Galphimia (malpiGhiaceae) in South america (malpiGhiaceae) in South america Christiane Anderson...

2005 C.ANDERSON:GALPHIMIAINSOUTHAMERICA �

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Contr.Univ.MichiganHerb.24:�–�2.2005.

Galphimia (malpiGhiaceae) in South america

ChristianeAndersonUniversityofMichiganHerbarium

3600VarsityDrive,AnnArbor,Michigan48�08-2287

AbstrAct.  ArevisionoftheSouthAmericanspeciesofGalphimia(Malpighiaceae)ispresented.Fourspecies,G. amambayensis,G. australis,G. brasiliensis,andG.platyphylla,arerecognized.Fulldescrip-tions,akey,andtwomapsareprovided.Eachtaxonisillustrated.Onenewspecies,G. amambayensis, isproposed.

Galphimia(Malpighiaceae)includeslargeherbsandshrubscharacterizedbyyel-lowpetalsandaschizocarpicfruitseparatingintothreecocci.ThecalyxlackstheoilglandsfoundinmostgeneraofNewWorldMalpighiaceae;however,insomespeciesofGalphimiaasmallgland,whichresemblestheleafglands,isborneatthebaseofthesinusofsomeoralladjacentsepals(Castroetal.200�).Mostoftheca.20speciesofthegenusoccurinMexico,adjacentTexas,andnorthernCentralAmerica,buttheremainderarefoundinSouthAmerica,southoftheAmazonBasin.TheMexicanspeciesmaybeassignedtooneofthreegroupsdefinedbydetailsoftheinflores-cenceandflower.Intwo,theflowersarelargeandborneonapedicelsubtendedbyapeduncle;inonegroupthepetalsaredeciduous,intheothertheyarepersistent.The third group includes two species most similar in overall aspect to the SouthAmericantaxa:thepetiolesareveryshort(mostlylessthan�cmlong),thepeduncleisrudimentaryorabsent,andtheflowersaresmall(thepetalsdeciduous).

ThreespeciesofGalphimiahavebeenproposedforSouthAmerica:G. australisChodat,G. brasiliensis(L.)Adr.Juss.,andG. platyphyllaChodat;however,mostbota-nistshavefollowedNiedenzu(�9�4,�928),whorecognizedonlyonepolymorphicspecies,G. brasiliensis, comprisingseveralvarietiesandforms.ArecentreviewoftheSouthAmericancollectionsofGalphimiashowedthemtobelongtofourspecies.Galphimia brasiliensisoccurs innortheasternBrazil (Paraíba,Pernambuco,Piauí,andBahia).ThevariableG. australisrangesfromcentralBraziltowesternBolivia,southern Paraguay, Uruguay, and adjacentArgentina, and the closely related G. platyphyllaisfoundineasternParaguayandadjacentBrazil.Thematerialfromeast-ernParaguayincludedonecollectionthatrepresentsaspeciesherenewlydescribed,G. amambayensis.TherelationshipsoftheSouthAmericantaxawiththeMexicanonescanbeaddressedonlyafterareviewoftheentiregenus.ThetaxonomyoftheSouthAmericanspeciesispresentedhereasanaidtocurrentfloristicprojects.

Akeytothefourspeciesispresentedbelow;however,theyarequicklyseparatedbygeographyandobviousmorphologicalcharacters.OnlyG. brasiliensisoccursinnortheasternBrazil,andhasstrigosevesture,composedofhairswithapersistenttuberculatebase,anduniformlyvelutinousovaries/fruits.Galphimia amambayensisoccurswithintherangeofthewidespreadG. australisandneartothatofG. platy-phylla; itdiffersby its tomentose sepalsand inflorescences,andpartlyvelutinousovaries/fruits; the sepals and ovaries/fruits of G. australis and G. platyphylla areglabrous.Asthespecificepithetimplies,G.platyphylladiffersfromG. australisinitsmuchbroaderleaves(length/widthratio�.2–2.3(–3)vs.(2–)2.5–7(–9).

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Key to the species of GAlphimiA in south AmericA

�. Sepalsabaxiallytomentoseorpatchilyso;ovaryandfruitvelutinousalongthesuturesandontheproximal2/3–3/4ofthesurface,thedistal�/3–�/4glabrous;Paraguay(Amambay).

