use in the lower p in their areas (under >.,. 'nng constitute tin agnciillm 1 It i.utiui r nsn mention and the -JNA.P), and lorn-. .„. the ,i, .i,..L. ,,...!. n.„, following: 305, 2006). R principl. esilience, redun- and current vegetati lus L.), now have management 4. Updating of . established protected are 1™ " h U U!t Kljl '"' THE R ° LE ° F The ci in Colombia is CARDER) have commissi t, J r : l ;|;;;j;, ';;, a synerj : management plans. j. banefn tU m imi nt and NGOs. poralioi oJ i hav. consenation biology concepts .' land an. This ' ]h,h,d llnon.l ,- .....1 wo. ,inl,.-i -t hi. hi, in. .•IV far. -r,, Iu.xp ing on satellite ima; sification. A key elem. is the emergence of network of protectee -em portion of several of the , academics when it [ ..\ulnn-Vl- ,1^1 I tin pif<< - lu pi. i,.)ii]o l ln\< ni.i. U< doi.it choos. their research questions Case Study 3: Consortiums of Organizations That S serve those ne. immitted in the long- A third model that has pimed successful in te ows them to collect n ,i n.1 Kn V (V ,„!,,,! p „ii. h, , , 'v U, I, j |„| ,,i m inn. Murcia & Kattan Application of Science to Protected Area Management rtail and potentially n 3es. This huge task cai , donors, the need for profe; neral public— 196-201. join forces to ensure information is i bold ways. Wled.e Contempo mechanisms in place that al 1 Open Ecol. J. 1: 14-19. ike.,,.,. J B <;, , , l„,.f,ii , j nil , »hsl,i._., in- ,. I- . la 1 Iim 15: 189-211. Olden, J. D. 2007. B :Io eoologi •70~'~. Sergeant. 2006. Bridging www. mbgpr ess . org CONTENTS Biodiversity and Conservation in the Andes: Introduction Peter M0ller j0rgensen The Andes: A Geological Overview from a Biological Perspective Alan Graham Climate in the Dn Central Vndes ovei Uolotuc. Mill, mnal and Int. i mnual Tiniest ales Christa Ptaczek, Jay Quade, Julio L Betancourt, P. Jonathan Patchett, Jason A. Recti, Claudia Latorre, Art Matmon, Camille Holmgren & Nathan B. En t Diversification of the South Wiicau Uitauna: Patterns and Implications for Conserve in the Andes Jon Fjeldsd & Martin Ire: Pli , •■ VI B Moss Diversity a Andean Speciati n and Vic, __ Jason T. Weir 410 •en P. Churchill 434 ropical Macrocarpaea (Gentianaceae-Helieae) Lena Struwe, Scott Ha v Jason R. Grar, Determinants and Prediction of Broad-scale Plant Richness across the Western Neotropics Trisha Distler, Peter M. j0rgensen, Alan Graham, Gerrit Davidse & hdn Jimenez 470 Andean Land Use and Biodiversity: Humanized Landscapes in a Time of Change _Kenneth R. Young 492 Application of Science to Protected Area Management: Overcoming the Barriers Carolina Murcin & Gustavo Ka.ttan 508 Annals of the Missouri Botanical Garden 2009 * Volume 96 Number 4 The Annals, published quarterly, contains papers, primarily in systematic l)oLany. ; ■ • [he ! ; den dependent revi the back of die last issue of each volume and are also available online at www.mbgpress.org. Editorial Co C. Hollowed " Editor, otanical Garden Beth Parada ig Editor, Allison M. Brock e Editor, otanical Garden Tammy Charron \ssistant, Missouri Botani Cirri Moran Press Coordinator, Roy E. Cereau Latin Editor, oldblatt cPherson For subscription information contact ArxNU.s Oh the Mtssoi in Borwicu, G aiidi^. % Alien Mar- keting & Management. P.O. Box 1897, Lawrence. 8897. Subscription price for 2009 is $175 per volume U.S., $185 Canada & Mexico. $210 all other countries, four issues per volume. Tl,c journal Novo,, is included in the subscription The . ' Rota G\i \l,s,oun liotuin.al (,ji.ln,. 2.515 I owe, (,,„u- Avcnuc, St. Louis, MO 63110. Periodicals post- age paid at St. Louis, MO and additional mail- ing offices. Postmaster: Send address changes to Ar\\\i.s oi' the Missoi m Botamcm O \uni.\. % Allen Marketing & Management, P.O. Box 1897, Lawrence, KS 66044-8897. stract/Global Health databases, ingenta. I liesearcl, Alert®, and Sei Search®. The full-text of Annals of the Missouri Botanical Garden is available online though BioOne™ (blip:// www.bioone.org). © Missouri Botanical Garden Press 2009 The mission of the Missouri Botanical Garden is to discover' and share knowledge about plants and their environment, in order to preserve and enrich life. © This paper meets the requirements of ANSI/NISO Z39.48- 1992 (Permanence of Paper). Volume 96 Annals ^j||k Number 4 of the ^^ 2009 Missouri Botanical Garden A REVISION OF THE MALAGASY Wendy L. Apphqm ENDEMIC HELMIOPSIS (MALVACEAE S.L.) 1 ha\e affinities to geneia such as DombeMi Ca\. 01 collections a, others. Many more recognized; one spec lular petals. The latte M..l> «-..,' . Vi . (lie lemaim.n. -pecie- Taxonomic History broadly distributed. with a single specie rrier, and two named varieties (not I ue to the lack Ini.ui.,,,,]^] ■ , « ,n..l.,m. .JNm^iirUu. [I< Ll\ T „ niUiii.,,^ V'^a VI',', |, t , , H.c '-,■■:■■■ :■ ... ,.,',, ' , ■ I d.SMlbM _, ,, , b<>. .... m. k -n n -...1,1 ...m - b.tuM.1, ( l„t ojaulll]al j )t ,l, ^,I.-,,4(„ /.V. ae i- plated within th '- ' .Kt'.i(i»,t , 1 h iMimi v .1, , ',> t ,,,/H..l>, h , ...mdei )i.,liKlilia\t aiimih In Tilnce.e », , li'o ! L<> 1 ! '', ' \ '/ " \«' I , ,." ' i.ll J...,"!,,".' s.l. (Bayer et al, 1999). 1 , P i. „,il<>5>' i.. i'ni ii/'i t -«, / t '; il'» .„ |, KU i es (Dorr, 2001). , ihi fi'is 1>.) Id 1. pl.t 1 itlim // /? ; t Ms , s , T]e | ,| u \,enes is bema l rpels and the 'i-.-l, .J.i-kIl'I... petals aie otheraise t ,r M n. ,.1 ot (he kill, i ^-w>- lh.mtw\.i =ns (15 to 30), and nonglandulai petals, «lule in li.tliun 7 - L l,u ,K (Cull t( ,. S 'iuMie\ Irenes ^ <>M mL< "' iarecliaiaUenzt.lUl.no].. .i-.n-all 10 ■ p. >t ,1 t., be t „ - o- I >,< ,k.>l--n. M ( w< n .n L1 ,,i,,lepidote.andaWs,aprdh radi.r oi.- ,i H vi -mh fjan-ini.,1 as.,;,- i. „i.„n aiiuiu th- 1 „,...! ■■'.■••.:.. v.lj.v, .liiuilV l.-iuili md with petioles fit quent »„,, II .fin /;_>,. „ i',.,l! (Vogel, 2000). n -. iltlmL si. -.>..! -;.' . ill.- sometimes l^e a :., : with subterete pedi dated at the extent and visibility Lens, as do mam other plants. <,<„! ',!,, >,'< mil O). t.tllr s,, clr-^. t . 1< , ... t| ,.|t,,o',, „ I ] ,1 1 1 1 10 the locules of the former are internally glabious, ti,e, m-ei it. » 1 In « U - „ < J c aid J) Imtanjoha ■ : ■ ■ . 6ozr»izz (Baill.) A , may prove to be embedded within Ovules and seeds of Helmiopsis s. str. usually bear an soei -,1 ap,.ii« n> i »pl . I . ■ il .1 . . 1 < two per locule; if two. one is .rding to Arenes) is whether it shall be carpel number to usually three is certainly an ,- ,i-'e> -„< whether these spe, dhision of the genus at the subgeneric Wl is th-i than Domheya. Winged ovules were ters. Nine species and one subspecies are recognized, but it is not clear tl 1 described. les small, ± lane ace usually lepi adaxial surface kl : until .IK lat.-ral. . atidcl .,..1,1 ,. seeds 1 .r thar IS, Will •' all .mat] lei ,. i of the :, where subteiete, articulated at the point of insertion of tl lelmwpsis are native free except at extrei le, lepidote, basalt. Annals of the Missouri Botanical Garden I. H,h . , * Discussion. Helmiopsis gh 1- , , M-l.ohhiii.IUIi I V^.r-. ..s'B.j, .,;" ,Kjll ;B,i,li.a,"?>V;;r/'S'iJi !\jen« .-v>. f ] i -.1 Adansonia 10: 109-] ypes, G not seen, P!). A 1 r / { ) ) /' i : / / / - I ' - ^^ - 25w 1944, syn. nov. TYPE: Mada»ascai Mahajanga b as al veins 3(to 5), the lateral weak; base rounded to i i P!, TAN!). Shrub to 4 m high or tree to 10 m high, 20 cm young leaves, sparsely so in older leaves, the adaxial diam.; bark gra) ish, sometimes splotched, « ith pink- surface sometimes becoming glabrous; scales of leaves ovate to elliptical, bi ceolate, (2.5-) caducous and * Inflorescences up to 3.6-7.3(-8.4) X (1.4-)1.7-3.8(-4.5) cm, frequently 4(to 5)-flowei elliform, or genii- 1 - i , ....ihi-.o.L, J-i.i fro -rsa > ■ a H. Pe rier wei <: * rib d at other [ I lo- , ,1 to)4 to 5, opp 2 J-1..-..-4.1) n e the lepidote, pale brown to blackisl ften ttlut.^iubes'cpni ' ], n's 'in I'm ' *' ' " ' ! '* tlv ' x > > > 1< b i , .]< i ll, ,,.,,.„. ,..,. d size. Therefore, there seems to 1 eeds (2.5-)3.4- !«,, il r ' II hi it, i i on -» i mdiili il" .1 T i . 1 1U 'I (10,1 | hs ,l| « ■*> , , I J . V, a- fnuhng mn occur at almost am nil « i V/ a Zh - «nLn IViun *, . . ,m ., at P. f, .,„ !.[■,.. Hmik ..II .hlnhlnn , i\m.' m-N.wiiH ! M ,|u,. •„! . n.l .Li< iJhK l',J! I is clearh nui enes (1956b, 1959) f('. y;v';,.,'.,'. v »" P. . isj.n .1 UK N R. 1 ])lUM(||)]1 ,. |k| I, t , ,> ... t| lt .( „^t!„ , p^m.e. (LC). has. Common names and uses. Mainah (Reserves ^ ™ letter condition, with buds as we! Z2666); Maintinaty (Sendee Forester 147!]%), \morm argument o\ei whether the specimen v (Sereice Forester e considered origin 25782). The wood has been used in construction Rf> Jt , st , ntutut , s edmem ewmined y (Service Forestier 12666). Orthography. Under Article 60.11 of the ICRN ^ (McNeill et al, 2006 ing ("boh™") ot tl» . [,ul < t .1 h / > o w, •<> i,, is tuated as an tin. [15 ?]'S (Wo ,.| i] m Nh -1, -\i i I r te i. |>' « ' .. )ljl' 1> >)< t( | d ]e ^lo^sca'les 6 thJ stellate "Xescenc-ris s larse af surface. Arenes .: 91: 230. 1944), m 4 ™' which was no as the description i only in French, with no Latin diagnosis, B/flMo/pV'"" '°~' which the IC axa published after ■6°29'E], v. Hist. Nat., ser. 2, 28: 417. 195 "\ .s-,.t ;,es, P[2]!l a „, b . „ . „,«„. »,,,, _d ^ ,»,« « ^f p " » 23 <"•"- * -'-•• grayish, shallow h le lenticels; '' ; ' ,,'li.,,,! I.p,.,,!, ummIK u.lh.n, Intlo,. - , „. , , caducous Infloiesren, , , tn, ll, , f .d i ,11,! pl «,l.,!,l m<- h -..m nm . itli iu iueial and ps« u.loteimiml chisteud at twig -u ,K nu«, h 1m, ,1,1, I !-!>_• nun l- t ..d-, U )■ f, . H ,» . «• t 1 i.muA 1,,, 14-17 X 9-11 n ,pex; wi ' , e%,l!,..,.lqu,l,i, lln-.tul, 4, flowering in January 1900. /{/GV figrf ^ ca ^ Qry Provisional ITJCN Red TiMi,.!. I, i, ii 1 |, .1 h lifted , ,1 k »l -1 i I 1 -,, „ltl, ., I occupancy is likely collections is n, of rarih. to be ^er^ small. Se\eral remaining patches of ,e< il ■ ■■ m ■ s ca. 15, in groups of hei ' ;a r, 2.5-3.1 mm ; .tamii), es are ere. 4, IlrlmiopM* hih We- Bull Mus Natl Hist \n hum' oli'« t (-> i> , 14294) Menarn Toliara: Betsipotika, Analaiva, Morondava . ..... ■ 6 m high; twig bai nent is unwaiTanted, as the peduncle (l-)2-ll mm; pedicels angular, articulated ^6- TongT natn , to ' !, Vi. >'"[>, lanceolate, 7-9.2 )ase of ada- Tree to 10 m high, or occasionally shmb to 6 m . ■ . ■ ■■:../.■■ i.-i.m », MiLiiw.,! |h» 1); matme fiuit- ectedi] ov : .'. [„,-|. ,4_,7_]ul , v n-ted as I) ,,,, ! for entire length. Discussion. Helmiopsis hily subsp. boinensis can uished from H. hily subsp. hily by its larger . to at least 20 m), slighth broader leases v on short shoots, larger : 600-850 m altitude y occurs from species. A si{ ndemic species are fruit in May. Cavaco (Lowry et al., 2000), Combretum boinensis eq., sp. nov. TYPE: [22°46'S, '044°42'E], °sZZ FoieZ, 14294 (P.°T\N). \ohemai [Ihaiana, 13 21'30"S, 050°00'30"E]), (fl.), Service Forestier 27532 >v>\ TiPE- Madigas.ai Mali, janga: foret de Bekapika sur le plateau d'Ante nimena (Boina) [16 28'S, 046°11'E], 12-1 Nov. 1957 (fr.), Service Forestier 18423 (holotypf ventral surface, with a tapering apical wing ca. ■ 5.1 mm. IUCX Red List category Provisional IUCN Rec Specimens examined M\D\G\SCAR. Toliara: foret de pse boivinii (Baill.) A 1 ^:i^ A] :;l^:. i.,.H»..l„t.. ... L,i 1 (l.l-)1.4-2.5(-3. • bark reddish bi ,g-elliptica'l to ellip , (2.2-)3.2-6.5(-9i le 4-10(-13) mm, d. 5), the lateral weak; Tree to 20 m hi § short shoots ca. 1 ( 6.4 cm X 12-21 m r,l,l„J.S >- id, ovate, 1.3-1.7(-2.4 < r . , Jl-J - mm, „ iJ» lI.lllN Mil *«: ^ M 11 - | si I , ! 2< 1.1, Mil ' I Di^ on p'-en- ' j & >,;.! i rhi ,.JI. t M l ( : . It s rollert il ... f I dr. h. •„ .,-, .,!,! Iu.il that a . '• /:■■..■ . . ' . whereas H. hily is < species. pairs, ± oppositipetalous; fila idaxial surfaces; it. p-,ii..-i> aie ((>-«'.- loivinii has only (15 cule, only 1 well deve] maturity, are' much e Discussion. Arenes (1956b, 1959) did not specify pohandra, it is placed therein. II. Hclmiopsis sod. (, Bull. Mus. Natl. Hist. Nat., ser. 2, 28: 416. repar 1956. TYPE: Hel dus (Baill.) Capmon e\ Aienes, designated heie [= Dom- opulus Baill.]. Trees, sometimes large, or large shrubs; twig bark I In,,,-], ,.,„, M , t . „s. .11, K,ll..t o. with up to sexeial oideis of bianrhmg oi genuiul, 0. H* Immpsis l.emieri (Baill.) \ienes. Bull Mus lanceolate, lepidote. normalh I w-.u . - , . . ' • \ uous glandular patch he adaxial seen, G not seen, P!). s' >l. il-.U.I.l. l/t )• jMl-H^tltt! stlj.LUs .. 1 '.- u)2 ' '. ,', I v , ', , " j, ,; , (M() j, 3 mm, scah to lanceolate, usualh immediateh apices on sho.t thick ,,,l k ^-A 1-1 .'..u;. ^ ^ ■ " ' 2.28:416 1056 Basiomm s,„ ,tn,ne~ M.l.tunl.-.l I, .null J. . o, mdiu.mal /' •'-< /".WW- B.„il.. ... >,,„AwV,<- u [12°26'S, ;-6.8) mm, inserted idly caducous; P! )- l.eaung small patches .,t ; ;l.. „l,da. Ii^nr- at Imsc „j tv.iv, p.,K h„d, te It a,^i„o..dh yai> 1-2 mm fertile st mm,, L< p . si.i . t .1 u m j ut ,» , , , [ t« . , i ., II , . i .1 I ,->m -,l l„tl «.b,. hi , j intdnt, ,,1,1,,) >n un , ,,t lml„ s.„ ( . m , ah lobed with a short apical nipple, lepidote; young seeds tei hi ^i ,. psen , ,t llmll I , |,m ^,ed ntai distal ends ■enology, and habitat. bermen is confined w .i up mh ill ,-l.i ,.,.,) ■,». nn.l. ,n ^- ,ul.., immediateh caducous; inflorescences and pel lepidote. Sepals lanceolate, 5-7 mm, lepidott patches inside at base; petals creamy or white .ecial corona 1.1-2 mm; fertile stamen V f V ) * + ^ cv"* 4, U> i Madagascar (• = H. rocks. Most flowerii recorded from May, and mature >nia 10: 108. 1871. Domieyo .. Ijh.i P..u- I "rn jur TMT Madag .nana: Nossi-Be [Nosy IUCN Rod !,«„,.<, P.,„,nlll.NH 1 BeH17\04S15»E| ,ll / U , /,„/„ < list category (2001) is noted as Leas, Concern (LC). ^ ^^ p , ^^ p!) Common names. Hafomena (Service Forestier Treg (o 1 u DBH; (wig bark 9389) - grayish to rede ly ridged, with small ■:■'.■'■ Article 60.9 of the ICBN (McNeill etal., 2006: cf. Ex. (-8.3) cm, denseh lepidot, I, - .1 public tion) Hon.u t. I ' n i ] f r mini li.il . . ii . » L in,' nces omo.e «ith -,.., . . : ■■■■' ■ • required that new ished after 1 12 cm; bracts .us; pedicels (3-)9- !.«mu„ M .l if e„t publication Capuion's combina- mse.t-l w II Ukn il . . , aim.t.lnt. K . a-U tion «as therefore not \ahdh published; it «as n ote. Sepals narrowly validated by Arenes (1956). lanceolate, (4 pidote, yellow-green with brownisl at maturity, usually bearing small nside at base; petals ., TAN)! 11-13(-15) - (5->0-,'i-J>> ( iu.i., Ik .'l, .!,nM, <>» , |„ llp „t w ,nni^L nl ,w\>,c,] „„ 1 I. HM.I.iai h - ue peme, lepidote on . Hu-aered, 1-1.8 mm. Infloies- bieadth ot the leave- m H. psendop.v ('„■, not t ,.lr„K-.1 «iti> it] ..,t s, ,1U> ^ s ,.,,!.!, T . L. n , i / , , > , „- , ,,»,,, ricWii and , have both been collected iron egion, and the two specie, aie clejih 'J.--I. ul.ud H hind- ,K n Nomenclature and olograph. As for /fefrm^ Forestier 14883). pseudopopulus. presents the small. -< h, Jc oi r'„ t >'" ' l •' '" s ,lUi( »" > )h ? ' ' '" ■II., I.V .,IM Leav !.: ' able. ulopopul as have I.' I I.UI- ' 1 ]'. 'I 1 I i • I i (MO [2], P). tions are f rom 5> one noted t0 be in . Helmiopsis sphaerocarpa L. C. Barnett, Ann. dl 7 decidu °us forest, and , .,sn-s .lr I., ijm.u "es r,I,l,s e. Hp la >T ""'"-' "^ ^c -.iJ.er 7 May 1970 IUCN Red List category . Provisional IUCN Red Tree to at least 14 m high, 11 cm diam., or large Helmiopsis from the same region hme been more J1CBBCU , W1IWCBB ccjn ^ ^»n B p^» UB Mcl . fl Annals of the Missouri Botanical Garden Forestier 22669 (MO [2], P). This collection, a white-flowered tree 15-18 m Malvaceae' 3, 23: 151-153. O", 1 ' Lowry, P. P. II, T. H.v ,,-un .,.> I -Y I .Vt. i" J.-F. Leroy & A.-E. Wolf. 2000. Endemic families o A REVISION OF NEOTROPICAL Jason R. Grant BONYUNIA (LOGANIACEAE: ANTONIEAE) 1 l »n li ( u i. I- !» 'wu> / I,.- ,s ,1 ,11, lm,nl • U Mtl, tie 13 genera: Aretoraa Pohl, Bonyunia M. R S t I hill V-< ,./ L , V , u-iPli nu bm<, lmalh iesulte.1 m the* need loi /full taxononuc BxUiul i .1 _><>«» \„ u.j „,! (mi nit ,,,, A ,,„„., I!" »a<-sh I iiU.il an.! J N.l ! u.J . f i - .i s < ,1,, f , If ., B . ]1 i.l Roll,,. T«o other species I, .nil I' (ID l< I \...li. . Kh.-.!., di > n,„ .k : elands of the Orinoco Rher watershed in gei ia Aubl. with nine myana region in Venezuela and Guyana. The g w endemics, each ,'.! tl. t . i<. . tll.l. .hst-.l.Ht.XI !- 1..J.U.1 \U ,..,!, / )' ( K , I orly known. These mostly occur in areas of high and T. occit; Weaver. Further : d are probabh narrow endemics. Both B. < /p.). In contrast to these less .... . • ... Light to repoi ealth of immigrants specimens including AAU, B', BM. BR, COL*, F, having onh 10 -,, . B i ,, a ^hibits a rathe FMB, G, GH, HUT • . k M. M v. M" N\ , P, R > , RB , eMuoidman bieadth w :: . . H-\ US/ . W . W i(T. „.,d Z ,,,„„,,!], ml.ilit 1- - , oJlertions since the . =« ki «i, \, I. ira. nix • < ,]'•» hul.-dilti. - ft n , -si , n J, .,,i 1 open savanna (B. i,.m i , u, llh, a I, ill „ In | , , , u ,„. / i ,,i. 2-JO.i Nil ,i tm 1. , 1 1 superba M. R. Schoi a forests and tend to be glabrous and generally from Brazil, which a decrease in height toward higher ele\ations: B. preserved. trees, B. excel e 4-35 m tall tepui- After those first two species, three more were b and B. superba are »L, H unthonouLs Gleason & ■■■■■...■.■ ..-...,. provided a brief s n Leeuwen- (tnate, elliptic, o\al, to obtnate), the leaf base ig the midvein this brous to densely hispid). The are ;nce (Mori & Brown, 1994). have specime .nits and seeds: B. have few taxonomi -haracters. In capsules on i per species, seed - -..■■■ han,,m i,t. ioMatdfh, p, li .1.. p. : : r f ■ i \ antonifolia ^ ::H-;«f/ . \/ og 1,F1. !ra (Martius) 6(1): 267, tab. 72. 1868 (lectotype. designated In Le-uu, i Branched shrubs or trees, 2-40 m tall, glabrous tc .*, Made- -H.sple . , , .. i.^r. «a ,1 MijU,, „A, on,. Lm.kIi.s i .l»» i, J. ,h i similar to the leaves, sessile to short-petiolate; teoles (organs on pedicels that subtend individual us (calyx, corolla, and s : .p.-,! rt it!. -ilonphi^ltul^t.. 2/3 oltl^«»r..l .o.-.ll . lube ...mI ,u..Jh K.be, ,1-n l..b,-,;Jal. 1 „..-. ; ud ', I „ . ' • , u .,,.,11. h.be-.n- .K,|u4l»< ',..,. ,mu-. s free portion less he ,,-. Il.^tnL.. i o" (t, sqviK.i m dehiscent, e] pubescent, erect, tan ters, especially ha rthe'cahx-I-Ily^lobes 6(1): 288. 18; Grosso: "In 0.5 mm; pistil ca. 8 mm; ovary ovate, orbicular to .■-..:•■:. ,:s 1, 2C, 3, 4, 6A, B. (excluding styl seeds per locule (e.g., surface-, tiunk to 10- ') , in (\P ,u I I, riit - secondan \eins; base cuneate to rounded; apex br< n-colored wings. cellate; pedicels 1-5 nu,, 1, ,, t le m^uLi ( 3 X 0.5-1 mm. Calyx campanulate, 2-3 X 1.5-2 ,i I in > I m ,.i < mm triangular, 0.3-1 X 1.5-2 mm, apex acute; corolla '""'i"'™&-> "— 17-30 mm; tube 8-11 X 1-1.5 mm; lobes 7-9 X IUCN Red lyunia antoniifolk Distribution and habitat. Bomunia antoniifolia ]road distribution in the Amazon River basin and its butaries in Brazil and Bolivia (Fig. 1) at elevations 80-800 m. It has the broadest distribution of all ICN, 2001). --,,., :■ ,: ■ Typification. The disparity of the type locality as Sen Riedi' 1J-.Q) is ips 1 .' 1 u fl- | J i i p " D , L ' ' ' " P lk * * ' " , ,'\i r ( i - I! a ; .u,.-'w .. • ininp I...L-1 Mm,, l! u,-. Ri.„iJ ^ """'iM «, HI l^. «„ ',,,,.. „ . ..P _M<. I a»Li. ,1 (p.g., Duckp, 193? ><\n Ki. .' ! Ni U P Rubiada In Mumntmi Jan, , I, if' ,1 ,">! h'.i n' . , that Riedel did not collect in Minas Gerais in \ \['- 1 (.',m L'^r'/l) II, .(.N.-Ti h,it-tVMi I. V, , F ( ,t,l,^d au,u U.Fn ,.n, »,,!, R which is near Cm, .:.■.-■,•■: a?.U u ,1 ,w.s .Irjv. ,,1.-1 lh. ip; ko«e\Pi. a Humaita, estrada Hnmaita-Jacarccanga, Km 150, I. 0. A. t r,...n ,u,i .- 1147 1. ui- ^ ;,;;;;„,', !7\ v u ,; ,1 J 1 ^ WV'.'V'-in,. Riedel, tke collector ited tke main ^ set of his collections Russia (LE), Caxi and the second in R MO and NY Ferr, sheets were distrilml he labels are (SP): specimen: "Indeterce a, Riedel." ™ A Mad 1: 211. 1935. TYPE: uperioris, civitate A t NY USZ lofsTenf DeC ' 1931 ' A ' Ducke 2376U $ ■a de isotypes, G!, K!, P!, RR [2]!, Figures 1, 2C, 3, 4, 6C, D. Volume 96, Number 4 Grant 551 2009 Revision of Bonyunia Branched shrub to tree to El Tuparro (Colombia) It i 5 assigned a prelmnnarv / on the midvein of the itus of Least Cone ern (LC) as set forth in les, peduncles, inflores- the IUCN Red List Categoi ries and Criteria (IUCN, ilyces, and core llas (glabrous only on 2001). if smfaces). Lea zlT^-i^z Etymology. The epithet is taken from the Latin 5.2 cm, thin-cor arker adaxially, lighter ^growing aong aluMalh. ad <•.,,] - UJ td, f -< nooth with some slighth [face with slightly raised ,224(NY[2]),ADudce354 moiiH-. m- 1 .lateral; apex rounded to oh°o'ei ? ra n Tunuh' ^R^frZ lorescence 3-9 c sessile to short-petiolate, Negro, Rio Marie, village base aequilateral; apex 198 (M, NY, US); s. loc., /. ;tioles 0-2 mm; primary / />>>lIVN 1,1: . COLOMBIA. Guiania: Rio 22 X 8-9 mm (ex( L S \dJnles ""1-1 i -)\ I n\ S LIS u VI,) Rio i. Its in v: =nce and calyx ai ,j;ta>»i,-. ..lnJ. '.ributio habitat. Bonym 3. Bonyunia ant, sp. nov. TYPE: en un sitio cercano a un tepui en donde protected areas such as the Resena Florestal do Leptuan B!). Figures 1, 3, HC\ Red L -r , ..-.v^n lius, .HO,,., JlOfll tl„ee , 11,. t! I,- .)1,P -,U f ,t. 1 llM ,l, , preliminary III rable (VU) according ncles, inflores- «»"/"<-> n,f ,|,i|1ilt ,. idken fiom the Latin Bonyunia magnifica J. R. Grant, sp. nov. TYPE: Brazil. Amazonas: BR 319, Km 190, Manaus- 1974, G. T. Prance, T. D. Pennington, M. Leppard, P. P. Monteiro & J. F. Ramos 22804 (holotype, NY!; isotypes, INPA not seen, K!, MG not seen, MO!, S!, U!, US!, WAG!). Figures 1, {ranched tree to 20 m tall, glabrous throughout, m. Leaves ovate to oval, short-petiolate, 4-10 cm, iole 3-5 mm; blades 3-9.1 X 2.5-5.5 cm, thick, iaceous, glossy on both surfaces, distinctly discol- d with the adaxial leaf surface distincth olive- en and speckled, and the abaxial surface a solid dulous vs. hispid), with glabrous calyces and ^ acuminate to acute calyx lobes. found in the Ararac , Mesay. 3 to 4 seeds i , 8 seeds per fruit); ar, 0.5-1 X 2-2.5 mm, a] nate to acute: ; corolla 12-14 mm; tube 6-7 X 1- „l,hi— • -.t.-i.i, 0.5 mm; anth, ovate, 1-1.5 X ca. 1 mm; style 7-8 X ■iuund l,i]„J,erl. each lol,^ ,, i.mn.al,-. 0.75-1 X 0.3- . n i. Car • ! 13 8-9 mm (exc ee, yle base), brown, erect, j distwli-e . and similarities. Bomunia excelsa is or« ,,w,-. -~t 'III iTi 7^7!\l,Ja .„.i ovate, to nearly orbicular or lenifoim a.ti I rounded to ro.date l t ..( !m- The .da' id I k,V -ml ,. , l '-'"• '' ' "'< 1 ,fl ' ,V l ~* mm; bLldes M " )3 2 " a 4 X tutl./ • <, , .ul/'r. /,, .., lt I,.. I.li. ,( ' J - ^ ' •liL'-oiiaceous daik.ia mm (ev< k lm° only known from the type collection. It is assi„„ 1 , »'- '"' ''«"' ' ""' "' ' '" ' ! h < ' ' " ' " (CN, 2001). "magnificus,"" meaning "magnificent." by Leeuwenbf | a i so accepte d here. 5. Boiiymiia minor N. E. Rr.. Trans. Linn. Soc. - ries considerably 7C." h^elaigeik-, si, bxl , h,n. p lt t« in In the ■ 11. Bull )(.!.. IV. ,., „ L „1 .„!,„ ,]-,, 01 l „, llt ,„ lj , .hstlll.lk Oldni, if McConnell 161 [K]). —T> U 1 .< t i> > , •, F r t i ml 1/ ' - -'-, 1 IUCN status of LeW ( oncem (Lt ) a-, stt ioith n, til -,', 'd ., \ »,..,. k u u m O Ihle, UTi, 11 i IN hdLi t< M .1 ^ " I' it« mil M 1^1 i i J .I.. .1 >,\ I n '.I , « .» r, m Etymology. The epithet is taken fiom the Latm a lo largo del Rk C h . , ,.i v ' ' , .1 N df 1 Salto Caiapo ill. rm\ h j in., -p-ti - jii 111. T-nii ^5,1 1 ,„.., at the time: Borayun , VY, U, US, WAG); Ros. io, \\ -I Specimens examined. BRAZIL. Roraima: Serra dol Sol, \( " J'™ ^ ' ' \ ^ 'l , ,',, ''.I'm!, ' ,' , , 5 mm; anthers ca. 2 X 0.5 mm; pistil ca. 4 mm; ary ovate, ca. 1.5 X 1 mm; style ca. 2 X 0.2- t mm; stigma bilobed, each lobe spatulate, ca. 0.5 0.3-0.5 mm. Capsules ellipsoid to obovoid, 22-25 8-10 mm (e: 1, tan, 2 to 5 seeds ge, seed wings dark Uumi.tf,™ M ot Lu,, , / i -t.nnml V I 11,,, k . , , »« u. 1 !< , J > , ., , I I. mil, lly, and solid gold- Uari, F. Tamayo 3132 (US). and B.magn nd described abo\e 6. Bonyuma nobilis J. R. Grant, sp. nov. TYPE: U discolored leaves and dark orange seeds. 14 Dec 1990. P Paku ios, J Estiwl i, P Fun ■ />, „.; IL ,,,.,,,/.", » 7?, >, » ,// „ n r IUCN Red Li unia nobilis is only \ J) accord- )i: ,1 I ,si ( ii.o, ,„ -,,,1 ! ,,i n, (LK N 2001). Etymology. The epithet is taken fiom the Latin ■•.■ml.il... "hummus "iinl.lr-." 