G. amambayensis�. Sepalsglabrousorwithscatteredhairsnearthebaseand/orwithatuftofhairsattheapex;ovary

andfruitglabrousoruniformlyvelutinous.2. Ovaryandfruitvelutinous;sepalsusuallywithatuftofhairsattheapex;petioles3–�0mm

long,strigosewiththehairsmostlyborneontubercles,orinolderleavesonlythepersistenttuberclesremaining;pedunclespresent,0.4–2(–3)mmlong;Brazil(Bahia,Paraíba,Pernam-buco,andadjacentPiauí). G. brasiliensis

2. Ovaryandfruitglabrous;sepalsentirelyglabrous;petioles�.5–6mmlong,glabrous;pedunclesabsentorrudimentary,sometimesto�.5(–2.5)mmlong.3. Laminaslineartolinear-lanceolatetolanceolatetonarrowlyelliptical,(0.5–)�–2.5(–3.3)

cmwide,length/widthratio(2–)2.5–7(–9);pedicelstomentuloseorwithscatteredhairsorsometimesglabrous;sepals2.5–3.5(–4)mmlong,�–�.8mmwide;petallimb3–5mmlong;cocci3–3.8mmlong;widespreadthroughoutsouthernBrazil,westernUruguay,Paraguay,andadjacentBoliviaandArgentina. G. australis

3. Laminaselliptical,broadlylanceolate,ovatetobroadlyovate,occasionallysuborbicular,(�.5–) 2–4 cm wide, length/width ratio �.2–2.3 (–3); pedicels glabrous; sepals 4–5.7 mmlong,2–2.5mmwide;petallimb7–8.5mmlong;cocci4–5mmlong;Paraguay(Amambay,Caaguazú,Canendiyú)andadjacentBrazil(MatoGrossodoSul). G. platyphylla

Galphimia amambayensisC.Anderson,sp.nov.—type:pArAGuAy.Amambay:30kmalNdelcruceBellavistayPedroJuanCaballero,enmatorral,��Feb�982,Fernández Casas & Molero 6190(holotype:NY!;isotype:MO!). Fig.�.

Suffrutex.Laminae4.5–6.5cmlongae,�.5–2.8cmlatae,ellipticae,sparsepubes-centesdemumglabrescentes,margineintegra,glandulis0.4–0.5mmdiametro,ses-silibus;petioli3.5–6mmlongi;stipulae4–4.5mmlongae.Inflorescentiaracemosa,tomentosa.Petaladecidua,subaequalia,ungueca.2mmlongo,limbo5–5.5mmlongo,4–4.5 mm lato, late triangulari.Antherae ca. �.3 mm longae. Ovarium velutinumproximaliter, suturis pubescentibus distaliter; styli subaequales, 5.5–6.5 mm longi.Coccica.4.5mmlongi.

Subshrubs to0.6m;stemstomentosewhenyoung,becomingglabrous inage.Vestureofcrispedorcurled,reddishbrownhairs0.3–0.7mmlong,exceptstraightandappressedonthelaminas.Laminasofthelargerleaves4.5–6.5cmlong,�.5–2.8cmwide,ellipticalornarrowlyso,apexapiculate,baseacute,adaxiallyandabaxiallysparselypubescentwhenyoung,mostlyglabrouswhenmaturebutsometimeswithsomescatteredhairsremaining,costaprominentabaxially,thesecondaryveinsbarelyprominulousornotevident;marginentire;petioles3.5–6mmlong,glabrous; leafglandsapairborneonthemarginnearthebaseofthelaminaorto6mmaboveit,eachgland0.4–0.5mmindiameter,circularandflushwiththemargin;stipules4–4.5mmlong,0.7–�mmwide,linear.Inflorescenceaterminalraceme,theaxesdenselytomentose,glabrescentinage;pedunclesabsentorrudimentary,pedicels4–6.5mmlong,denselytomentose;bracts2.5–3.5mmlong,0.7–0.8mmwide,linear,bracteoles�.2–�.5mmlong,ca.0.5mmwide,linear,bractsandbracteolesabaxiallytomentulose.Sepals3.5–5mmlong,�.3–�.5mmwide,narrowlyelliptical,abaxiallytomentuloseorpatchilyso,glandsabsent.Petalsdeciduous,subequal(?),yellow,becomingredinage,abaxiallysprinkledwithhairsontheclawandonthelimbonandadjacenttothemidrib,orsometimesglabrous;clawca.2mmlong,ca.0.5mmwide,limb5–5.5mmlong,4–4.5mmwide,broadlytriangular,baseacute.Stamenswithheteromorphicfilaments but subequal anthers; filament opposite anterior sepal 3.5–4 mm long,

2005 C.ANDERSON:GALPHIMIAINSOUTHAMERICA 3

FIG.�.Galphimia amambayensis.a.Leaf.b.Detailoflaminashowingmarginalgland.c.Portionofinflorescence.d.Openingbudborneonpedicel,withportionofinflorescenceaxis.e.Petal,abaxialview.f.Gynoecium.g.Coccus,abaxialview.Scalebar=a,4cm;b,8mm;c,2cm;d–g,4mm.(BasedonFernández Casas & Molero 6190.)

filamentsoppositeanterior-lateralpetals4–4.5mmlong,filamentsoppositeanterior-lateralsepals3.5–4mmlong,filamentsoppositeposterior-lateralpetals4.5–5mmlong,filamentsoppositeposterior-lateralsepals3.8–4.3mmlong,filamentoppositeposteriorpetal4.8–5.2mmlong;anthersca.�.3mmlong.Ovaryvelutinous,butinthedistal�/3onlyonthesuturesandadjacenttothemandglabrousonthesurface;stylessubequal(?),5.5–6.5mmlong.Coccica.4.5mmlong,ca.3mmindiameter,velutinousliketheovarybutglabrescentinage.

Galphimia amambayensisisknownonlyfromthetypecollectionfromeasternParaguay(Fig.5).Itisreadilyseparatedfromtheotherthreespeciesbyitsdensetomentumonthecalyxand inflorescenceaxes.The leavesaresparselypubescentonbothsurfaceswhenyoung.Theovaryisvelutinousinadistinctivepattern:thevestureispresentalongthesuturesbutonthesurfacesonlyintheproximal2/3–3/4.Inthetypecollectionmostofthepetalsareabaxiallysprinkledwithhairsbutafewareapparentlyglabrous,thoughintheseinstancesthehairsmayhavebeenbrokenoff.SuchpetalpubescenceoccurssporadicallyinsomespeciesofMalpighiaceaethatusuallyhaveglabrouspetals,andonlyadditionalcollectionsofG. amambayensiscanresolvewhetherpubescentpetalsareindeedcharacteristicofthisspecies.

Galphimia australisChodat,Arch.Sci.Phys.Nat., sér.3, 24:500.�890.Galphimia brasiliensisvar.australis(Chodat)Chodat,Bull.Soc.Bot.Genève,sér.2,9:96.�9�7.—type:pArAGuAy.Guairá:PrairiesdeCosme,entreVillaRicaetCaaguazú,Nov�874,Balansa 2393(lectotype,heredesignated:G!;isolec-totypes:K!P!).

Galphimia brasiliensisvar.pubescensAdr.Juss.inA.St.-Hil.,Fl.bras.merid.3:72. �833 [“�832”].—type: brAzil. Rio Grande do Sul: ad ripas UruguayjuxtavicumS.-FranciscodeBorja[SãoBorja],Feb,St. Hilaire C2, 2495bis(holotype:P!).

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Galphimia australisf.angustifoliaChodat,Mém.Soc.Phys.Genève3�(2:3):23.�892. Galphimia brasiliensis f. angustifolia (Chodat) Nied.,Arbeiten Bot.Inst.Königl.LyceumHosianumBraunsberg5:22.�9�4.—Type:pArAGuAy:Guairá:páturaged’Itangu,prèsdeVilla-Rica, �7Feb�876,Balansa 2394(holotype:G!). Fig.2.

Subshrubs or suffrutescent herbs to 1 m. Laminas of the larger leaves 2.5–7 (–8) cm long, (0.5–) 1–2.5 (–3.3) cm wide, length/width ratio 2.5–7 (–9), linear to linear-lanceolate to lanceolate to narrowly elliptical, apex apiculate, base acute; glabrous adaxially and abaxi-ally, costa prominent abaxially, the secondary veins barely prominulous or not evident; mar-gin entire; petioles (1.5–) 2–6 mm long, glabrous; leaf glands borne on the margin near the base of the lamina or to 0.8 cm above it, usually a pair, or sometimes with 1 or 2 additional glands, or sometimes only 1 gland or the glands absent, each gland 0.2–0.5 mm in diameter, circular, flush with the margin or prominent or sometimes with a stalk to 0.5 mm high; stip-ules (1.5–) 2–4.2 (–5) mm long, 0.5–1 mm wide, linear. Inflorescence a terminal raceme, the axes mostly tomentulose during anthesis, becoming glabrate in age, sometimes glabrous already at anthesis, sometimes the vesture retained in age; peduncles absent or rudimentary, sometimes to 1.5 (–2.5) mm long; pedicels 2–5 (–10) mm long, usually tomentulose or at least with some scattered hairs but sometimes glabrous; bracts 1–5 (–7) mm long, 0.4–1.2 mm wide, linear to narrowly triangular, bracteoles 0.5–1.5 (–2) mm long, 0.2–0.5 (–0.7) mm wide, linear, bracts and bracteoles abaxially glabrous. Sepals 2.5–3.5 (–4) mm long, 1–1.8 mm wide, oblong, glabrous, glands 0.2–0.5 mm in diameter, sessile, mostly 1 but some-times 2 glands borne in the sinus at the base of 2 sepals, or glands absent. Petals deciduous, subequal, the posterior petal sometimes a little larger, yellow, becoming red in age; claw 1.3–1.5 (–1.7) mm long, limb 3–5 mm long, 2.5–4 mm wide, triangular-ovate, base acute to truncate, apex obtuse. Stamens with heteromorphic filaments but subequal anthers; filament opposite anterior sepal 2.5–3 (–3.4) mm long, filaments opposite anterior-lateral petals 2–2.3 (–2.5) mm long, filaments opposite anterior-lateral sepals 2.3–2.5 (–3) mm long, filaments opposite posterior-lateral petals 1.8–2 (–2.5) mm long, filaments opposite posterior-lateral sepals 2–2.3 (–3) mm long, filament opposite posterior petal (1.8–) 2–2.2 (–2.5) mm long; anthers 1–1.2 mm long. Ovary glabrous; styles subequal but the anterior two always a little longer than the posterior one, anterior styles 3.8–4.6 (–5.2) mm long, posterior style 3.5–4.2 (–5) mm long. Cocci 3–3.8 mm long, 2–2.5mmindiameter,glabrous.