7. Bonyunia pulchra Ricketson, J. R. Grant & i tall, glal i thr, -5 cm, thick 'c. distinctly disco] died !< sluf.d .11 Ok ice 6-10 cm; "r,,, 1 ,L ■ . 4K >,h V ,lll „ si hlU ],„,. base cuncate; apex obtuse, rounded, to acute. i j,.,.,! iv u,.l e nl.nil." n «°. Bo numa -pe. Ulnli- [ I d-n -,) n( T\ Ft 0.75-1.5 X 2-3 n tube 7-12 anthers 1-1.5 X ovate, 1-2 X ca. 1.5 mm; style 5.5-7.5 X 0.3- el at the branch apices morphology, it Colombia). Distribution and habitat. Bomunia pulchra is a In s 1-2 cm; bracts • : ■ ■'■ - IUC\ Reel List categon. Bomunia pulchra is > iCN, 2001). i the Latin 0.75-1 X 2. Halm & Gopaul 54211 [US] is illustrated. ia! t ^> V.Yfo/»'-> ,.!)!, k< to .,.-, mi.iale fi.fJ-.i, ,, f . - i that the es 2-6 cm; bracts o\ate, elliptic, Ian. corolla 15-17 „„>< tub, 10-17 - 1 VJ . , ■ n |.e ;,, . 1( „, , - oil, ; .1 .1, Mo. i.l Ro, >ima along the each lobe spatu l a t e , 0.75-1 X 0.3-0.5 mm. Capsules . : IUCN Red List category. Bonyunia spectabilis is <.'ti..lU Eil.Ij.i>i..1 it ['< a-oi.b,. t.UKNO., 1 is! , ria [UCN, 2001: Etymology. The epithet is taken fiom the Latin lent (van Dam, 2002)," 1842-1843 [28 Oct.^. Dec. 1842], Robert Schomburgk 614 [= R,< ,-ml "i h -nl.'ii^ '• "| (lectohpe, designated In Leeuwenberg, 1969: 158, K!; isotypes, BM [2]!, BR!, F [2]!, G [3]!, GH!, K [2]!, P!, U!, W!). Figures 1, 2C, 3, 4, 8C. ,atulate to c e j, ,itM - r I Re, Li ot the umteisidesol leases, petioles, pedunci l bl T , ,_ ai „„ )TUjN sfatus of ovate, oval, i -,kMjih.l,d,t.,ab,i\,alb.^],.-i l H, l<„.fir i,cn. Tlit l.-ikTium L,l,t Is ..J <,/„.,»/»/, — m.ln, ,ems: base ion ided to c lineal, . ,,. v lh.- t,„ ^nU^ m-I^ „i E, V , ■iPdin 2(HI2, Tib- lust number, 614, 4-7.5 cm, with 3 to fa pair- of arching secnn is horn ijobeifs second collection senes ..hi.h ,e„^ petiole 4-6 mm; blades 3.5 7 m-h,!,.,, £ s«/.«6u \ccoidmg to un Dam (2 however, Our Village was neai the settlemei pears to be most similar to B. excelsa and perhaps so to B. antoniifolia. It differs from B. excelsa in ..„ving stems ai .florescence equally -ii.1 \ l ,.i, II. ''2\ '' J,7/rrrirV| 1 ff!^"'in 1 k.,,'!L\t'VTi' l d longer Distribution ami habitat. Bonutnia venusta is I \ 'F nil B,l»>. 2 Id Lrn in i, Manaus, at elevations of 50-1 .00 m (Fig. 1). Of the four S, US). d< as B. aqua, L999: 564). 10. Bonyunia veil p. nov. TYPE: -t, Kp. Jl I"', - 59 ",' \\ '•-'•"'•" ti-m the hpe collection, fiom a i..,i»,ilK 5. Sousa 1103 (holotyr Cntena (IUCN, 2001). Etymology. The epithet is taken fiom the Latin . 3, 4, 8D. Bu.1,1, If,, el' ni..]l In plnUiis .-.], stems, peduncles, inflorescences, calyces, and Ibu.H..,. I.H.sh ',bTJ^', cream. Leaves ova ate, ceae. Amc , M. Thn J V, I .(,.<.. A » R Pepper, T. J. ■" [2]), 2641 (MO \2]). , J.'ittV L J < ••'■ ■ <" '-'>'M-. (MO, USZ [1]). ■■ ■ ■ (AAU, NY [5]). Jnl I" i . NY. L -. V, u- [">]! l'„„„. I !' j"IO f^ 1 1 T — TV \l,|4| ' '"[Ml ■ (MO, NY [5]). MO, NY, USZ [1]). MO, R, NY [1]). R ,H >. IT .»,, tX — 1 11 ' „ I. RB, US, WAG [1]). (K, MG [5]). WAG [1]). . WAG [2]). PHYLOGENETIC POSITION AND A TAXONOMIC CLASSIFICATION OF '< Mmke * AETHIONEMA TRINERVIUM md Klam Mummenhojf ' (BRASSICACEAE): A MORPHOLOGICALLY VARIABLE SUBSHRUB FROM SOUTHWESTERN ASIA 1 ~ets «itu ...|.lil'.«.ndl ,Uc ]-, a'\ilil ?m,\ ,-ili v FuH-n> ■ .■,.... .■ . :■•'.:■. display a subpuhinate habit (Hedge, 1965, 1968). str., along with other genera (e.g., Alliaria Heist, ex ally, the evidence for placing the Fa da Jacq.), in tribe possesses auriculate or amplexicaul lea\es, the v hie structure (Davis ■■..■. ..■.■■■. 1, l- i„ ,1 «., ih, n. , IM t UiH J,« t, > , o. t. uel . , net tl„, „i ho otth I I 11) il . ik 111 i .1 ilmn <■ mi (, II I i Lamond 1980) .) 5« >„),„/ r ;-, M ( ,i['( > Fi.,»ifl Kai|a;,'.. J'J'j.i, UMi.i. „hU, ., t, Jm. . •, «i.lu, A'" v" - 1 '". - 1 »"'"/, (R Ji f & lellen) Hedge 1a Hedge it! Ive tl il < Ilium nl-,'1 ,1 .1 l"<)". ', bhd lis. Il ,lh Ion a! -, al, 2001). Thus, it is unclear in typical A. triner- segregate A. trinen: it num. with each flonei's pedicel separated In ■■ : ■ eilstein et al., time of anthe pical 1. trinervium 2006) also pimed that Thlaspi s.l. is a pob .acted racemes at lly using sequential pairuise comparisc of the aligned sequences of the ITS c lf -b* ,1 h, 7,,'us,,, - I I 7 ,i-ik H il U )6 Mummenhoff (Zunk et al, 1<- ay et al, 1998; famih of the Brass c ted as the Resul i ■- II.. Iiii.iI .I..I.. - ■■ ■ — > (Table 1). arch. Pairwise distance sequence divergence of ITS is calculated for each accession pair in PAUP* ing the Kimura two-parameter model (Kimura, these, 188 Total DNA was isolated from dried leaf material of wil The results of the ■ ■ -■ il'IKi i.sj„oTls |lmlri . mm II, .|,„ rll 'W7.i, I.; Wli & M ami. -nl.ofl 2.H/I; at 94°C, and 35 eye (1 min. 94°C, deae. Also, t im clearly belongs .■ . stadt, Germany). dneigence between these two main clades is iF: 62 2 >l !3 :i; AY154812 f (36152/ J 3« . 3 Kama campylophylla F. K. : K kurdica (Hedge) F. K. M, .1. trinenium and other .ltf/i. ITS sequences to clarify the pin logenetic leLifi... .1 ■.[ r< jp, ' uioush assigned t. , 1821; Iberidella, Boi Jer, 1867; Eunomia, P ffarian, 1996). These g£ : rices in Bornm.) are ispi s.l. and are better itrast to treated eith '. K. Mey. or Pseudo- asy and sempervivum 'ig. 1). Finally, Thlaspi 1991; s. str. is mot Peltaria, Alliaria, and lenhoff, Thlaspi s.l. ium F. K. Mey. and ( these Kotschella F. K. Me}, than to any other of Meyer's 3uter et segregates c .)• Except for Thlaspi s. .■.,:■.■. . -.' '...". .1 M.a ,,-<<,, jf t i.i ,mi i. .ii..;)Ih! i, -l. h',,,1 I 2>n, Il„ ,, , Meyer (1973, 1979) , "I '• iceras F. K. Mey., Raparia F. K. Mey., and «',(",/./, I k M.n ,!,. 1 Ii, rhe most itcent I the remaining 77^ tribe Noccaeeae. The priman differen difolium Boiss. = E. beroi »-i oi th. tube \etluonemeae is runn imestigated In one of the authors (M.M.). The genus Eunomia has been most recently treated . i 11 i I. I ; / l> iM ' unpublished data) ,ii.»in the left part (see Fig. 2). description, see M photo!, one sheet with two specimens) beais a show a firm texture, while the stem lea\es are E ichaux [scripsit y]." 1 • -luk | id- unl ..pi.uk.t- ; • / z,< DC [srtipsit desriiptioi.s !,■ dr L , til. ill -I l.l.UMph.i ,1 ft« (1889: 3), or Bois-in i Ji!(>7- v,m| j>„« - mk.im.mn.. Mm. a.™, hi .l.n d.< i-.i.tMl m la il.-n 1 , '. ii. ..] collectors ir .Iw.Jnt la. k •>} ml. hfe „ il (1845: 61), whose close fanul herbarium, libran, and archiv that Michaux traveled in Iran from 1783 to 1785 and F. U M, i 1 I" > i ' d i < ]i ^ «,i„l ,. went to Hamadan (according to the map in Jaubert & appinach um,,. the bootsaa,. Evolution W '., -7 M Spach [1842-1843]), certain dossmg r „, ■ • ^ ^ L ^"'^ ' - R * P "- 1W Alwand Kuh, like! 1797 (Boissier, 1867: xxvi). Finally, it should be iu.M (.^..m u 11 ',1 Km i i A (, ] dHllledlel^tolhl" 1 It, ,,V h "l t'lVir.lu' ri " '' " ^ '" '»'* llt ' '"' "" '■' * I. m - U Mliln i"!' \m>lu tuj, Hid <.(.-.. irii-,1- .."". leMMtcd MoKuldi data io> |.b I... . „ ., P., -MS~',22 \l in.u- L- L-IUkI , I. A. Al-Shehbaz, F. T. Bakker, H. P. Linder & A. Sa Mi l.lli ui*.,,, _'■", Hnl . p mni-nok i. «l l.,ti »i Z uil k K '.Km, ,mii .if \l L< , h l II ll.ikt l"X. A SYSTEMATIC REVISION OF Simon T. M arlotte M. Taylor GAERTNERA (RUBIACEAE, GAERTNEREAE) 1 i, S. Teo Ping, J. Thomps rche Appliquee au Develo] 1 Progran f : , other substrates, to in that region. M->i n JO.H- r. 'I'uMiit V M n n 30i., tlic i u, illillt. oldln .hlldlll ivi,. «., !,ifC 1 r^T* •! | ^W^^ -'■■'•' / 'p^wVn -x. ^<1r ^ - 1 / £J ■ : r 30 S vi*^ jn \ 700 km i^ 1=1 30; E : 1 i J i specimens. All othei measmements (e.g., leaf f n >th - h,.ns i , „. I !id U ,.l . ,i, matenals Tims, we ogical charac- sary. descuption In the mdmdual speries ,U srnptions S1 , L , ^ ( : . , ■ e (e.g., Dorr, 1997), v. ill in this revisio ia imposed In the An index of all taxa treated in the current revision li L999). Preliminary I,- ,-.*■•..., , [u. senie.l m 1 ,, ' nuclear DNA (nDNA) data srt- loi ',•! -ti.-u, sp- . i* s ' hyj • 'jp ]»( .r . Hi,..,' L \i AY046331 AF333870 AY046361 Davis, 2005) were and PepC-S sequer capitulata Malcom e 1). Th i Baker and G. 31 Gaertnera LanL.i. M..L- „a. . 12 species recogn HE3 Double- : the ITS regio] s ITSLEU1 and ITS4, and the two copies of the fourth intron o in 5l)illll i ) i ill ' i IVJ ! nl .S.A.). Plasmid DNA ine lysis/PEG precipit , 1989) prior to sequer to four clones were seque s were described in ing reactions, following the recommendations of problem of I ius names, Kuntze PEPCINTF, PEPCIN listory of Gaertnera. W,l>\^ i\j.Jul d. msI.us it,,iilH P^\ l„ ,1 su,,nl i!< , s a >.]•, 1 m. i' sequencer. Pielimman alignment of the ITS. PepC-L, and bet. vn i.ie.Ueir >u 1 membeis of the Rubiaceae, the Sequences were ana iaceae and Rubiaceae. Brown 1 parameter, and proi belonged in the . .., j.jnki'ujljtj'^ei. i^rlt< ex,,lu s ,Ul iLM'l i . • .i,»d two genera again, inside (i.e., adaxial face) of the corolla lobes in its anthers m mate to the petals Pagamea (Taylor et al, 2004 t; this last condition has not rl, i ' - '!,„ >t,h, t j '*,<■•. /i »„ ,...s t !,»..< I.ii/ .1 I. I„ut. surrounding the base of the petiole in G tlyx and the anthers which is apparentb however, this all alternate to the corolla lobes, with no cafe ) DC. ex Meisn. foi a plant that was piobabh a G. cooped Hutch. & trther. There was, howe\er, a poll. i. liif.jiiioJ.'N in.\ iu.»i.| .n.]it,..ivl , iM],...,., tn.isr.v „ \!( .\^Ih >. w ill. , 'In ii ir ' 1, h. pi ti, tlu ill x I , mil 1 n mil- in. nl . mi t hi uillm ^ t« . II. .... ill. .', tin.. ..m ^iitli. l.m.|s u'll..!/ an. il ,i i . .1 „.,., , i.i .1, , , I no., thus the identity of authors (van 1 ,ut others have been comprise,! the Mascarene species with sessik li < i- m liluimi >nm< . w I im I ,n a teiatolo^ralh ..-HM. -MtU ^s.le f |,e.l., ,-ILi rhoL.it.n, 1\L,<1)...-. ... ... M.dnol sciipt dccnption had hetn assigned a position small tiees up to 15 m tall jt'i. , I n-M A.. , '.i.l S/-c u.ll.l .horn ,11 _ , £, ( ,,.„„/< . n^aitts. 6 „ nop folia) an Malcomber and Da\is (200? .d on all plants of a n .tn.,1,1,1 n.hn.r... (h- -e.nl.,> -fn .!-..■ , „i .1 vu-tim - ,U, ih> name for the gei es, to densely pubescent with i !. arenaria (MAD)** G. brevipedicellata (MAD) . drakeana (MAD) macrostipula (MAD) /7/sp/tfa (MAD) G. madagascariensis (MAD) G. pauciflora (MAD) (MAD) - I — G. inflexa (MAD) i G. cuneifolia (MAU) jt- G. longifolia (MAU) JL G. psychotrioides (MAU) L G ecfenfafa (MAU) amosa (SEA)** G. belumutensis (SEA)** schizocalyx (SEA) G. wm/nea (SEA)** G. caprtu/afa (SEA)** fracf/ffexa (SEA) G. g/06/gera (SEA) G. aphanodioica (SEA)** G. iunphuhniana (SEA)" (SEA) G. ob/ance (SL) i L) eri (SL) ag/nans (SL)** iowryi (MAD) ■ u/afa (AFR)** ginalis (AFR)* (AFR) (e.g., G. tuuln^li iH, ui T M A. Petit. ( ('>,/ >-..' m. .1 -. t.l . 1 II. p,rs r, - « 1 >.u. 1 sp. lt sh, ,W,„i, < -].i iin ii at it a ,,„ M( | ( ^arr^l.le rn mam of the interpetiolai ale down the The form oi ra is rather unusual ... .. of G. obesa mentioi ,eh related genus - nli ,1,. ,nU, lf , ,m„ ill. .iimulul. J it Ip ih. /',«/.-,. I",. . ,i,i>L- ,. h.,.,1 „i jJitll. I« ,r i removal of old branches. (e.g., G. sralensis (Pierre ex Pit.) Kerr) to well- illi 1 .n.,1, i<>< ir;,. t,n, iicLli >1> -laiKt.Hmr „ , {. suhsessile to usualh petiolate «iti, , , . I] I , P . .1 . Il .1 \, I ,p-.l t. I.nl... m,, Ml,, i --75 mm long) that ternifolia, 0.5-2.5 X 0.1-0.3 c h or occasionally and obovate or broadly elliptic (to 55 X 19 cm, G. leaving a p ise. Two different van Beusekom (1967), 'the leaves are so ^,,il a ,», ,* .lamaging the sheath, which undersides of the leaves, in the ,mN i ,],. , , )<< i -w Ml, o t„U t! ,1 „ft u «, I" )s t m nd the top to forri a fmmelform or ^ase-bke shuctuie yGaeitnaa -e < subrapitate) This (DC.) Baker). Some species have six lunsytudm .1 , .1 s [,„1„ ,r ae amen on the stipule sheath (G brevipedicellata). The inflorescences are sessile to usually peduncu- brecht 11988), the iml.ii. -. f n. - - ^<- ,ci.,idi . i.i W t mmm. „fl Rubiaceae "tripartite" or "termmal and sessile." It subsessile to usually at least shorth pedicellate. »,'l U 1,1 1, , . ll I ,s .1(1, , I th~ sUU i, s tU 1 ! , ! „ I ,s > s i'ai lei korai 961 • / : : . ay ,ra. hln.K J rally cup-sl 4- to or shortly 4 ,- to J )f the lobes (e.g., van Beusekom, 1967: fig. 4). ver, most of the internal faces of the corolla are glabrous or, in a few species, glaucous. In wm .pedes, the , .' ire ei ;ular to ligulate and flat, but in a few species they ickened at the apex and prolonged into a hooked al protuberance, and sometimes these apices are ,li,l a,e Fl ,hv, ,1 ai.d « u< pHate at the apex and dhU h w U , ,J, V ,\ ,al,v 1,1 snilu.l rh. ..llll I- .,, I 111.. (..-I lul-lll ,1 ,! „1 th H„., Ml nil. ,1 ? flowers of an inflorescence or onh on a few D; are found in the vers in I thei timber may vary on a ieoes. The 3 calyx siz 5-4 mm and is not ften is described in ■ it was not found to be I,< iiilMiu.iUxefoi sepaiatmgcloMh iehted-,1) n mh rl. il II ^i ml i i- mi. d t, position may be ioun-l -,> < m • • il n ( i t ,im depending the numbei of n. f - tl „t -U-l .,. j on the species. The the fruit also depends on the s i mi a' tl.Mi Ih ^Jlh u-t.iliell, ( .-,- 1. -. u ,1^',/f./ -]> i. <1M pels obs ), this is i et al. (1996a). common path (Piesschaert, 2001). style of the pistillate flowers. Rubiaceae tha crepuscular and may observed, all black os ire protandrous. few species from Mauritius, the fruits ha\e been The flesh}, drupaceous fruits of Gaertnera are reported to be whitened (Bojer, 1837), but this has not piesumabh smnlai to those of othei Rubiaceae m • !/ fi.,m J-JO < i-Li.nun. iMih,i7, i -,r( W ,-. l„.iu . ,- ol,-e, ..■ I dimuo t| 1]s -liul. ,„ I xanmg to some extent between species and also n is have been found. noted by van Beusekom (1967) the fruits do not show (Fig. 4; Malcomber. 2000. 2002). \an Beusekom „,„.,!, -,ul lf l-l,,s , t,< , , f 1, ,] ,,« ! V >■ ,U, i ].» s „ M -,, ,1 | )llt tl. I-.: pollination studies In S.T M in Beiakas Foiest h . colonized regions, Resene, Bmnei, in 1998; collections fiom this especially or, igascar, Mauritius, biology is kno«n are either eUdenth dioecious or Genera, 122 mainly present in Africa and/or Mada- dishlous. The breeding biology of se\eral species gascar," app. part on the work of Asia are expected to onstrated to be dioecious. The e\olutionan aspects of the breeding sho« a cons,. , distylous through BlOGEOGRAPHY, HABITAT, AND DISTRIBUTION species of broad geographic range with mixed Islands (nine species in Mauritius, one species in The 12 Af, .era are found from Reunion). Accordin presented to Senegal in the northwestern part of the continent to J 1)1], i. li-Ku !s ( ,s M,k,i, i^ioi) < i- / ( .,*';;' M>-i,ilh sjm,i. s,.ic ..,lu.<"! t( M ( st ,., .it, -I ,1l,t ,1-nt ,1 m tlit M.I.. Mil than the actual range (e.g., G. Ion an region. (Sdiveint ex Hiein) E M \ Petit k>. um lit ! ill ^-<-»,. ■ . lias ,1 , • „tei . 1 ,| . < i^s iichuess (as well fication of White (1< s cmienth each kno«n fiom a single one of these ■ ■ ... ! .nc I'- j l< a ,',,'. I /»/, VI *.»i,sts ,,,4 1, J.IA • ,„ u,I wl., l,,i,, • ■-. .. ■-. spicata, G. traclusnla) of the Dahomey Gap. gixm additionalh to exclusheh on sedimentan : . ■ : several generic distribution patterns of \fncan along the entire eastern part of the island, through the "Africa-wide" distri not present in natural vegel re) and westward of the island; G. vaginalis is the most commonh G. tlrakeana Aug. DC, G. hispida Aug. DC, G. ■.,'.■ . •■ M.ll'OLllL,. \s ,K,k-.| I* l).,Ms .,,, ' .. ' ■ . Gaertnera is found in three of these: the Eastern "'■''"' '■■■■■ : . . ' .'.•■■ , > -■ ■ I setae. 01 with only setae. Most Gaertnera species have InM, k i i u.li n, „.U 1 J i f, ,,. muln, u, ,. - t [ i ,1 • >m!ieast Asian species. Sampled Sri fhmers. Cal^ lob - ,«.e n-.^lh .elathch small and 30 mm long. long been legaided as a sepaiate spei w- .11 • > .11 (-1 ill i.-aional populations considered a hybri ieri, which is Such broad pread taxa that are ih... l.Nlx.l ,. 1 it,l j- ,.iiku..v ■ .1 o ,hi ,.1,.,. , (eg \mk [1970] grouped 39 local "entities" into a Thus, ft/flftiflri p.cn'i.K ,n.wu,!« ,'.| t] ', \ : Van Beus< in his 3X pa a list . ndnea 1 : lose]) ■ , , !u, 9 hat it i clear duel - the syi ^ , tk ,.t \- i m. k»li,n R..n.K. .i.,.l tl . W-,,,, ., this classifica atest in this region, within Gaertnera (] the classifi- work). Thus, in this case the addition of more data h iLl' ,( < llol M.l( J I, I, l,'l I! I'll- II- 1 1. ,11 tt-lll IMf.s-ll, . nil- Hl.s conomic Treatment Lam. ' ■ f>'> G s eop 13 Feb. 1792, nom. cons., non Gaertnera & Vi i I : Pristidia ThwHites. 1 1.1.1- )1 Zr J 2 J -I" ]!>• HP! iff : reduced to 1 or fc flowei 1 racteate ((; nlata,, p.hm, k ,m™s»i Ltmrfo^U - h. , 1^,1 W.v.mn, Hi) , „ I 7, , ( , , ;i ,), , l!lllt 1 1IE 1 (1 •der axes up to 30 mm, deltate to linear, o\ate, or m i,l,,.< I .t - \1 cl il ^ ^ die. ,m 1 ,1 Hihill. .nt! ^ ed or ruminated. Regional Keys to Gaertnera Species ■ 22. - J 4b Nl,,H. - (Ivmiv. d '"^"■"•"'- , cles not flex uous, with inflorescences ascending 24a. Caly t, 5-7 mm lo ng with at least some lo liar or ovate to linear or narrowly '. 60. 53. a 26b. Sti , usually regularly persistent at least 1. , l.^h .1 i Il.is h ii. me . I " " M. <•, Mah-mbei & \ P. similai in general aspect to some plants of G. us species is so fai known fiom few Bdthie 17098 (holotype, P!). species to be Endangered (IUCN, 2001) based 5(to 8) pairs; dom ies 0.9-3 cm. 2 . Gaertnerj ler, sp. nov. TYPE: •minal on principal and/or axillan branches, gy. This species has been collected with -3.5 mm. Flowers 5- ide glabrous ; ies to 0.4 mm long, of Ambatolampy, at elevations of 1400-1550 m. H( m.n .1 io> tl.nuiel 0l - ] lhn fj , ei« ! ts 2 ,u>rvd inh-i «o,k\ ! !■ Utn J, Ion j.Mii, Is. anl ,. I,Hk I, L flowers. The measur. la width given ■ ' ' •■■ '.■ . . across the corolla 1 here it is known from Borneo, fig. 4E. 1967. TYPE: Brunei Darussalam. Belait: Seria, 18 Apr. 1957, S. Smythies, G. Wood & P. Ashton S 5909 (holotype, L!; isotypes, BRUN!, 1.5-9 cm, smoot: -24 X 2-9 cm, oblanceolate or ovate, apex shortly mm, deltate. Inflor ii.ij-.mu1 lilt -i),u -'">" ivl • 'V / i>J. j B min-Miia j- B A r,. f -'■ '■■■ '■' -•-■ " ■ ' " 1 "" i ' " " ' " ^ t '" U< * """ " " '/ 11 HI >-l, S,,„la,. s HJ i »l , .1 / , 9 mm, 2-4 mm dial i upper third, ■■■ igure6H-N. ■■■■■■■ Trees or shrubs, 3-10 m tall; branches terete to 3 ™ 2-5 cm, smor ate. Leaf blades 5.5- • - *-_0 > 2-J'.|-lJ 5) mi elliptic to oblong oi li^i ^..l^es^m XiU.L ,Mjlr. -in (J|)>ii| , Phenology. This species has been collected with les 2-11 mm. Stipules tubular, ' '■ ' ,. U.U. ,. Mil I- LI, IvUM ,1.- „■,,„,,][, n „ lMll p.^lllleU, ,. '' ' .'. |. !"«.!«:. i, ]>\. h j. I- l-J.if-,.5) mm; bia< teoles ........ which includes the hpe specimen m an L. - k mi | l|f ljU t j, //,,.,,] x , „ s h a ,, c d 2.5-4 mm •, Beusekom suggested, but functionalh dioenou- ulli , tll . ull !t, roiolla unite, ihomlxi 1 comber, unpublishe, a tube 10-16 mm, 1.8-4.5 mm diam., inside villous in H ho. . . of 'he ;>,ti.s GwT; ,,,,,„, rluul, I. I»s ' -i i mil., Ii t . ' . : . within the corolla tube. This species be )-15 mm, 1.5-4 mm the G tagmans complex; see also the discus- diam , lobes 3 5-5 5 mm; anthers shortly exserted, distinctions. ary through March and September through Anomadiobe. .S,7«m R-V 0512 (MO. TEF): Ambilobe. Wcter- of Gaertnera in Madagascar. Gaertnera Reserve, Malcomber 1152 (K. MO. P. TAN): Emi Jla tube, 10-16 mm long. ^ 50 m, 1962, 1. /. .1/. Leeuwenberg 4154 elevations of 0-100 m. (holotype, WAG!; .otypes, BR!, K!, L!, MO!, PI). ^^ ^ ^ ^ been ^^ ^ Haec species Gae,tne,ae coopen Hutch. & M. B. Mo-,-, U with fruits January often lea\ing a pel am, tube 10- l',,!,',',,.,,'.,., 1 '',,',!.,'!,!,,,',' |_- | -,_. , .',,, "| ' l',i",!, '■ »•'■'■'■ -•/:■ iv -•■■•''■ iv- \i".iv / ./. ii,/,/. /.•■.,\\ \.;, late. Long-styled j-shaped, 1. 2.5 mm wide, glabrous outside, with hair-ring insi truncate or lobes to 0.2 mm, tria alcomber & A. P. clavate in bud, when open salverform, outside Davis, issouri Bot. Card. 1.8-2.3 mm, ligulate or oblong, acute; anthers a l ong trail to the summit of Marojej\ Est, NW mas 1.8-2.2 mm. SI filler & P. P. Lowry II 3978 stigma 0.9-1.2 mm. Drupes \ iolet-black, subglo to ellipsoid, 5-10 X 5-8 mm; p^renes ellipsoi plano-comex to hemi-elhpsoid, rugose, deeph sured, endosperm ruminated. , ■ 6 . , I H i I ooth. Leaf blades 2.5 oules tubular, glal or 4, 0.3-2 mm, filiforr >,,,, 1 U'i "to e :r;!l::; : i:: J',./,, .up shaped J .'1-2.5, ies to 0.4 mm, tri a^da?cot1r^ther z^r-^z ^W^subglobose, Distribution and habitat. This species grows i >wn from the province . a. Here, it has been found in hum] "-C'^nUM r0 R!ient t-owm k g aI f 850-1235 m. locks at ele\ations , Phenology. This species has been collected wil 1 U '«'" fI„M..~ r.l.MU.N til,.. gl> M^,, ..!■.[ MO, P, TAN). 7. Gaertnera I .omber, sp. nov. TYPE: Malaria. Johor: Kluang, Kluang Forest 103°33'E, 1000 m, 25 May 1998, 5. T. Mal- comber3024 (holotype, MO!; isotypes, A!, AAU!, BO!, K!, KEP!, L!, QRS!, SAN!, SARJ, SING!). 1 I But 1 fat Biuxelles 29- 51 1959. 1.2-3.5 cm, elliptic or elliptic-lanceolate, apex sho [Equateur-Orieni.il. ' ; « d>o!.. hpe, BR!). vl.i (Bil. FHO. G is r J roi.fl.lMll.-. h«,^,- -4 oi E . U. S of N-soii" Rnei. 5 km tt of Sm»lx.ii '••' UiE ol El-.-i Vara Meer 7785 (WAG). , u „, .1 J.-ii .H ..U ihhi .s »,', V.2 ii 200", hl't RLi ,m.i iiu. t . hird, lobes 2-3 mm, ligulate or ovate-oblong to 21°16'S, m, 5 Nov. 1997, 5. 7 mm, glabrous or , 0.3-1 mm. G!, K!, P!, PRE!, TEF!, WAG!). liam., lobes L4-3.5 mm; anthers fully exst llaments 2.2-3.6 mm; style 2-3.8 mm, glab mIp !>i„>; intemodes 0.4-4.5 cm, with 2 !: elliptic or ovate to elliptic C11U1C ' ally, 3 to 5 fasent; petioles 1.2 c Republic J uu.i,ln, f u fr/.m- >.,*.. '.,!<, ^,„oU tl, ,u I.'.', J- 1 -,,m, .mlII. 'hi, "I >-,.!., J-ilnin ' '" "i'«»™'l..^ v - J«JI 1< t. 1'., > m"'.\,h,", ..itKuiat,.]' l,^'\.,iddl, fL",'\' 800-1000 m. (ilJCN, :or.]i j,.,uk.i.i. .Ui, !, I.- u.itr-.i range . The localities ation have not been recorded. f y ] Phenology. This species was collected with L,„, ;, /.'7; [MO TFF). \hhombe, 2iWh ,MO 1EF). JWi , in H ,n I. ,.,..1 ha , !.. ui r oil . .\|ii. II I . \1T M.„muu. f .. m-l '^bLl IwC,!,, mV^m "1 "'.'] n.l.i.iPnh,) nio.jh l.r . -IK I, ' , < . < t , / u ... 1 ^ if ^ ^ , ])hdes 4 ^_ 2Q , { ^_5 8 - in dlh.ti. >M ' ! ' ''"' '"' f ' i! , ,ll,,«.olH H X Ml" 10 .. .1. 1 I... |.t. .esedichei, inducing Mi! h,.>t- ievi,n 'i.,, '!^i" ,l ,' 1 ,:. ;""i „.'' prominent loi irising beneath the W. M..a (.',.[.,! SUM] l| u „ rl, l)U>l! 1 Sibat S glabrous, dning c >us or rarely 13. Gaertne & M. B. Moss, Fl. les, tube 4- W. Trop. Afr. 2: 21. 1931. TYPE: Liberia. bi. , fifo... > s u<, -« .in., iK.u 1 t-....ui I (t , <>L >[i ^ iil ( ()i , j([ (| n mM! |])(( , ]i( r< ed t0 drying pale 1 iam.; internodes 1.5- mm; 8 cm, smooth X 3-10 cm, elliptic shaped U al.sr, , 1.5-3 i n L.5- to h ts deltate, 1.5- mceolate, 0.5-1. flowers: calyx 6ntlre - orders, rathei deltate or linear to : .:. ; Marie, Mananara- rk, and the 4.5 mm wide, puberulent outside, L(ll ,|] Illt | nal , ,,„.],! tuliilk Bddum 11121 (k. MO). Bodji K* ., I,,/., inflated and cucu .rtly exserted, 216. 18 ssifolia, orth. var. sti mi 13-2 5 mil I) i jrl I] 1 ]> I || l](s ]__].) , - nn II ig to elliptic, apex subalobose oi didMii.A, .-10 V , nun ,. .. ... s t „ M \ iA lW|] A , K ( ( , |ma t , ( j J(l , (1 u Jlil|t , kj „ nate. through December. species. styled except mm, 3-5 mm diam.; Gaertnera cooperi was first published with an a s ca. 1.5 mm; style Goudot s.n. (G). MAURITIUS. -,. loc. Behmge, 101 (P); marcescent, lobes 2 or 4, 2-2.5 mm, deltate to hi Herb. Horns s.n. (K); Herb. Roxburgh s.n. (BM). principal an Aes, glabrous, sul le, subglobose, 2-4 X 2-4 cm, del 216. 1837. Silesia cuneifolia (Bojer) Kuntze, Revis. Gen. PL 2: 425. 1891. TYPE: Mauritius. Forests of Flacq, at Nouvelle Decouverte and on the peak in the middle of the island, W. Bojer s.n. (holotype, P not located; isotype, G-DC!). Long-styled flowers: c; Trees or shrub : to glabrous oi flattened, glabrous, 3-5 mm diam.; internodes 0.5- diam., put 1 cm, smooth. Leaf blades 3-5.5 X 1.4-4 cm, 7-10 mm, ong or obovate to cuneiform, apex rounded flowers: sii violet-black, ellipsoid, ca. 15 X 10 mm; pyrenes t< im diam.; internodes fissured, endosperm entire. blades 4-6.5 X 0.7-3.2 cm, linear-lanceolate to . «. ■ -J..1 tons; serondan icms Hat t- ,,i>.!xi, B. Clarke, in Hook, f., Fl. Brit. Indi. TYPE: Sri Lanka. Near Galle, G. H CP 2991 (holotype, K!; isotypes CGE!, G!, K!, P!, PDA!, W!, WU!). Trees, 1.5-3 othenMse teiete, glabious, 1.5-3 m nodes 3-10.5 cm, smooth. Leaf blades (2-)2.8-8.5 cm, lanceolate to ellipti Phenology. This collected with axi wrl. iMlu,e.! U...