Phenology. Collected in flower and fruit throughout the year.Distribution (Fig. 3). Southern Brazil, adjacent Bolivia, Paraguay, northeastern Argen-

tina, western Uruguay; open woodlands and grasslands, cerrado, caatinga, campo limpo, campo sujo, matorral, and secondary sites; 90–900 m.

representAtive  specimens. argentina. corrientes: Depto. Santo Tomé, Playadito �7 km W ofApóstoles,Anderson 12362(CAS,MBM,MICH,NY);Depto.SantoTomé,Ayo.Chimiray,Krapovickas & Cristóbal 25195 (CTES);Depto.Mercedes,Mercedes,aItáCorá,Ayo.Pay-Ubre,Quarín & González 2036(CTES,ENCB);Depto.BeróndeAstrada,46kmWdeItáIbaté,Valencia,Schinini 14017(CTES,F,MICH).—entre ríos:Depto.Concordia,ParqueRivadavia,Burkart & Troncoso 27703(NY);Depto.Federación,SantaAna,barrancadelRíoUruguay,Burkart et al. 29359(CTES,GH);ConcepcióndelUru-guay,Lorentz 563(BM,F,G,GH,K).—misiones:9kmalNWdeConcepcióndelaSierra,Krapovickas et al. 15148(C,CTES,MBM,MO,P,WIS);Depto.Caniguás,PuertoRico,Montes 3995(CTES,MICH);Depto.Apóstoles,Apóstoles,Xifreda & Maldonado 381(MBM). Bolivia.sAntA cruz:Prov.Velasco,SanIgnacio,�00kmNcomunidadSanMiguelito,Bruderreck 24 (CTES,NY);Prov.ÑuflodeChávez,EmbocadadelCarmen,ca.40kmSSWdeConcepción,�6°38'S,62°26'W,Krapovickas & Schinini 32151(CTES,F,MBM,MICH);Prov.Sandoval,SanMatías,�6°2�'S,58°26'W,Krapovickas & Schinini 36157(CTES,MICH);Prov.AndrésIbáñez,JardínBotánicodeSantaCruz,�2kmEofcenterofSantaCruzonroadtoCotoca,Nee 44434(NY);Prov.Sara,lomeríadeBuenaVista,Steinbach 7370(A,BM,F,G,GOET,

2005 C.ANDERSON:GALPHIMIAINSOUTHAMERICA 5

FIG.2.Galphimia australis.a.Habit.b.Nodewithstipulesandbasalportionsofleaves.c.Budborneonpedicel,withportionofinflorescenceaxis;notecalyxglands.d.Floweranddetailshowingpetalapex.e.Distalportionsofstamens,abaxial(left)andadaxial(right)views.f.Gynoecium.g.Styleapex.h.Fruitsubtendedbycalyxandfilaments.i.Coccus,abaxialview.j.Coccus,adaxialview.k.Seed.l.Embryo.Scalebar=a,4cm;b,8mm;c,f,4mm;d,5mm(apex4mm);e,2.3mm;g,0.4mm;h–j,5mm;k,l,2.8mm.(Basedon:a–g,Schinini 14017;h–l,Krapovickas & Schinini 32151.)

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FIG.3.DistributionofGalphimia australis.

MO,S). Brazil. Goiás:Mpio.CaldasNovas,�2kmdueWofthecityofCaldasNovas,Heringer & Eiten 14190(HB,K,MO,SP,UB);BR-�58,5kmSEdeAragarças,Krapovickas & Cristóbal 42948(K,MBM,MICH).—mAto Grosso:Oasis,47kmEdeCáceres,BR-070,Km680,SerradasAraras,Krapovickas et al. 40260(MICH);Mpio.Cuiabá,Rod.BR-070,ContornoSul,Hatschbach 65660(MBM).—mAto Grosso do  sul: Mpio. Campo Grande, RioAnhandui, Hatschbach 21813 (MBM); Mpio. Rio Brilhante, Rod.BR-267,entroncamento,Hatschbach 25071(MBM);BelaVista,�0–�5kmO,Hatschbach 49160(BR,G,MBM,US);Mpio.PortoMurtinho,Rod.Bonito–CampodosIndios,FazendaAguaDoce,Hatschbach et al. 74009(MICH);Mpio.RioVerde,baseescarpadelaSerraAlegre,ca.20kmNdeRioVerde,Krapovickas 29878(CTES).—minAs GerAis:ca.5kmSEofParacatú,Irwin et al. 26193(F,MICH,MO,NY);estradaSantoHipólitoaDiamantina,Km69,Shepherd et al. 3837(MBM,NY).—pArAná:Ibaréopp.,Morun-gavapraedium,Dusén 17413(GH,MO,S);Km�27,Mpio.LaranjeirasdoSul,Hatschbach 23123(MBM,MICH).—rio GrAnde do sul:BR-�0�,Km6,CampoBonito,ca.8kmSWdeTorres,Krapovickas & Cristóbal 38484(CTES,MICH);I.A.S.,Pelotas,Costa Sacco 407(F,HB,NY).—sAntA cAtArinA:Rod.BR-470,Km354,Mpio.CamposNovos,Poliquesi 495(CEPEC,CTES,MBM,UB).—são pAulo: Dist.RubiãoJunior,�–2kmSWdaFaculdadedeCiênciasMédicaseBiológicasdeBotucatu,Gottsberger & Campos 14-301072(F,MBM,MICH,UB);Jales,PastosdeRetiro,Hoehne SPF13903(MBM,RB). para-guay.Alto pArAná:Ea.SantaElena,PiraPyta,Schinini & Caballero M. 27152(G,MICH).—AmAmbAy:32kmSEdeBellaVista,Ayo.Negla,Schinini 21495(F,G,MICH).—cAAGuAzú:ColoniaPindo,camino