-S l (.J--l-ii(-15) X (0.1-)l is loralh common aioimd Hmi.luma pait.culub m ^ ^ ^ ^ f< ^ |imK)|1i (p ( , )b ; if k\' J'L M jjim [.,!,» K 1.1 ]« . 1I1PJ(U ,n a .! ' i(> fl outgide gkbrou '' >' puberu g inside, truncate or ; ■: ,' .-■,:■■■. Etam, 1891, H. J. ctotype, villous inside in upper third. Drapes violet-1 <>>' MM,' l"^ MTiPI i\l 1 x i h ^ ft*,' sphen.al oi hemisphencal, lugose, fmeh flss 7429 (SING!).] endosperm entire. •3 (PDA), / :' intermedia Ridl., J. Fed. his species grows in . TYPE: Malaysia. Sel Here it is known in Peninsular ;! >- Mi, is known from the found in humid TvLndaTn , c ;;x;t™ ,1 Abd^w.v lll.il l.N jU 5 unisexual flcn d by van Beusel .u.mIuoT; . acute ]..-, (li [. it nil .I. to M».. E i^» -. r '" " M M '"" )'" omJi!,) -.„ FtJ -j.j ,\ i L s\N '-INQ INegeri Sembilan: . ' ^ . ? 2 or 4, 1-4 mm, ,r to deltate. Boissier, Ser. 2, 1: 58 outside glabrous, tube 6-7 mm, 1.5-4 mm diam., glabrous, 1.5-4 mm diam.; internodes (0.3-)2.8-7 "' (-11.3) cm, sm. 5.5-16 X 1.5 Trees, 0.3-6 m tall; branches flatte ulate, acute; anthers included, filam. i upper third of corolla tube, 0.2-0.5 mm; style 9- chartaceous, glabrous; secondary \eins prominulous .^ mm ^^ j^ Q 3 _ Q ? mm . coro]k ^ g 5 _ winged, arising below petiole and extending to lobes, 10 mm; p)renes spherical or hemispherical, rugose. 616 Annals of the Missouri Botanical Garden collected but ha is ':.;.■ iUis,l,/l> < x,< ,t. A, U\ m„ styled except \.,/..•>,„ .,/.-,,/.,/., (I;,.,,,, K /.-. i:,*,~. Gen. PL 2: 425. 1891. TYPE: Mauritius. Forets is an attractive Figure 10C-I. of the stipule ma) or ma) not rune setae, but four lobes are cor: This species differs from G. hirtij lly glabrous corolla, .hh*. j-rinii^lim mt.in.U , t -, , aril ,j 1( L uf u , ll, , t .1 m (1 . al'l , F ■>> .1 , s 2-10 X (1.1-) (D H Lorence5 perg comm } ,-...•] i-.m.u-. j;):.'.....!!-.: -.-.•ok.Li' KM!- k >,ible and flat to prominulous abaxi omatia absent; ,. .tjob-sl i-l.-,", ■ absent oi 2 to corre i ated wlth ha l )iral lt 1 ^ >., lb M, , .,. ' ".:; . . ' 10 mm, ligulate to linear, acute; anthers included, 1914. 1 lastern State [Akwa filaments inserted ,11a tube, ca. Ibom]: Eket Distr., 1912-1913, P. A. Talbot 1 mm; style 9-25 ;gma 1-4 mm. 3391 (h< Trees, height not noted; bark with longitudinal third, lobes 2.5-3 mm, triangular to ligulate, acute; ridged. Leaf blades 8-12 X 3-4 cm, elliptic to glabrous, stigma 0.6-1 mm. Short-styled flowers: Phenology. This species has been collected with marcescent, lobes 4, 4-6 mm, linear to filiform; setae" fl(raers in A P ril and Ma J but has not y et been found 3-5.5 X 3-7 cm, branched to 3 to 4 orders, lax; biacts ,„ m „f Q, pa niculata, but, as noted by Petit (1959b), ieltate or trifid, 1-4.5 mm; bracteoles reduced; , numerous setae and pedicels absent or to 0.8 mm. Flowers 5-merous, the brown-black baik «ith piomment longitudinal shaped, 1 1-2 mm wide, glabious, tmncate oi lobes «hich lacks both of those features. Gaertnera eketensis is when open saberform, externally glabrous, tube ca. see comments about their separation under this latter 3.5 mm, 0.8-2 mm diam., inside Pilous in upper s, ketensis similar to G si i ;,/■ "■ ilJi.u, I M \iMilh - > . tlKinuM V,, a.,1 ,. il. .11. nl, i -. koiiiilM. - 1.. 1 ... i ere as doubtful as to application b n i x.l i . s- I ' i m . t . . ! , m, shows that G. fractiflexa is still extant at the type 375, fig. 8A-C " Sri Maha Mariamman Matang, Peakes Tea "" ( '' 23. Gaertne ill. ex Vatke) Mai- Missouri Bot. Card. 104: 386, fig. 2. 2005. Basiomm: Ps^chotria furcellata Baill. ex Vatke, Rutenberg s.n. (holotype, P!). 3 mm diam.; internodes 0.5-5 cm, sm, blades 4-16 X 1^ cm, lanceolate to elliptic or to 3 to 4 orders, r, 3-30 mm; 1 mm, with ribs 4, narrow h winged, arising below :at. This rneo, Madaga in, ■ where it [he Sai.n . II.... .1 elevatic J>ha<„<,s: fed with Phen 1^,1 I I., I ,ll{< - . , l, ]. '1.1! J .- ,A ill I.J >„ ,,h H„ Volume 96, Number 4 Malcomber & Taylor 619 2009 Revision of Gaertnera branous shreds at the leaf nodes and inflorescences can be found in humid forests at elevations of 30- 700 m. Phenology. This species has nsidered the co yjT'tir,"! ^ an<1 Wkh fmkS H-e.-K-, 1 • 1.. . TIN, 2001). TZZ^Z '/977 (K, P). .ndrazaka, Conn 1203 (P), lose calyx tubes. The specific < amber, sp. nov. TYPE: Gabon. Iwateki. : [), ■. o; c '. 1 .p. 8575 (holotype, P!; isc .types, BM!, BR!, MO!). JZ^Z^'V^V (K, WAG). Nyanga: 4 (WAG), '..„.. ;ardneri Thwaites, Enum. PL blades 21-30 X 8-11 cm, oblanceolate to elliptic- Zeyl. 202 I. it, J..r„ m .,,i.,l, ust'i, ,u 1 i| d m. lift i i I^NLIPE Sn Lanka, G.ff. K Thwaites CP 363 01...M 1 K.'- M.,^ . t.i(..l ,-;_ -, „ 04 _ o ( _ L2) ' S6 ' gkbr0US mSlde ' bu 2 9 mm, 1.4-2.1 r, third, lobes 3.5- [>,«.., Intl .H'v ,;,. ■ - wno-,e and fe. I H,,m l.x,.m,,l ,, nil tonj.-s ,o.Ilt.-,1; |,hIi,tL 2-6.S nun. F/,.< calyx cup-shaped, 3-3.5 mm wi«., oi,is,d, i i „ lo,e Auth indum l'l-2 jii hi iL.V.l... . ell lut. ], m n .1e >,).-_%) ... , ul l , , „, IIVI.uu V r , ,„ // lt l, lU . lioJi,!^' mi - - '. H.l.i.h H.ifi, 4.5 mm diam , lobes 3.5-1 mm, imh. is -h.»,h ^t-ulmj to lt| ,ex; ' apex «ith 1 oi 2 incisions. fissured, endosperm entire. sessile, subglobose, 1-3.5 X 1-3.5 cm, branched to 3 4 orders; b -2 mm; bracteoles / . ■ ■ o eel trur at. wet montane forests at elevations of 1400-1630 m. tribution and habiu U. This speci east Asia, where it i: : ak (Malaysia) sectoi -. Here, it can be found in Phi les has been co .. . ... . . ...... I ,itl 1 . : Discussion. Although this has pre\ioush beer '.'•'.. .■■,... . . ■ . '■•-.■■ .-'.•■ by their usually va, e based on a note on the type label. However, despite .■■■■■ . - -. - . •- el :ALAYSIA. Sarawak: Panjang to Ieku / 1 II Minw «. '), V , Hn.o,',,, 'i, !, M (holotype, SAR!). Figur Hook, f., Fl. Brit. India 4: 92. 1855. Sykesi grisea (C. B. Claike) Kuntze, Reus. Gen. PL : with indumentum drying pale yellow or gray-white; Jard. Bot. Geneve, 11-12: 115. 1908, non secondan leins , lo 11 pairs; Gaertnera guillotii (Hochr.) Bremek., 1963. . . ^ ; , 6 Trees, 2-8(-15) m tall; branches terete to quadran- below petiole and sometimes extending partway along guU _ ^ ^ ^^ ^ ^ Distribution and habitat. This species grows in sh. rous, lobes 0.1- Southeast \sia. «heie it is kn e, elm ate in bud, Indonesia, near Sim diam., inside villous in upper third, lobes 2 ir or ligulate, acute; p*. i tu- • v, v. n * a anl 0.5-0.7 mm. Short-styled anthers shorth exserted, filaments 2- tyle 2.4-3 mm, stigma 0.8-1 mm. Drupes Phenology. This species has been collected floweis Mss though Decembei, and with fi coriaceous or chartaceous, glabrous; secondary veins 5-12 mm -.jili t., rt .il m.l.lV ' J Ml " s ' ' " " ' "' ' ' u ' ' T " • ''' ' ,M " n ' r " ' "' ^ u> ,v sv ,\, '", 'i „'" i! .'.T.'.Wi", 1 . sl ! ' » ' 111 I I- it I u In. I li l\ , liiMt 1 ^n.l u H s ^ uli in s diaiactenstjcs of nocturnal 1 29. Gaertnera liirtiflora Verde, Kew Bull. 37: ceae flowers may be nocturnal or crepuscu 535. 1983. TYPE: M Cilhems well. Distr.: Macabe, 540 m, Aug. 1980, C. Puff . M i URITIUS 800825-1/1 (holot; (MO), Vaugi Lorence 1541 (MO). Trees, 5-7.5 m tall; branches flattened to t, glabrous, 3-5 mm diam.; internodes 1.5-3.5 cm, smooth. Leaf blades 4-11 X 1.5-4.5 cm, elliptic or 30. Gaertnera hispida Aug. DC, Bull. Herb, oblanceolate to obo\ate, apex obtuse and shorth Boissier, ser. 2, 1: 58; i 1.2-8.5 cm, smooth. Leaf blades tnan ^ llar: ' 5.1-16.1 X 1.1-3.1 c or 4 , 0.5-2(-4) mm, deltate or linear. Inflorescences ^^ ^^ ^^ ( or peduncle to 5 cm; branche ;ascar, where it has been form, 1.8-8.5 X 1.4-8.5 cm, branched to 3 to 5 orders, lax of Toamasina in Mananara-Nord i s to pubc lobes 2-3.5 mm, triangular or Hgulat, tube, ca. 0.4 mm; style 9-11 mm, | 0.7-2 mm. Short-styled flowers: simil; except calyx 1.5-3.5 mm wide; coi 4 mm; anthers shortly exserted, filarr style 3-6 mm, stigma 1.5-2.5 mm. black, globose to subglobose or didyi .i.:.. .- , ;:-l l real, ru spec tloweis m Octobei, and «ith fimts Januan tin, ., Ji . i • jjr < ,1 _ 2- J mill i I. Ifl.^ April and in December. corolla tube 3-4.5 mm diam., lobes 3.7-5 mm; Discussion Gaertnera his ida can be rec nized 3 " 2 " 4 mm ' S ' yle 5 " (,.-,:/,«., '. vf , 4 V . ,<" ,s /",<, D' 'i « il 1 1« m.asm. I! 1( . n j, i mud .., I.umi \»,- h],-,,,!,,,,,, „ (K, MO, P, TAN), 3346 (K, MO, P, QRS, TAN). narrowly ellii scence bracts and France 45: 352. 1899. TYPE: IVhdagascai NE dei "^ " ' "" n " M il »i.il i - .clditjm.l under that species. Trees or shrubs, 2-10 m tall; branches flattened A!, BR!, G!, K!, P!, PR Ho„ei, 5-meious, heteiodistslous lose with indumentum dnmg gia^hite becoming "'''II,,*!,! t iiiiju'l' 1, lJ l ISl W,1 "i 11 V«- I " .uimu Massif M ,»indrano up 3, 0.1 ure 8F-L. Up sl^p I 1 V-2 5 1PP V V L,U r ' ,. Jard. B irtHH' .Unl.u.l .h-m.^n .r,U"ii..rm. outsiJ. be 7-9 mm, 2-2.5 mm di Hgulat, Tr ees, 1-6 m tall; branches flattened to tere ; glabr ' shortly 3 1-] i.i- »f!-] urn, -,!„>,, ,',/,, 9-10 mm, 1.5-3 mm di. . 1 1 ■ 1 .1 th. Leaf blades 4.5-24 X 1.2-8 en Phenology. This species has been collected with December. P.(,..*.,',,, iIA0U,V-< \P Viilshauaiia: Vo.l,,].. fo»,s 'ftii ( p IVM * li 1 I. \nkob.hmiPeik, lizi diam.. glabr. res 2-10 X 0.3-2.5 cm, : 4-12 mm. F/owers 4- 3essile. Long-styled flow- £:;: corolla white, clavate in mtside glabrous, tube 3- "--'»'" inside villous in upper anthers inserted at ca. middle of styled except calyx 1-1.5 shrubs, 1-10(-15) humid forests at elevations of 150-500 m. Phenology. This ^ pn)c Lmn goc ^ u2 : ■■■■■.. rough March. PI. 2: 425. 1 Discussion. Will iidL> F1 Maky Penm - 1 ['LA ,-r t 1, - , „ .1 u (holohpe SINf,') . , ; . , (,„ n „ n i I ill lull ill Inf. ' t I St the stipule lo : .• . . : ' ' ' ! : ' hagmentmg t. I.. i- 8- ged, arising below peti it, lobes 4, 1.5-7 mr 34. Gaertnera juu: ed. Ind. 2: dal, 1.5-15(-18) X 1.5-17(-22) cm, lax, bran 383. 1856. Sykesia deltate or trifid, 3- 15: 388. 1967 [1968] \\V) Iml ,n ji -inn- < i •< -li [>< .1 2-]mm wide, outside glabr , oi pub. luhni tube 2 -.-1 mm ] 5-2 2 mm l|f ' " m h M I hl ] iui b(! '' 1 !f " ' 1S '" ' '"' '™ tl ' ' t 1 >_i ' " h I n! i d nl 1 1 » ' i b\l I SI?. VT 'I J \'_ ' t ' l " '" ' Y tj " 11. ' .mbdkRrvnf Koihummeii VRl^^K, I V. k. Kl I'. J™ 35. Gaertnera kochummenii Malcomber, sp. nov. ose, 5-7 X 5-8 mm: anu: 10th Mile > Dun " ns of 0-1500 m. pubemlent with indu- . : ; Phenology. This spe TYPE: Cameroon. Betw. Tabo & Akoi [amfe, 2 June 1975, R. Pamtypes. CA Mamfe rd., betw Letouzey 13702 (' ■ ,,, , HIV. IV >■* <* M,.U, •* „,., _'■• kn I\\E ol M.,,-^ U L.l K!.MO!.P!.V \i p '" '" T »1"»"'W 'I' -A ll laid R-H Lui Btuxelles 32: 186 distinguitur. Shrubs, 4-5 m tal d near stem Belgian C Republic of the apex, otherwise teiete to subquadiangulai, «hen Congo]. blades 16-33 X 7.5-13.5 cm, elliptic-oblong to G. W. J. Mildbraed 3206 (Rf).] --. r 2 ]<>< i i , in s ;I / t.il.lu 1 I .ii ' 1<*> BR') iceolate or obovate, apex cuspidate or acuminate, ' 'Vth h'nf .m' m ! ' j'T' 'l» '-^ '.T *""' '^'" ""' ''' 1 "''l'- 11, ^'" , ' >* ^ ^ "^ n ti„ ...oil , -MnK is ^"^^ I^'," 1 !,," 1 ,"', 1 , ''iJ., ,(„,'' i^Vlli and stigmas unknown ih , i >1< t i 1 , k I. 1 > . , | ,t 1 ' I HI 1 erodistylous. Long- lounded, antheis uuJnk.l nl,ni-iN jmIuI i" •,.; M d 1,-1. I '>-i . m 111. ij, t, .1 1, ui, ... antheis fulh exseited, filaments 1.5-2.5 mm; style petioles 2-8 pubescent, drying 2.8-3.2 mm, glabrous, stigmas 1-1.4 mm. Drupes , tube 7-10 mm, with ribs 4, violet-black, globose oi subglobose, 6-10 X 6- n petiole and extend- endosperm entire. ca. 9 mm, linear to filiform, setae numerous, 1- Democratic Republic o ion. Here, it F " """ is found in humid f. if 470-1150 m. [ Phenology. This species has been collected with { flowers June throug i fruits January j through April and July through December. j Discussion. Gaer similar to and '' may be confused w itral Africa, but ' r, lev, .ithn.u .W[BR P S U„ E, iln Mai. I i < Jhi.i , .jr-r *> . " ltJ ''^ ]"• I) - <- "I .'ll" i '1" i al '1>1 / / / ' H. K u, , M - tl „ , / , , > | Mj , l, H.tJn.i-.i. , H <,ill il hi p. , it, I a •m. Kew 1937: 62), this name was validly W9 published in their 1931 work with an English diagnosis; the ICBN requirement (McNeil et ah, Cooper 277 (holotype, K!; isotypes, A!, BM!, Bojer s.n. (holohpe, P not loca FHO!). W!). ubescens Verde ck River G mnated b • i ll,.,l ,l,~l, . Thei 1.5-3 featui. .ation „l!, t , *,:, .us, adaxialh 2272 ' m,\' 1 ,, >m','i- 10 - ^^'-^ii-i-alwiigH-agiiialis (Schweinf. ex Hiern) t ™["''].u, T, l M V FltlS B !l Jj " ! ,5,,t SA.tFJlvell ,2'' chotria lon t , .ill ^ !,. ,. V 1.1. .us ■ „ h,l.„ M x K' ut, d Trees or shrubs, l-6(-8) m tall; branches terete, ■' ■ .11 .l.tiisf .,.ti, sh.1l! lul.ii.t ms.it. .I.ii p ,| osu i os , j nln£ , menl ] )lan t , -,,„ 1S t ,,], ■ ...rolla tube, 0.5-] mm: shle ]8- p „ slst ent ) )ase f_2 mm. tu 1 . t .-A mm i.h i.b, led to blue, obovoid to ellipsoid, 23-28 X 12- g ( Distribution and habitat. This species grows in hii 6.5 cm; branched oblanceolate leaves rolla white, clavate in bud, when open ; Lorence 2224 note ce. Verdcourt 4-6 mm, 1-3 le villous in upper i,mLHi« ,i i'. ,< \ -idr Ihis \aneh v is onh w -n 1 m upper third of similar to long shied except cahx 1- m.Lli. i(b^ m.Mih, »|»IIjii rui. ~li.ii," ■ ilu» " '"' " "''''"« Hu- m -lv j, , , ,,, raafe differs from G. vagiimns in its tubular stipules Phenology. This \aneft has been collected with rli.t.lr nuii.ln.i.oM, . I,.-, h . mw J ,) t] 1( *,« aid 1J( , )C . „, | |„ ,| ,i,M>, Jiout the year. l')V».,) ll.i- • ...,h ,- mniIji to « A,,/,, Bull. Jard. Bot. E 1,9. 1959. i ...bli- ..I ihe t'ono.,] Ki.m pnBnc7)F'TH£ Lauren* s.n. (holotype, P!). !] WOl Si V 1C R] plagiocalyx K. Schum. digital image available ed. 322. 1900, nom. mid.]. TYPE: Belgian Congo ^ _ ^ ^ ^ ^^ ^ (] . [( (|K .^ ^ j ^ ( f __ .^ |f lie- .. *-"i,tiJ-, 1 -I..-,'. ) n, mil ../.-/ />c 2- -..-.)! „/,; ,BJ. HU>, L .. n .l.iwll. f . < -s - > J r FR 1 tin -)5.5-18.5 X (0.8-)2.3- lliptic-oblong or oblanc , 10-15 mm. Stipules with tube 10-28 MA \u , , , .'» luiv,! .1 ..ltK i 1 VIS.m K^ UH\! \ IU1NC1-, <>OLM] J\/.. Noum 1 , <>, 2-3.5 mm wide, lobes 0.1-1 mm, trit .rolla tube 4-6 mm, 1.5- -.UKl.v, ,t.J filaments 3-3.5 mm; style 3.5-4.5 mm, stigmas 1.5- Madagas< 2 mm. Drupes 5-8 X 5-8 mm. West Coast, near Antalavia, 0.8 km NE of «;=,-., T onno H™ u „„„ k„ f„„„,l ;„ u„ m ;A ( nraaic , „t Figure 7C-H. with the blanched poition 1-6 cm long with two or . 5 cm, smooth. Leaf - ,.r,!,H ,— jrous or less c tei P"b . loi : tubular, glal f,tl,-n I., f.^1.1- it, ti. u. 1i,U _»>-i>ti mm ,, -im raa.-roI.otrv-. F! >k, ered, terminal on axillary isotype, P!). I, lades 7.5-26 X 3.8-13 cm, elliptic- tubular, inflal d, glabrous, dning deltate or trifid, 1-10 ir >us infloiescence and iefle\ Madagascar. 6ntlr6 ' Distribution and habitat. This species grows in 60-1100 m. ; • W\!«Vi;\-( \F, l,.AUiasina: Manaiidia- I u. t;pt>le- tha .■.c-r third; lobes 2.5-3 mm, upper third of corolla tube, 0.5-1 mm; style 9- rugose, habitat. This species grows in ad. It lives in humid 43. Gaertnera macrostipula Baker, J. Lin, Soc, P^gy. This species has been collected with K..t. P« s < .i U 1 l_- I. 'I iilr Dt ,rtl ' "'* »"»',J.iin lu . hMiMnlu, r,ml922 November and D . i : .,, ^u.Ul then shoith cuspidate oi acut. In. ^' '»-< ! »<< -^ - '-"' Thp t] — are iSO (MO P) Toamasma: Alantadia National Paxk Disunion Tin- -pi-ri, , « <.- o.i -Jiuill p I'l J l.J J. ,_ ( ]\Ph '',"'". ' tl "' K! )' densely appre ;rminal on prin. '77 (P. TAN). "! f; 5 ^ mm :ii'! g ! S l 1™: 4 5. Ga,,-.,,,-,:, mirmphyllu Capumn ,a Mnl.-«,m- l' iM.is Mn..o, -n-t B»t Mj- ..in Bot. Card. 104: 390, figs. 2, 6. 2005. TYPE: Madagascar. Toamasina: 6.6 km SE of Andasibe, Malcomber, A. P. Davis, D. Gower & J. A!, BR!, G!, K!, LE!, MAL!, MAU!, P!, PRE!, 3!, WAG!). b, t i>, >:,n unci habit al Tins species g 10 u Madagascai, wheie it is widespiead in the ea- escaipment. Here, it is found in humid e\erg il<. k hi lil mi .nth. - 11. Ii I l<«iti> 1>- 1 I ( < f < U I.-J'j.™ si.i/dllil, , .„d.. wo,. ,..!„, .ni.irhi.uir.i., ;''- 2 -;7;;;^^ ' -'■-- ! ™'- '■-- '•- March through May. apex and its stipu plit into four bud, when op. »us throughout, tube narrow, persisten n, 1.5-2.5 mm diam., lobes H5mm, .-, I [<- EU" S I" ILT" \Si,l,iii Hi.h-.s. ., f „ > ; , ,,, 2 , -, mu lai to 6 ure 12A-D. macrobotrys, Med from it by its 47. Gaertnera obesa Hook. f. ex C. B. Clark, Hook, f., Fl. Brit. India 4: 92. 1885. Psychotria ["' obesa Wall., Cat. n. 8328. 1847, nom. nud. j""' ntze, Revis. Gen. PL 2: 426. 1891, nom. superfl. TYPE: Singapore. N. ^ reduced; pedicels absent or to 1 mm. Flow, obo\ate to oblong, apex acuminate to acute, base tlmd, lobes 1 8-2 5 mm, ligulate, acute; staminodia nous thickening; secondan \eins prominulous abaxi- 1.3-1.5 mm. Staminate flowers: similai to pistillate ally, 9 to 13 pairs; les 1-10 cm, except corolla tube 7-10 mm, 2-3 mm diam., lobes d habit, ' ar, 3-10 mm, - lpP er third of cc ■ • ■ ■ grisea and presumably a natural hybrid. ± rugose, finely fissured, endosperm entire. .... ■ h.- Boi™ .,„ !, - l,. 1ados.ru (SING); Up PVm/ .-, Tin. sjjvk, ha. I. .ii t s f .,i,l S>» 1 , terete ,» ,, .1k-u ul a lu.'< u- 2-0 mm iuhij 4-ll'rm. ol-Ki.eolit, , . -.Gh >- ,,^ , , -,ul «, lo I'-iv.,. -I, piotologuc I acuminate base attt.ui. ie lo cini^fr ,|, Mll , ,.„, - !t '"> '' --"' ' 1 |, n, ]„i ... ffon^, FRI 2891 (KEP), Malcomber 3021 (MO), 3022 (MO). Phenology. This spe 49. Gaertnera obovata Baker, J. Bot. 20: 218. ^™ 1*™*? throu g h March a » d Ma >; ^ 1882. 5yteia untze, Revis. Gen. PL 2: 425. 1891. TYPE: Madagascar. >ll -r.fl.k ,]li.,t„-(»K ,f< < |lj|.n. - ' .si- ovate, apex acute to rounde Inn , il. II 1 ' >i .1 malh shbious or bleand flat . pn oil do is 5 to L6 pa domatk 3X margins and , rs, whereas \arieft ''■' : " inside villous in up} e 49 a . Gaertnera obovata Baker var. obovata. lo .dJsph. -, ..Ion*, .nut .ntn-i- la 0.8-1.6 mm. Short -st ;■ h mi j.h.n. .1 Inflorescences with ped 3.5-7 mm, 1.5-2.5 mm diam., lobes 2-3.5 ligulate to elliptic-oblong; anthers included, filan 0.3-1 mm; style 5.5-8.2 mm, stigmas 1-1.6 1.3-1.7 mm; style 2.8-5.2 mm, stigmas 1.7-3 Drupes 4-5 X 5-6 mm; pyrenes hemispherical. lana, R.-J. Dufournet s.n. (P). Toan 7 (P, ' f 10-2000 m. October through December. Rrj^ntatnr s P ru,nms namlltrd MADAGASCAR. Madagascar. Central Madagascar, R. Baron 1243 ^alTen P r< en rib Trees 1.5-12 m tall; branch**. fhh< .. I -k ,l i.K t,i im, I.,-i r ,Llir ,,1! N , . t ', > . t> h< gular to terete, glabro er 74475 ( TAN )- intemodes 1.8-6.5 , ides 4.5-23 X 5Q Gaertnf ^ M R ^ ,ip,! ^'"^ '""'^ ^ h " ilh " '" (llt " I,.., II J 4_T> LilUl lU'E Nii,eiid Grand ^ Bassa, 5. Fo^e/ 72 (lector? pe, designated here, jairs; >; Trees or shrubs, (1.2-)2-9 m tall; branches flat- bracts 0.8-40 mm; ] , 2 mm. Long- tened to teretl reus or puberulent, ! _ ,b __ - k > 'I , in s, u , , tl, , i nl, ,!„>,*!<. Ijeaj blades 8-18 X shortly exserted, fil m; style 3.3- 5.8 mm, stigmas 0.8-1.4 mm. Shorts-. U< • similar to long !yx 1.7-2.2 mm; Drupes 6-10 X 5-10 mm; •1 , ; '../,', I 111, \ llH* _!«■> - 111 \„Uii n n. 1 .no. \nt,ii rt n„iji. Ku.ui.-ntsoa, Toamasma. elections of 0-1600 m. Phenology. This \arieft has been collected with Discussion. This \arieft is similar to (kieitnera -..-,■ •■:: :'■ ked there. 4 mm , l_ 2 .5 , Ulcus in upper third, lobes 1.5-2.5 mm, ligulate to linear, acul shortly exsertt ed in upper third of I ,0 TTT ->i)H) (MO TEF) 2<>2c (MO TIT) J- lmU\ ti 1 1 1 u.l . im , \uu.k I., nm FmlmaFw,. . im wn, I I,, in. M.I., Nr..h, ll^ Fi M , r l,h, «1 (1 ;, Togo, and Zambia. Her v , J ,f |,[«|e, |_; n, elliptic to elliptic-oblong or oblar, II (-.„,,•(■, iH-K; K F \T if h | t ... PI -J ' 5 f ,<-- i ) T \ \< ^1 il p ! RT .11 Madagascar wheie it Antsiranana and Fiai 5 known from the provi] 72549 (BR, G, G-DC, L, P). SENEGAL, s. loc, Perrottet <. \l<) >\ \, Di, ,.htr:>» , 1,1 !•«',>:.,' liu - ~, { „ r les £,t,.v - m Gen. PL 2: 425. lauritius. of 50-650 m. i S °'" ^" ( h0l0t ^ e ' P P^ofogy. Tins species has been collected wkh fi m ., OT o ;„ t„„„„,^ r.u,,™ „„,i n„t„k r tt,™„„t, 8 cm, smooth. Leaf 'blades 4.5-17 X 2-8 cm, elliptic lo , absent or present, often crypt-type; petioles 3- 53. Gaerta Baker, J. Linn. ceous, caducous Revis. Gen. PL 2: 425. 1891. rpub. ' lent; ts mini. :;-i , plahrou, . : d xl.a Z-^ till,.- JO-2', mm. H ">--! mm hm. . I»l . s 0-10 mm haulue oi mate-Mo.u .nth, is ,h,uth , -ti r, 4 oidti,, conae,l,d 1 1 .< I rliiif K lu,.,i '- Bo, n >T',t) ^ jih ,lu tJ n < uil, , Elections it corded as glabrous or pilosu] ibrous inside, similar, the i that sho« all the ■ • ■ . ,■■■■!.■:■• ' ca. middle or in upper third, lobes 1.5-6 mm, d only recently. 0.4 mm; style 9.5-16.5 mm, glabrous, stigmas 1.4- K!, P!). The latter is selected 3 mm. Short-styled ;ause it is a more complete and til ,. i hi-, i In, vb.miu iMiiiih. to -.Hitai, ,1 k, V,. j IK U tt,j, i, , u, 5 mm, filaments inserted in upper third of corolla Gaertneru irently hybridizes :■••'■'-:, ■■.■■■.;: rugose, finely fissured, endosperm entire. hybrid betwe ebia and either G. huwioz* I- 1M -I— f „ >■ » V ,"5 ; tlil ■. Md. .,■„ 'w„,k K, , rl> ers Januan thiou-l. M -,< 1 iul 1 , u- th-.,i..\ «"'<"' -'--< ; P> ''■'■■" iMt P,. ■< "'"Mo . " D' ■ r ii i I | * t 1 1 1 w II "^' * T «"' rlllevj i»hjll«> , -*'lMlii K.)l-r 't Kuntze, Revis. Gen. PL 2: 425. 1891. TYPE: scar, R. Baron 1920 , , llii!(l|1] » li „l,., 1 „, ^K,,... !m ,m, „ll,,i s„ti- ihoJohpe. k>, isohpe P!). Figure 14A-F. ..'■•.' see additional com; sly hispidulous to m<, j.'.-i , n ;itj, ,nM,t,t„>\ - ],h>fi- e , < ,,1 pil Irs, „ ,,,1-, ) 12 deltate to linear. Inflorescences cmose. mam -flow- 2.5-3.5 mm. Drupes violet-black, suljgJolx.se or di- ered, teiminal on axillan blanches, pilosulose to chmous, 6-7 X 7-8 mm; p.renes spherical or biform, 4.5-14 X 3-19 cm, bra ?rs, entire- lax or somewhat congested bracts lanceolate to Distrib ution and habitat. This species gro«s in Phenology. This species has been collected with glabrous inside lobes ca' equal in s 1Z e or 1 lobe fW ^ m Janua1 ^ Februan, and October through larger, 5-9.4 mm, '- and ™ th f ™ ts J™Y thr ough ^ and » ovate, white; corolla white, clavate in bud, when open December, infundibuliform or saberform, outside denseh pilo- Discussion. Gaertnera phy llosepala is an attractive upper third of corolla tube, 0.3-0.7 mm; style 14- recognized h } its denseh hispidulous to strigose or i- oL'.o.i, U, )i,l,Mi.l.nr I n, h.* hgulate 01 elhpti. -oblong, a. te a les, cahx lobes and biacteoles 6- ill corolla tube, 0.5- tmncate or 1- to 3-lobed calyx limbs 1 mm; style 11- tigmas 0.7-2 mm. 1.7-2.5 mm; corolla tube 10-16 mm, 2-4.5 mm liam., lobes 3-4 mm; anthers shortly exserted, 2.5- ; «„.)'3/.'M( I\My„ ■' ,', 'iMD r< \I„i,i i - ^.lieucl >i u-nij.i.limnl = u j, >*< . i.n.b J.xxiu. -! \\ l fiUJD ] PIS, ' , P /' 2 Mvh .. ii M. i H i, I uinn i> ,.i V „ t elev idagascar. s. loc, R. Baron 2327 ber. igh May and in (lectotype, designated here, K!). November and December. Trees, 5-10 m tall; branches terete to flattened or Discussion. The calyx . vV.lu^ilii, I I>'c->is M., la.. 1 I i . 1 uh lobed but the . le ai ,11. |„-,,,, tent, tube 10-35(- inged, arising ben< as bracts'. This IcU - tJ ld it «■-. plants with relat ^ese structures nre interpreted here xpeciex ix moipho luous stipules (e.g., Miller & Miller avimbahoka et al. 69 [MO]) to robust ively large leaves and larger persis- )es 1 c ■r 2, 1-2 mm, delta'te to lin ., Schatz & Miller 241; ; ml i [. - peduncle 1.4-4.5 n, 2-10(-20) X 5 .rders, lax or somev to ovate or trifid, ous, abaxially glabi nceolate to ovate, 15- _ L|PlO]i ]| , riu- -,>■ i-, i L- nix J lv,-.. her characters. collected in western forests near ( i n, hil ■ds al.-.-ul : or to 2.3 mm. Flm 1477. V). On**, zhya is similar to G. loUx I.Kx , . Ihj.t, «, ... . *h "IJl[H .1 rlJl, t. M.U, S I ,1. .11 psichotrioiJes (DC.) Kuntze, Re\is. Gen. PL 2: 425. 1891. TYPE: Mauritius, s. loc, F. W. Sieber 57 (holotype, G!; isotypes, E!, K!, MO!, OXF!, P!, W!, WU!). Figure 15A-F. rois Hots, W. Bo ertnera psychotrioides (DC.) E chotrioides DC, Prodr. 4: 531. 