2005 C.ANDERSON:GALPHIMIAINSOUTHAMERICA 7

entreItaquyryyCuruguati,24°35'N,55°20'W,Schinini & Caballero M. 30243(MICH).—cAnendiyú:iteradYerbalesmontiumSierradeMaracayú,inregionefluminisCorrientes,Hassler 4524(BM,F,G,GH,K,NY,W).—concepción:propeConcepción,Hassler 7572(BM,G,K,NY).—cordillerA:Piribebuy,ColoniaPedroPabloCaballero,Rojas 12731 (S,W).—GuAirá: Iturbe,Montes 12595 (CTES,MICH).—itApúA:CapitánMiranda,4.2kmNdelHotelTiroldetrásdelBarrioCONAVI,Krapovickas & Cristóbal 44463(CTES,G,MICH).—misiones:�6–�7kmSdeSantaRosa,caminoaGeneralDelgado,Arbo et al. 6141(G,MICH).—Ñeembucú:SanJuanBautista,Purvey 390(CTES,MO).—pArAGuArí:NationalParkYbycu'i,NWcornerofparkalongArroyoMina,26°0�'S,56°46'W,Zardini & Tillería 28962(MO).—presidente hAyes:Primavera,Woolston 192(NY,S). uruguay. pAysAndú:Chapicuy,orillasdelRíoUruguay,Sta.Sofia,Rosengurtt B-3278 1/2(GH,MO,SP).

Galphimia australis is a widespread and variable species found from central Brazil to Bolivia and eastern Argentina. It is distinguished by glabrous leaves borne on very short petioles (to 6 mm long) and small flowers with a glabrous calyx and ovary. Although the youngest branchlets and inflorescence axes are pubescent, the vesture is soon shed and the more mature parts are usually glabrate to glabrous. In specimens from Rio Grande do Sul (Brazil) the vesture is retained in the racemes and leafy branches, and only the oldest vegetative branches are essentially glabrous. This variant was named by Jussieu (1833) and accepted by Niedenzu (1914, 1928) as G. brasiliensis var. pubescens. Such exceptional populations with persistent vesture occur occasionally in other parts of the range as well (e.g., Argentina, Lorentz 563, Entre Ríos; Xifreda & Maldonado 381, Misiones) and are not accorded taxonomic recognition here.

Throughout most of the range G. australis bears narrow laminas, ranging from linear to linear-lanceolate to lanceolate to narrowly elliptical, that taper toward the apex and are 3–6 (–9) times as long as wide. Occasionally, some laminas on a specimen are only 2.5 times as long as wide. Chodat (1892) assigned specimens with very narrow leaves to his forma angustifolia, based on Balansa 2394 from Paraguay. Schinini 14017 and Schinini et al. 23447, both from Argentina, are two similarly narrow-leaved collections. In some populations some of the leaves or sometimes even all are wider than usual, and approach the leaf shape found in G. amambayensis and G. platyphylla (e.g., Hassler 4524, Hatschbach 25071, 49160, Krapovickas 29878, Krapovickas & Cristóbal 25195). These variants match G. australis in all other aspects.

Galphimia brasiliensis(L.)Adr.JussieuinA.St.-Hilaire,Fl.bras.merid.3:7�.�833[“�832”].Thryallis brasiliensisL.,Sp.pl.,ed.2,2:554.�762.—type:“Frutice-scensherbapisonis.Margr.bras.79.f.3”[theillustration;p.79,inPiso,G.:De medicina brasiliensi;Marcgrave,G.:Historiae rerum naturalium brasiliae,�648.]

Galphimia brasiliensisvar.pubescensf.ovataNied.,ArbeitenBot.Inst.Königl.LyceumHosianumBraunsberg5:22.�9�4.—type:brAzil.Bahia:withoutlocality,�830,Salzmann s.n.(lectotype,heredesignated:G!; isolectotypes:MO!P!).