183' J.C.Pu.Ji ') .54 eM.niaJK »lal)i..u , , ox.ljj, Ui-nti ei a psychotrioid.es .nee ou Bourbon," s. migh t also b edentate, but can be rf C ^ 8 oiform inflorescences fer flowers. ,-■ rees or shrubs, (0.6-)1.8-12 m tall; branches it 1 ; In lljpur .I.Jon. Iln.tif kin- though o^eilappmg condition- u late, or elliptic, apex rounded or acute to . , g to t he species ""i' 1 ' I's. .i.iu-,1 m «^ I>7.i <',.,.,., r.ni.M'uptamvll... tlu l-hu.' I. . . , t t ^ LI'' !• m '->l-.S! m Mil. W flattened, glabrous, 1-4 mm diam.; int lowers in January, I rid December, ind with fruits February through June. ( , I t , „ / ,' s ,,.- , In -II , acks setae at the I tipules tubular, glal inged, ansing below petiole 4570509!; isotypes, BR!, G!, TAN!, WAG!). s lediued; pedicels absent tPlelP ra '['^l.,n M ,l„. .J.,].,,.,,,. .'-J mm -I, >,u . 14) X l-l(-5.5) cm, elliptic to oblanceolate or 0.5 mm; style 6-10 mm, glabrous, stigmas 2-2.5 mm. * _ ' > '' mm ' vl ' ' ' alK " - ll "'<-/'> for related species and their distinctions. Van Beuse- upper (hird rf .^^ ^ |} ; ._ 5 mm; gtyle g_ , ,,, , -s -ubs,, , m„h, ,w.ui , \li'.< ,oi, f .,.„„..., ,s sinuLi |o lijf „ ^ )iUi| t%(| _ p| ( jK forms of G pmqhuhmana. it is m ,_m -d h ■ - as ., j ._, ^^ ^ ^ ^ rj ^ _ ,_, . ^ ^^ »« '«* *>* T IT " ' f ? (A, K, KEP, L, SAN, SING), Henderson SING 23390 (BM, ' lanaianNoa and iotuia, m th. i x .i... „ ,t «.„, \i,..iin H1(> MNu. rmseA Hill. Pmsegbve 4227 (A, K, L); Pine gitia, and \ndol. Itt b N >1iu< R, senes Heie, it is /aHe 4876 (K). found in hum cms of 150-1150 m. 58. Gaertnera r, sp. nov. Phenology. This species has been collected with «,V,/,,v ' i<;,;,,tlu ,s s Hlij.ti' l..U',jLU,i 1 x i 1,1 "- mm Ion. W i !. t.mi.t i, '.,in-i r:,i,o>,> ■ Lmi ■ 875 iu (An •> A tit 1 MO ] I ' ' ' ' • " !h - > - \ i i>t * , „ ,. -> /j, 1'T^lir s" j tl(!l , (t 250-H00m. 1472 (P, TAN, TEP ahela Reserve, J,il~ ,V\ \%, (/, ,„ /„, JJ7_' (MO Phenology This species has been collected with 77078 «■ its ridged int. ,ces reduced to a 3- IVn'lj) ki.nL.x. R-ni- C- not seen, BR!, CGE!, G!, GH!, L!, P not seen, some other species of Rubiaceae. Walker s.n. (K not seen).] Kanneliya, W air rreelets or tiees, 2-5 m tall, blanches teiete, MdmllkL ™ dttd T««^'«-' (E, GH, K, L, MO, PDA). 1 flat to prominulous abaxialh, 3 to 6 pairs; located; isotype, G-DC!). ig in basal part of 61. Gaerti .mber, sp. nov. TYPE: lear. Inflorescences Antalvi E, Nov. 1989, G. E. :, many-flowered, Schatz : 3-3769600!; isotypes, Figure 2. UK. lb U.lv i,-s ■ • lit <■ -n.iiUi to long shied alp impa [late, 2 3.5 - id, , oi s.i. ■ . h-< it J , 1 n, v .. t«,.m P mas, 1 u species is named aftt of ca ' 10 m ' the type specmei .cant contri- Phenology. This species has been collected with Discussion. Gaertnem schizocahx is not well 62. Gaertnera schizocalyx Bremek., Bull. Misc. The species can be recognized In its subglobose. Inform. Kew 1« He, sessile or \en shorth pedunculate : isotype, K!). is similar to G differs in its smaller :.'...■■ ■ • , ''.':.' 2.5-6 mm diam.; in smooth. Leaf s 6-14 X 2-5 cm, elli rf.l.M.JU -ldllC Kavu. Maws mer s.n. (SING).' ;:' , ^7- , \ .:'"■ 63. Gaer ■ ndahl, Sibange 1 inn 5 i Aug. 1879, Ti^^uV'lTi sotype, P!). d,ym § TuteTo-^l";?; zp:ir: (hc.lc.tj ,':. .- Bi er,inO i ... Afr. 1 (1, pt. 'or with 2 incisions, n l-1'fs.fllt W.«- • shrubs, 1.5-8 m tall; fcroncfces terete or ,e, 1.2-3 X 1.2-3 -, tteie cited in tht ]iiot i II. Souun 17,", 1 1! .V-liov.1. ' rmolla tuljt-, . tert . at sea level or very near it. 4-13 X 1-4.5 cm, elliptic-lanceolate to elliptic or floweis \ugust thioi h (i til.ij .I'll >,ilh t , lis ,11 d tnud e ° c,mP '' e - <-nm? r > dr actous, y .« n.ii,- , <)i 'st,in[. ud.li-h I id, u. .,i ■ :■■..... : r ■ . ; . is apparently enden is ca. 16 mm, with ribs 4, narrow h winged, arising i— 1.5 X 0.5- tre Ha; the name P. .,! jvdl-L i.. i..".lw.« -|. -.n.lJ-1 t. ail ill, .111. I ... t -|.t M 65. Gaertne dtes, Enum. PL Zeyl. ( .ml.xiji TVimsuhr M-ln-ia, Thailand, and Viet- Near Adam's Peak, G. H. k. T'n, i 600-1800 m. Phenology Thi 'ected with GH! ' K! ' P! ' W! ' WU!) ' [ SYNTYPE: G - H - K - Threes CP 457 (K!).] figure 14G-M. through April and t ember. Gaertnem walk Benth J Proc Linn Discussion. Van BeuseLom (1967) considered !;;„„'„,;,, ;";:;.,'' „',i, ,' '!,'. ' i ■■" ' ■" l " "'''"''"' u '" ''■"■ i '' ""■"■ i ' "■' . IV. . inn-,.,. U-IwjiL-.I- /. B. L. Pierre 12, J), /. B. L. Pierre for this species by Pitard in the pro mgitudinal ribs. Leaf blades 0.5-2.5 X 0.1-0.3 c Volume 96, Number 4 Malcomber & Taylor 655 2009 Revision of Gaertnera Basionym: styla Hiern, Fl. Trop. Afr. 3: 213. 1877. TYPE: W tropical , ill. jjdli,.,!!-. Africa, Mt. John River, 1°N, 1862 1791 (holotype, K!). Gaertneru Jink^ei K. S.hun,. Bot. Jahrb. Syst. 28: 88. h i: style 3.5-6.5 mi Drupe! ol lack, globos, ■u-. i;-a f.-8 mm; pyrenes BR!, E!. G!. 1 !. uTI!). / s j ■ d "» ii nl I .1. i Fl Tie,, Vh 4: Distribution and habitat. This species grows in Sri !, P!) Lanka, where it lives in wet premontant ,mi' Trees or shrubs, 1.5-3.5 m tall; branches terete or Phenology. This species has been collected with isT alractive its itlatneh small 1 „.,t. 1 ,-- . - 1, u. , i,nl 2-15 4 mm 10 ^ bronT^ de^ety entue oi «ith 1 oi 2 inosioi.- mi.cr-.<'i,t If I., -, I C °Van'Beusekom (1967) cited the Th.aites CP 440 ' •' ' "" ll " . .I ,u^«m..I..iei, Ij-hloi.., L„ ^ ,h/e ( //?o« ere: calyx ...b,.',. mi Kl.uJU.H,,,!!,, ,.,„„ 1„, ulloui, ,.,,,„,,, „,,„,, ,1 , KHln 1L1 , lH Ml , . . ..- '- . \.s s.l..L,o„ ,.j ill* ,,m- l.-.U.t M »- 1„1,^25 „l,l.„i« .1 /.'.,,, 1PIH /)j; ^ V| b]i k 1(|| ^ n L j o 4^ e 6 _ 9 x fi _ , MO, PDA). ' ' rugose, dee ,erm entire. Phenology. This species has been collected with December, and vjfi, uml- in Uiu.u. I synonyms here. deltate. Inflor. any-flowered, termi- m.miio. s (upU i,s \u -,„ ,11 ql ,1K , uj.hi .. M i„ u- , ,-j| o- i. i.ml --"-. on 1.. .1. '. < and exemphn sptcim.ns is with most coll. . h», pmh ,n uinl.it ,m ,. , wii.ii.l .] (2 5-)10-24 X there along with the general Rubiaceae collection so it 5 mm Flou ei s 5-meinus. heteiodisU lous /^-sh /<■ >' <-i 'A <■ - -b V 1 i)-lm n " Wi rnl .p 0.5mm: V ^ ( " ' Kft " >ex) stigmas 0. (-1.5 mm. lort- ^ai;.. ' _ s ,_>- „ 67. Gaertnera vaginalis (DC) Merr.. Enum. Born. ™ olet - black - subglobose or didymous, 4-8 X 4- ■• DC, Prodr. 4: 520. 1830. Opbio.wlon arboreum m " n>e - deeply hssured - endosperm entire. Distribution and habitat. Tin- >pecie> grows in Sri 1836, nom. superfl. Meg. Gaertnera koenigii V ! !'' Gen F'l. 2- 426 Jecen.lie.. and «,.h l.mls January through Apnl and •E: Sri Lanka, s. loc.. in herb. Van gU> ' "" ' """ ember - Royen 108, /. G Koenig s.n. (holotype. L!: Discuss isotype, L!). Figure 15G-N. call,, van (BM f,H MO) kudne ttiddu, h L,t « fk.ln uJ.t, el^l.iw fclmo JIm lli.is 1, , lu , ,h„ r » -, Im.'ioi- .1 li i lib jf „ t , , „/ [ |r „„ (R M \o llc . Il e 1983), but because of the lack of a „„„. Ill . -■/. .-, ..7,„ - IH H>1 f '" ^"" ' ril '- -I'l"" llth f "^'»> «'" • l'« - ! 1891. TYPE: Sin; (holotype, K!; isotypes, BM!, CGE!, K!). Representative MALAYSIA. Johor: smooth. Leaf blades (3-)4-10 X (0.75-)l-4 cm, N; tgle, Mafco m &er 3070 : '- ■ . I" 1 ^""" 11 """- " l,,f i- 1 """ 1 "A ul.- i SSoi-el ,m 19T 1 1 '!(.,*] ,',0 1 ilni,Jl-l (2 Trees, 1-4 m tall; branches ^terete, glabrous, 0.7- ■..■■.. ' ■ ... ' ■ ■ •■ ' . • , (pubescent near apex), stigmas ca. 1mm long. vied flowers: calyx -. This species has been c t! , is I ill u tin. fli June an.! m N t.s l„>, n'lh.,..L, !.'..! ar to long styled except corolla tube 8.5-12 mm, 5 mm diam., lobes 5-6.5 mm; anthers shorth ad habitat. This sf .ecies grows in Sri Lanka, where it is found in premonl :ane and montane forests at elevations of 750-1650 m Phenology. This species has be en collected with floweis Febman though Septembt :r and with fruits ;h November. Discussion. Gaertnera walked is generalh similar to G. ramosa of weste; d to G. rosea, also , ith hoc, ,..,!].. 1.1- , m,\U ,, •vi.ti. ..nuN L to 3-flowered and fasciculate inflort :scences. In areas where G. walked and G. temifolia g (e.g., Adam's Peak), the natural hyt id * •■ mine, occasionally occurs. Gaertnera Xgw ■dned differs from G. walked in having mixed opposite ; and ternate leaves !!J . P1)V. ^35 arpa Drake, Bull. Soc. 1 [1899]. TYPE: proba i iM'i H. ... />7f Dctalu 8 ala Forest. ,er the is annotated by Drake rted and the descriptio 259, 2: 168. 1840. TYPE: Mauritius, not , Hern) E. M. A. Petit, Bull, designated. Jard. Bot. 29: 382. 1959. 12. 1877. TYPE: aim ms, ju m ic i Gabon . Es Mann 1721 (holotype, K photo!). 45: Only a pho specimen at Kew has vm s.n. ^.oio.ype, r no. ioca.eu 7 . generic placer confirm because the tity of this name is unknown. seemed to hi 'nera liberiensis but !. 1910. TYPE: B< S Gaertnera thouarsii Baill., Bull. Mens. Soc. Linn. hi, Winkler 3321 (holotype, BO not seen). Paris ] md. TYPE: Mauritius. P. Duj iotype, P not located). 3eusekom. The descrij variously identified as located, and its 2(1): 68; Malpighiaceae (Steudel, 1840)]. ( ies C. E. C. Fisch., Bull. Misc. .en. PI. 2: 425. lnf orm . Ke- rhinocerotis kom, 1967)]. , ■•'•• : for species previou ?28: 411. 1928 [= Chassalia ex A. DC, which is by him, but he Milne-Redh. (Petit, 1959a)]. Gaertnera par. etz., Observ. Bot. 6: 24. 1791. Gaertnerc . pongatU, orth. var Sphenoclea :e)lanicu ; Gaertn., Campanula or Lobeliaceae (Steuc lei, 1840)]. 19, t. 18. 1795. B; isionym: Molina race Cav., Diss. 9: 435, t. 263. 1790 [= Hi} benghalensis (L.) Ku rz, Malpighiaceae (Ja, 1955)]. tnera richardii Drake. Bull. Soc. Bot. Franc 355. 1898 [1899] [pr obably = Psychotria s 216 8.' l . )889)J (r ,' ,, 11 1,1- 7K ['ill I I ! Mil 6: 164. 1915 [= Psychotria sp. (a 1967)]. : ifrjqu. Jahrb. S.st. 48: 430. 1912 [ = ]- Chassalia, a,„t„lu DC. (Wi 9: 35. 1845 [= Chassalia 1989, as "G. coeru/, |- /■/„„„,/,„ /...,..,„„ h<. ,\,., Occ. Franc. 1: 444. 1920, nom. nud. [= Pre, quadrifolia Schumach. & Thoi (Petit, 1959a)]. ) Literature Cited !sl t IV u- l..i PI J -IJ"> J,V,I ,-, jc Wfl . V)11 L 1W8 \ , ,™ of th ». nu ' , & P. T. Li, Blum :6 [= /xora (Poir.) A. Chev. subsp. lanceolata (Verdcourt, 1983)]. • Mauritius a 181-207. ichusetts. Doyle, J. J. & J. pid isolation procedure Allun'L ° ' Drake T 18W f • * m,!™,. ,,.- ', ,ml, « < - , 200! III. ^ trills. ,t . p s .vi.nte.id Vision Nepokioefi M 1 P , . .< I A K ^t .,. J<»> l».,».ifM Ho. „i,i, - t .llnt.i I'.J ,\, 2 ,( i-H i u ii. 1 C> j_> i ,.' i ' .tit L D M 2524 eaux, G. 22 (5 r V> ,tf CJ 1 ,.l i ill H f „!,, | 8_1 (40b) l,u(Hl 186(50 <, '(.. Be , .1. . I ll 150 it 1 ,, ) F r < V i I t,j i I 1 D > 11 I I H i I I . > - ,'_''_' ii". >53<> il it, ,"., 4 1 4-t ■ » . 1> sh .,j.l I irf i .1, -\ t f n !',( c "i Jl trt l. 7.jS(l> J ), n, t, f 1 . . 2' HI 'I W.I ,...; .'.'.'. -if... !',•>.. (,1Hm,; ,.-,<> (1-1 Dm Petit- Dioua^. P. 100 (10). (68), 11621 (68), 12839 (44), 14413 (49b), 20811 (68); (50); Enti, A. A. GC 35586 (50). U 4l>_-!4 i=. . " i\ .731 111 1'j\m,--|-4, l c " -r - r i (5, -P .!.,.,. -' ->' I I.Euu.1 CM. 2827 (50); BoMton, L. S. MALI 1080 (20), 1085 (52); Boyer, G. 59 (8), 6327 (50), 6448 (40bi 66l8il l!2l!(i,i £ -11 \ (68,. Bi.mu B 895 (5>'i. 1 Mir. ,65 llj,!!, ilium j -HP (101., I40bl 5458 (40a, 5513 (50) 5974 . ,0 C > 5 >( , i ,1 lur, ,-' 111 , . I f lib - N '1810(34) Fliun V i'MiJ'tO >->J 7 i VUi ]'i , V ' M|i'l'Vn,ni Si \l ' 1- U ij I .- 'i i'.,,. ' 1 !( 1 IV; l< ( .) Bui . «. , H V i 1 «- - >. r >U ,27 (49a, 117(44) 401a (44) TVIk m R 56500 . '• I Si , •, L. 2514 (43), 032 (13), 1107 J.J 3 (50); Gillet, J. 2809 r. 1 '. > i r .(. l.i ,r ,2 ! r ,i> l])',' Uu. 17 (58). Lemidid. ] 84 i SO 1 ' (",('. z-l\ J. , <7 ■ T4 ; Hill ! I i" l-Ki^i 21 i- -4i _" ( i" '(I i ._' I ..I. I i Lull > l> H 2 . 1 1 , > i40 ( t'l, I I'.'' i 1 iJ '. i H, Hall. \ 2 )1 MJ.'.i 443, (06i. 46] 7 1487 \ (13), 1487B (13); Lisowski, S. 16116 (8), 18904 (8 -f i '.) Hmldnd (, D 3460(54) Hendei-,011 909(37) 1WI i i'i I1U nd U),lM' , U- ! «1 M.R. 11476(18), 17769(57), SING 23390 (57); Hepper,F. (40bi 188] (401, J!!"-, iM 2006 |5>!> 22 r 2 ( .7, 2 .i ><> ,R'i "''-9 (57) 11211 (37), 11586 (37 20600(54)21997 132 2201 , , >', _>_■>_• t , »2. 220 >,. _'! ;,j 2229 (4) 2585 (42) 2589 1 55), 2594 (43), 2637 (4), L. 28 (50). Ilias, B. P. S 39183 (18); Imbert, T. 94 (28). , 1] , _'■ >>2 1 1 .,•>>. 12 2 * t H 2812 |4) 2814 (54) ,1- n ]<.,,«„ 151 .,!>. l+ ". i J,'l'U*'»i 284<>,'«> i 2.>-7 .4 '1 , _\> 'Xl>i,> l-l,.3 ] „,, ,d „ t '(4i, - , I 'if , 4 j i, J ,.