Fig.4.

Subshrubsandshrubsto�.5m,oftensprawlingandscrambling;stemspubescentwhenyoung,becomingglabrousbutoftenroughenedbypersistenttubercles.Ves-tureofallvegetativepartsofstraighttowavy,lightbrowntowhitehairs,0.3–0.8mmlong,eachhairborneonapersistenttubercleupto0.25mmhighortheepidermisonlyslightlyraisedatpointofattachment.Laminasofthelargerleaves2–5cmlong,�–3.8cmwide,ovateorelliptical,apexapiculateoracute,baseacute to truncate,glabrous but often with a few scattered hairs on the abaxial costa and along themarginnear thebase, costaprominentabaxially, secondaryveinsprominulousor

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FIG.4.Galphimia brasiliensis.a.Branchwithinflorescence.b.Detailoflaminabase,showingmarginalgland.c.Leaf.d.Petal.e.Stamenopposingposterior-lateralsepal(left)andstamenopposingposteriorpetal(right);abaxialview.f.Gynoecium.g.Fruit,subtendedbycalyxandfilaments,borneonpedicelandpeduncle,withportionofinflorescenceaxis.h.Coccus,lateralview.i.Coccus,adaxialview.Scalebar=a,c,2cm;b,�mm;d,h,i,2.7mm;e,f,2mm;g,4mm.(Basedon:a,d–g,Anderson 11737;b,c,Harley 16312;h,i,Blanchet 3904.)

barelysoabaxially;marginentireorcommonlywithafewpersistenttuberclesnearthebase;petioles0.3–�cmlong,tuberculate-strigoseoronlythetuberclesremain-ing;leafglandsborneonthemargin,to6mmabovethebaseofthelamina,usuallyapair,orsometimeswithonly�glandortheglandsabsent,eachgland0.�–0.3mmindiameter,disklikeandprominentorsometimesdrawnoutintoatoothto0.3(–0.5)mmlong;stipules�.5–2.7mmlong,0.3–0.6wide,linearorverynarrowlytriangular.Inflorescenceaterminalraceme,theaxestuberculateandstrigose;peduncles0.4–2(–3)mm long,glabrate,pedicels2.5–5.5mm long, strigose;bracts�.2–2mm long,0.4–0.5mmwide,linear,bracteoles0.5–0.8mmlong,0.3–0.4mmwide,linear.Sepalsca.2.5mmlong,�–�.�mmwide,narrowlyelliptical,glabrousorwithscatteredhairsatthebase,usuallywithatuftofhairsattheapex,glandsabsent.Petalsdeciduous,subequalbuttheposteriorpetalsometimeswithawiderclawthanthelateralones,yellow,oftenmarkedwithred,becomingpink/redinage;claw0.5–�mmlong,0.4–0.5mmwide,limbca.4mmlong,ca.3mmwide,triangular-ovate,baseacute.Stamenswithheteromorphicfilamentsbutsubequalanthers;filamentsofstamensopposinglateralsepals3.2–3.3mmlong,ofstamenopposinganteriorsepalca.3.5mmlong,filamentsofstamensopposingpetalssubequal,ca.2mmlong;anthers0.7–0.8mmlong,cylindrical.Ovaryvelutinous;stylessubequal,3.3–4.3mmlong.Coccica.3.6mmlong,ca.2mmindiameter,velutinous.

Phenology.Collectedinflowerandfruitthroughouttheyear.Distribution(Fig.5).EasternBrazil(Bahia,Paraíba,Pernambuco,andadjacent

Piauí); in caatinga, cerrado,atedgesofwoodsandgallery forests, in thorn scrub;280–��50m.

2005 C.ANDERSON:GALPHIMIAINSOUTHAMERICA 9

FIG.5.DistributionofGalphimia amambayensis, G. brasiliensis, andG. platyphylla.

AdditionAl specimens exAmined. Brazil.bAhiA:�0kmWofSerrinhaonrdtoConceiçãodoCoité,��°38'S,39°W,Anderson 11737 (CAS,MBM,MICH,NY);Valente,6kmNofValenteonrdtoSantaluz,Anderson 13689 (MICH);caminodeFiladélfiaaPindobaçu,a�4kmdelaBR-407,�0°48'S,40°�4'W,Arbo et al. 7334 (CEPEC);BR-324,�2kmNWdeJacobina,caminoaUmburanas,��°06'S,40°36'W,Arbo et al. 7359 (CEPEC);Iaçu,Fda.Lapa,�2°42'S,39°56'W,Arouck-Ferreira 256 (MBM);PastoBarreiro,Faz.SerradaMonta,Itaberaba,Bastos 162 (CEPEC,MICH);withoutlocality,Blanchet 1046 (G), 2184 (BM,G),3904 (BM,F,G);SerradaJacobina,Blanchet 2674 (BM,BR,C,G,MO,NY,P,W);ConceiçãodeFeira,margemesquerdadoRioParaguaçú,�2°32'35"S,39°03'06"W,Carvalho et al. 544 (CEPEC,MICH);Bar-ragemdeBananeiras,cachoeira,valedosriosParaguacúeJacuipé,�2°32'S,39°05'W,Cavalo 201 (CEPEC,MICH);entreJeremoaboePauloAfonso,Gonçalves 41 (CEPEC);LagôadeEugenia,southernendnearCamaleão,Harley 16227 (CEPEC);64kmNofSenhordoBonfimonBA-�30toJuazeiro,09°55'S,40°�5'W,Harley 16312 (K,MICH,MO,NY,RB);SerradaJacobina,8kmNofSenhordoBonfimonBA-�30toJuazeiro,�0°23'S,40°�5'W,Harley 16500 (K,MICH,MO,NY);Piatã,Harley 24152 (K);Itaberaba,Fda.Morros,Hatschbach 1986 (MO);Milagres,arredores,Hatschbach 42452 (MBM,MICH);Itaberaba,Fda.Morros,Hatschbach 48206 (BR,CEPEC,MBM,MICH);MorrodaGarrafa,�2°45'�8"S,39°5�'W,Melo et al. 2043 (CEPEC);PastoBuffel,Faz.SerradaMonta,Itaberaba,Oliveira 544 (MO);Mairí,Oliveira 647 (MO);Faz.LagôadoCanto,Ipira,Oliveira 682 (MO);Faz.VárzeaIpirá,Oliveira 713 (CEPEC,MO);SantaTerezinha,0.6kmNEdeSta.Terezinha,naestradaentreSta.TerezinhaeMonteCruzeiro,�2°48'56"S,39°32'05"W,Queiroz et al. 1540 (CEPEC,F,MBM);SantaInês,Km20daRod.Ubaíra/Sta.Inês,Santos

�0 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME24

3070 (CEPEC,MICH);beiRemonso,Ule 7180 (G);SantaLuz,30kmWofQueimadas,�5kmNWofSantaLuz,��°09'S,39°28'W,Webster 25670 (CAS,MICH).—pArAíbA:withoutlocality,Coêlho de Moraes 2116 (A,NY,S).—pernAmbuco:Sertânia,FazendaCoxi,Alencar 3 (MICH);Area-projetoSuape,CabodeSto.Agostinho,parteanterior(EstaçãoD),Andrade-Lima & Medeiros-Costa 48 (F);Cabo,Area-pro-jetoSuape,CabodeSantoAgostinho,Andrade-Lima & Medeiros-Costa 48 (MBM);nearPernambuco,Gardner 944 (BM,GH,NY,S,W);20kmdePetrolinaemdireçãoaAfrânio,Heringer et al. 185 (UB);20kmdePetrolinaaAfrânio,Heringer et al. 975 (RB);withoutlocality,Houllet s.n. (BR);Olinda,Pickel 676 (SP);without locality,Ridley et al. s.n. (BM);Alagoinha,SWvonCaruarú,Vogel 129 (MICH).—piAuí:LagôaComprida,Gardner 2077 (BM,GH).

Galphimia brasiliensisischaracterizedbythe“tuberculate-strigose”vesture,themostlyovateleaves,andauniformlyvelutinousovaryandfruit.Theandroeciumisalsodistinctiveinthatthefilamentsofstamensopposingthepetalsareonly2/3aslongasthoseofstamensopposingthesepals.Thehairsofvegetativepartsareborneonsmalltubercles,whichpersistafterthehairissloughedoff,sothatthestems,axes,petioles,pedicels,andeventheabaxialcostaandmarginsoflaminasareoftentuber-culate.SuchtuberclesarealsofoundintheMexicanspeciesG. arenicola C.Ander-son, G. hirsuta Cav.,and G. tuberculata (Rose)Nied.Theglandsoflargerlaminasaresometimesdrawnoutintoatooth(Fig.4b).

Galphimia platyphyllaChodat,Arch.Sci.Phys.Nat.,sér.3,24:500.�890.Galphimiabrasiliensisvar.platyphylla(Chodat)Nied.inChodat&Hassl.,Bull.Herb.Boissier,sér.2,7:294.�907.Galphimia brasiliensis f.platyphylla(Chodat)Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 5: 22.�9�4.—type: pArAGuAy.Caaguazú:danslescampos,Caaguazú,Nov�874,Balansa 2396(holotype:P!;isotype:G!). Fig.6.