| >i »'(■>, 2MH >.i ''If |.j 2910(541 2911 ( 4j) 004 (7), 3054 (7). i2» -i("'«il i UH»i4 > ?iM-J i -t, .Hhhi \J hi 417, t,0 J>W0B(68> 4102 (68), 4 ,, i\ 1 .0| *(._'>' ill, ' (' I _'i 74 (18), 116 • m. fMi it: ,.• i. J i)288 (67). 10374 (17K 10461 170) *>»i ..'(il ftiU, ft 72iH»l.i =.i°. H-'M.i. ," iK 71' ,,().: \«orho«e.A.823(13). (59). 1465 (59. 1483 (79 1)11 117 ]r,<>) 0-.7 (.(',. W >.d , 15584 (57), FRI 15585 (57) ^i~\ (41); Zollinger, H. 3051* (34). Name coffeoides DC. psychotrioides DC. G. bifida Bojer ■ E. M. A. Petit G. longiflora C. F. Gaertn. G. longifolia Bojer p M««W«r/n« BJ.,1 *nf,/„ f ff V sifolia (Roxb.) DC. (Mai (Hiern) E. M. A. Petit G. thouanii Baill. G. vaginalis (DC.) Merr. .use & Gilg •v , o \\ an alis Schweinf. ex Hiern P. stictophylla Hiern S ,<>,,,'• i}S v i hi - W, 7,, f „ I -m-hIi, 19aL i & \\ J lei, 1999: Delpli & Sakai & Weller, 19' an in then sexual expiession pie's Republic of China. CEDEX 05, France. Cryptic Dioecy in Nyssa yunnanensis 1:1 i ..is iiu.,1 x \»<;.b M>v I iN>-uiai. lt , tui.i I »li > , wi in I Imu ins < '.jUs , ,1-ii.nrt di-tnbution I has a rich fossil ([v.l . Dl.il T',1 l,'„l,»llif1J\. s,s( ,„ „1 \„w |... .hoe j u, m.lnu l.t-,»„l],,nt..|ln I. ih n^, ts ln ,l orly known. ;. Q. Yin ex H. N. Qin & Wangerin as closely related taxa with the two sf ed, Wen and Stuessy (1993) treated it as belong the A jauuuca complex and used A jauuu represent this presumed lineage in their phyloge mid p.-..|« J s f -d ii.t- transfer iron, Rank HI to Raul: I JuglansL.,Perseaf in from morpho (101 06 E 22 25 N* xi. - L'i i x \ ■ ij. ... iO,M-n| u( > .ulh.ti-hllr,, «<...•„- the o.xlM.lfflu IR. ,,...1 n., -1U - >• K-xsx, ■-,.--■ Cryptic Dioecy in Nyssa yunnanensis k- ... f ....i^-i.iit]. pi.-oil,d i- -i.-i,,a-..f hw ...ipj, ut I I.A.. i si, ,lh ,tamtns from the than ont mdic ite t f 1 1 k ' i .s ' i i , ! th , I, ,1, 1 ) ,| 1( . t ] lt t|„ | i anthers of N. 5 active to pollinators. Discussion and our res, it set is higher in ■■■■ isis. Thus, although the leless be important for the functional,, dioecious. At least 78 species in 20 1 ch as bees and flies, (1991) to exhibit functional or cnptic dioec,, and r < p f^ored in forest , m , uii„ |. cj j.i v In. n i«]luuu .1. 1 li , .... . n-m> ,! un n, Ik m 1.1 .1 - 1m rs I his lesult may be .■■■■• '• ■ . opmed f.nhei lliau functionalh female fkraei- but now. it, .!<]),* -wo, s,, ^ tV.],-. v\ *Lilr|,i - j ( ''» _ i .\ I^VM.l. I'l", Sx.nl lu.'j. - h „ (e.g., Melamp; Ipler & Bawa, 1978; MATING SYSTEM Sassaman, 19' va's (1978) study of ' r< -t tree, indicated that a female-biased sex . ,lh f.inl i' .... nip,!!. I 11,,, u ,. h.-l, H. ..1 t! . o I. . n n ,1, ,l„,i„ n, JO ,] I »k vv H. U. « , | ,1^ t „' ] i( ' | , ,U MI ," n ", 11,^200 ludmdi" Is dNi.hln \, •(,+„.,. ik'^nhLi^.n'V.t'Vl""^ n ^.u^lte i^n'.Tl','! botanical gai ;sent study shows, "•»/" 1 ^ H.. >,n|. i .iw 1 tlr I >,s ,i biodiversity (Turner et ly restricted i mpactf populations of /Vys.« rig chan ge tlfl.il.ln i .Ii Jl< -,'] K t. « t U» i . , 1 „ | tll]11) ,,.„ -, |„ Ulnnlf around the Puwen Ti on and the microclimate comersion of large areas of forest to culthate i ,„ ^.mI.I. ,. , , „,„;,,, h.,n. L.m.h -. U ^ ^J ?* ^ * >™ '^ ^ ...*. I, 1:2 51 lem, N. C. Garwood & Oxi Cryptic Dioecy in Nyssa yunnanensis ,, M. A., P. P. Lowry II & D. G. Lloy Polyscias pancheri (Aral >hea, M. M., P. M. Dix< Bot. 80: 26-30. Song, C. S., R. Z. Xu & Q. H. Zhan Spe les Voucher 1 + + nectar ; (Mei dae t - nectar SYSTEMATICS OF THE SOUTH Es od f. Stuessy, 3 and AMERICAN HYPOCHAERIS Karin Tremeuberger* SESSILIFLORA COMPLEX (ASTERACEAE, CICHORIEAE) 1 -it )M« f 1 rtliLliltt Ji i T l il I i i doi: 10.341 .7/2006131 i, «.1J lt\ » rlu i'iIuiljIi.j'i If]. 1 \ ■ In: o. m I n .i.'li .nun K .« 10 L ) -Ijni.slMiS.ulh \ui ura I L timaim l] <>".H, i.-.lu, ,-d Moiphological characters and states were obtained SI, UC, US, W, and WU, plus oui Ht,( .It. din I i L.]', l! ^ I .rris :nd Harris W), For study . ... iza ■ n ill..- i.i.,l IT- (u that the South tic and that tr ;. Maire from M 2(J>*'. Fi met-!,. >,... ^ jl , 2 i\FIP) stuhes h !, 1 i 1 t- ! ,lm fi . -,111 blages within the South Amencan ta\a i t ' I , ,!,! \ rh.s, t^M u n l\ 1 ,1 included plus a morphological cladistic analysis. acaulis, H. echeganni Hieron., H. erem Hook. f. ex GriseL . H. nm,kl ■ margin is entire or dentate. The epidermal cells are lular epiden the apex, being al ,ih >L»h.or e ,/,.,,''' • , i-lrtll... -ul // . ' -," < <-l H «. " ii « ji.Liiis ti \m Jl th, ,dleso-nt sptnes of yellow in ff. acaufc, / r hohenack- Hypochaeris ■■■■:■■ mucida. because the a\ailable specimens are too r . ig hted equally and faded oi ;■ and we have not collected the taxon. support va lue alculated with 1000 epidermal cells that [gated « ith RMIUS The anahses resulted in one most parsimonious tree of 40 steps, with fit = 110 and fit* = 57%; ,ih i- tli.> au.L,i- l.i.aia. tt 1<>, Ml .[..., o'« ~1i -^ ,u«l^l,i|. :■■..■■■■■..,,■ r ' state 4), whip trichoi,) - mth .1- a iK„J a c ,u ,li u. -<> .fh i. 1 » P. 1 n.,1 , - ■:..■■■ 1 ■ kb< did I Hila-w -..hillan - .i ,1 ,i k ie (< i> u .< t i lU^htu » h, n.h.tf, ,al s rloes not "at all completed. pu/urfris and h are also found at • •'■■..' , ■: J i .M I Ti i !, I ml n till . ■, (espernlh lm tempeiatmes an.l . Hid) //•;.,>, :ic',i- tomentose with ses 8). They are the Andes int, rrain, 1984; Taylor, ■ H. argentina 10 64 1 H. chillensis i I H. e/ate I H. caespitosa 9 m m H. hookeri 1 56 i H. sessiliflora - H. taraxacoides H. hohenackeri I H. mucida *—•— I H. eriolaena - H. meyeniana - H. echegarayi - H. eremophila - H. acaulis . Jl. 2: 810. 1753. TYPE: * :■ .*'■ , \ • K- "V , 1 ' .... ■^-i 2 4 , j, ,, K „ /'u * * c •■' + " i W " ~ V ^ * \ V" • -.- - ■*-- \ * B 1\. ; . ,| ^ ^ 1 ' % - ■. .*«•• « ^ ' V 1| • D ° 2 °° . "° 6 °° 8 ° ,]UK ^>|th Ul«.|, .1 1^ 1 . U> 1. ,rf ,1-1 ' . L.. I .11. and 5100 m. Volume 96, Number 4 Urtubey et al. 2009 The Hypochaeris sessiliflora Complex Linn. Soc. London 1(, LTV k,-. > I />> -p/> ,< p]»nfb> . i p. - 2 ml 1 0.5-1 mm: filaments 5- I!). Figure 3. 2007, 2008). Hypochaeris sonchoides Kunth, Nov. Gen. Sp. [HBK], quarto Habitat am wchaeris sessiliflora M. Hypochaeris sessiliflora flowers A. von photos LP!, MO!). et al. 382); Peru, Macbride & Uo^oupe. piesumajj estio.e . ,, TO t,„ ., ^ Herbs to 13 cm tall. Leaves linear-lanceolate or wi dentate to slighth d. ..I ,V, l.cl, ,„*!;.. - ,U„, , s ,., „t tl„ H , „// >/, , , , ,mi|.l- I her divided or entire. Another variable lanceolate to linen 1 '-_ , --in l,,i i I 1 ' ,A d t. iclmophoi is be ibatus, Schultz Biponti- . .... •.... ,.. :.,.,. j.mpL- fii. !,,.n.- „i,.I oul-j .,„J imclcll. r h,IW rh.t i,e .li.].l.,l ii 111. <;.^ i her without description 3 "^ ^ ^ P" » i^ec ^^ names of A. giitoui ?"> re 59405 ( NY )- prohahh had material ,.l //. ,t< tro r l,i!a. I- he ciied m/^, ,/->/:„ K:nv.„ . \i.x , (,,-n. PI IC'i. lud. I .",<.!) a f , , „ , R, [ fl -l-'i I <' 1 ' ■ ] » I u.wJI s ' .» < r M\'/ I - « ii..- I < ' ' i -i 2 ..1 .:-■ . .:. ■■ -. is sessiliflora Complex . Habit. — B-E. Phj mw.l.M-ie MlMi.luw.l. 11-2?. V|0 Mll plnl] , j , . , . i.ih< t , lb pi u A V a\,Uf ,i. . . ,f " ;■ Jin. oliL\ .)!,< v lourdrl oljoi,, h,,.icH ■ M \ 11 ttli. :■....'.,.. -.-r: - • underneath, 15-21 mm; tube 7-9 mm; ligule 8- b 12 mm; \asculariza is 10-17 mm; Steudel havi led the name A. ■ oi m ;i • J, Mexia 4191) H^pochaeris. r . aware of the earlier tozr/ts i- leadih } j^,, , K1 (J ) (|jl f ,, ,,, ,, ^ n • i - n .. "1 .. otam- pie-uni- a latti homornm ..] // r u ,«„«./V,« Moench (Moench, ,|K , tll |, , t l i,,,)l ,,.,,„ | tl li> '..t-ul. - tataxacoides Pourr. ex Steud. of Chile, Reaion I, Rvm. . t P > W II ,V',i7< 4B, circle). & Walp.) Wal xephala (A. Gray ex In ^iil.UJai ' ',, hoi li,i j„ Aalo «m!.i Ku.il |Li ut/. 1 8|l t „ . • M'-ihfsvrL.'jisstenorephalavar.integrifolia >,,hila twauuoides Meyen & Walp., but iiom .aiieh to/ ,w L . ,/ . ... 1 jt ni^,^ e-itn< .. wall. From Smith & Bu,iJ-:i,, Cruz 254 (LPB). Pot. ie la ciudad, Schult, ■ LIT.. -,■ , ■ -. • ) Walp. by We In il t li n. V s » ?,';< 7 ,, , j i j „,,„„ , , / , , , \\ Li,, ! I u„ 1 1 1 - » I i Ihi. Ik ' ,7i « J L ) 76 00S 11 Ul •- Smith ' ul ■> ^( (Sch. Bip.) Domke, vXTatn rA to uSSff^jK^SfA 1 «X 3900 Bonplandia 4: 54. 1856. TYPE: Peru. "Tobina in ■ 1 , hi ni ,t i > 1 1 [{,< i l uiUn . I i I 1 I 1 i probably fl . ., ii . u.umdid de Molo-Molo £o « uV IJ'Bl Iju...... l'll..-1'll.. . siu.H.ion, s .l< I,, V,,!.'.,!,'., If ,.,- <"'< HIT /„,( ',, -tJ'ii »i, 1... ,a ,. I 11.- is sessiliflora Complex 1.5-2.5 mm. Cyp: th whip and son Itill. p.-tM.,-, „„,.,. (-11 Him tic. 1- . . 12 ( i.ioJIa )--ll -, ished earlier In Schultz Biponti (1855: 236) as a nomen nudum. Although wit! .akeri"} was publishe, Bipon' ; -- Uahltat and dls i from Peru (Puno) and Bolivia (La Paz). It .unfits the And -4700 m, associ- 2111a), which was I (1856: 54) as the original specimen to accompany his valid LLla "^°>- r! „,„, ,;>:: spetimrns BOLR I\ Co.'hal,«.i.*a : ,„8 (US, photo LP). 1 2V) i »j(, TM'E Paz: "Mittlere Anden: Siidlich des Titicaca-See 00 ft., 12 Feb. 1903, ^ bo) -<. \ .>i. - r. i '.) i.i-r p ™ ",.». _•_ .| J- i-iiv, , r P' h 5. Il»|«.,-lin-i-i ! > .-riol:u-n:i IN I. iir s u,.l i ii.ii- tint "• liu.i, ill il' Il i.y iji [>-, (.i,in ( ..nil . /■"_ ,L]-I « ln^i w. a: i 1 . n i ., :•■■'■'-' ■■-■■ , Vacas, Cardenas 203 (LPB). La Paz n ,i . . >0 ! i , M t . - -I > I I „ 1 I b 1 1 , w ,„ , i a - Herbs to 5 cm tall. Leaves lanceolate or oblong, 20- TS LP* M..H...VH b,',m /... ,' "'', 'Jlilt, l»^>V" , ) , -'(! 1 n m. plTll.u. ,"'' I- ' l' ,., i:. I. ///-. A /...-,/ IV. a -, \, : i, 3100-4950 m (Fig. 4C, triangles). Phenology. Hypochaeris ec Nac. Ci. 4(1): irgentina. Sa located; rsotype, LP!). Figure 9. ^TTZU to the latter. are faded. ages type 2 or 3, 0.8-1 mm; filaments 6-12 mm; Ul " ' ll V ' ' ^ ' 2 ' "' 'w'riou"« n' i >,<[', llh ) 'U una '"ll is In its peduncules with La Plata 13: 22. 194! : two specimens. h-ie, SGO 65208!, SGO photo I ! ^ "' ' ' ' "J ^J^ ^ ' ( ^ ( "' ' ~ u ^ ^ ^ '"' ige!). Figure 10. >' l 'Zi'l\"L,!ii 'll'I /,'.!'" 'm] '< ll f»««'Ai »?»"•»" J ^'" (LP) PERTF P.u.o « ! - .1, I,.], -7 Feb. 1! 94 - Gar. ,. .l».l.,r M SI! • : ' : . Leaves ob with divi ■ ip 1 la ■ X 10-25 n a. 13 . yellow, 10-14 mm; tub* ca. 2 mm; f a h tt, with bra nches ca. 10-15 mm. In = 8 K m tl„ s. tribution. X 150 km ,/.. t, *1, . It occurs between 1 Jh- -h^Llh 1,11^1 lll.lt! I'.t Volume 96, Number 4 Urtubey et al. 2009 The Hypochaeris sessiliflora Complex -l I U J .i. i I "Imm .if i'm.J. -> „ wt Bae.a 15)71 But Jahil. >• < „ J I ."(.-W H>U HI I \lio .1 \il, i, . .n, rim, mJl,,] //. <<■'< - s I',, .61-363 m A Eimlei K< ji«m T\ 1U 11 1 lo. ^ 1 i rol'n. t - , ■, A 1 ' / ' Papavero, \. l'»71. Essays on the History of Neotropical Washington, D.C. ' Paulo. Goloboff. P. A. 19 Gray, A. 1861. Chi '_' Sot " m ,i,J " ^d™ 03 01 01 12 123 H. mucida 1 - 1 , ill IP. 18080, 18083 (LP, WU) ^ ' ^ ' Pan: 6754 (91, 7723, 7'Cf, (61 '.' >77i (H<- „'» > ' -. "" J ( 325j5 (8) (mrif & S'ut _ _* i , ' . • , ^ i. . 78064 (LP) 25 (LP) ., lf-w, , , I ,' ,,",-,1 oiamZZos 7272 (US). . . H77IZ« ffiflora: ■3177 ■ :) : ■8 (1); 1198, 1235 (6), 153 - \.;,i-nr!l, IJL'n ('.".: i'Wmy;,. 1955 (1). nift (1); W |C . Jiohurdr, T-12 (2). 131 (6), 8. ANNALS OF THE MISSOURI BOTANICAL GARDEN VOLUME 96 2009 169-520 oJ www. mbgpr ess . org CONTENTS A Revision of the Malagasy Endemic Helmiopsis (Malvaceae s.l.) Wendy L. Applequist 52 1 A Revision of Neotropical Bonyunia (Loganiaceae: Antonieae) Jason R. Grant 541 iniylogcnctic Position and Taxonomic Classification o\ Aelhionema Irinemun, (lirassicaecac): A Morphologically Variable Subshrub from Southwestern Asia Ahmad Reza Khosravi, Fernand Jacquemoud, Sasan Mohsenzadeh, March Menke & Klaus Mummenhqff 564 A Systematic Revision of Gaertnera (Rubiaceae. Gaertnereae) Simon T. Mc te M. Taylor 575 Cryptic IW<\ hi \.sv, vwnannu,, (N\^i(,ac), a ( uticalh Fndangdcd Specie hoin Tropical Eastern Asia Bao-Ling Sun, Chang-Qin Zhang, Porter P. Lowry II & Jim Wen 672 Systematica of the South American Hyjux-lmms sessiliflora Complex (Asleraccae. Cichoricae) Estrella Urtubey, Tod F. Stuessy & Karin Tremetsbergei 685 Checklist for Authors 717 Author Index _____ 720 Subject Index 722 idl.)E.L. Cabral,(