Subshrubs to 0.5 m, from a woody caudex; stems sparsely pubescent when young, soon becoming glabrous. Vesture of all vegetative parts of mostly wavy or crisped but sometimes straight, reddish brown hairs 0.2–0.7 mm long. Laminas of the larger leaves 3–7.5 cm long, (1.5–) 2–4 (–5) cm wide, length/width ratio 1.2–2.3 (–3), elliptical to lanceolate to ovate to broadly ovate, occasionally suborbicular, obtuse or acute or sometimes apiculate, acute to truncate, glabrous adaxially and abaxially, costa prominent abaxially, the secondary veins barely prominulous or not evident; margin entire; petioles 1.5–6 mm long; leaf glands borne on the margin near the base of the lamina or to 0.5 cm above it, usually a pair, or sometimes with 1 or 2 additional glands, or sometimes only 1 gland or the glands absent, each gland 0.2–0.5 mm in diameter, circular and prominent; stipules 2–4 mm long, 0.6–1.2 mm wide, triangular to narrowly so. Inflorescence a terminal raceme, the axes mostly tomentulose during anthesis; peduncles absent; pedicels 1.5–8 mm long, glabrous; bracts 1–5 (–7) mm long, 0.4–1.2 mm wide, linear to narrowly triangular, bracteoles 0.5–1.5 (–2) mm long, 0.2–0.5 (–0.7) mm wide, linear, bracts and bracteoles abaxially glabrous. Sepals 4–5.7 mm long, 2–2.5 mm wide, elliptical or sometimes oblong, glabrous, glands 0.3–0.6 mm in diameter, each borne in the sinus at the base of 2 sepals, or occasionally glands absent. Pet-als deciduous, subequal, the posterior petal sometimes a little larger, yellow, becoming red in age; claw 2–2.5 mm long, limb 7–8.5 mm long, (4–) 5–5.5 mm wide, triangular-ovate, base acute to truncate, apex obtuse. Stamens with heteromorphic filaments but subequal anthers; filament opposite anterior sepal 3.3–4.5 mm long, filaments opposite anterior-lateral petals 2.7–3.5 mm long, filaments opposite anterior-lateral sepals 3–4 mm long, filaments opposite posterior-lateral petals 2.5–3.3 mm long, filaments opposite posterior-lateral sepals 2.8–3.7 mm long, filament opposite posterior petal 2.5–3.5 mm long. Ovary glabrous; styles subequal, 4.2–5.3 mm long. Cocci 4–5 mm long, 3–3.7mmindiameter,glabrous.

2005 C.ANDERSON:GALPHIMIAINSOUTHAMERICA ��

FIG.6.Galphimia platyphylla.a.Habit.b.Laminabase,showingmarginalglands.c.Oldflower(thepetalsfallen)borneonpedicel,withportionofinflorescenceaxis;notecalyxglands.d.Petal.e.Stamenopposingposterior-lateralsepal(left)andstamenopposingposteriorpetal(right);abaxialview.f.Gynoe-cium.g.Twococci,adaxialview(left)andlateralview(right).Scalebar=a,4cm;b;8mm,c,d,5mm;e,f,2.7mm;g,4mm.(Basedon:a–c,Hassler 9132;d–f,Pedersen 14726;g,Krapovickas et al. 45943.)

Phenology. Collected in flower and fruit in March and from July throughDecember.

Distribution(Fig.5).EasternParaguay(Amambay,Caaguazú,Canendiyú)andadjacentBrazil(MatoGrossodoSul);campo,cerrado,campolimpo,openwood-lands;300–400m.

AdditionAl specimens exAmined.Brazil.mAto Grosso do sul:Mpio.Camapuã,CapãoRedondo,Hatschbach 33058 (MBM,NY);Mpio.PontaPorã,Pacori,Hatschbach 45921 (BR,G,MBM,MICH);Amambai,rod.p/PontaPorã,Hatschbach 48498(MBM,MICH);Coxim,5kmS,Oliveira 162(MBM);Mpio.GuiaLopesdeLaguna,onrd(BR-267)fromMaracajútoGuiaLopes,ca.�2kmWofErvania,Pedersen 14726(C,MICH). paraguay.AmAmbAy:SierradeAmambay,Hassler 9884(BM,G,NY,W);alrededoresdePedroJuanCaballero,ruta5,Krapovickas et al. 45943(CTES,G,MICH).—cAAGuAzú:propeCaaguazú,Hassler 9132(BM,F,G,GH,K,NY,P,S,W);Ruta2,Pastoreo,KmA-�97,Krapovickas & Cristóbal 13355(CTES);caminoaIhú,�4kmNdeCaaguazú,Schinini et al. 28326(CTES);Cnia.Pindo,caminoentreItaquyryyCuruguati,Schinini & Caballero M. 30112(CTES,MICH);nearJ.E.Estigarribia,Zardini & Guerrero 49141(MICH).—cAnendiyú:inregionesYerbaliumdeMaracayú,ApepúetTajai-raguay[?],Hassler 4327(BM,F,K,NY,P,W);iteradYerbalesmontiumSierradeMaracayú,inregionefluminisCapibary,Hassler 4393(BM,G,W);MbaracayúNaturalReserve,aroundAguará-Ñú,Zardini & Ramírez B. 51023(MO).—Depto.unknown:“AltoParaná,”22–23°S,Fiebrig 6348(G).

Galphimia platyphyllaisdistinctiveinitsthickandbroadleaves.Thelaminasrangefrombroadlyellipticalorbroadlylanceolatetobroadlyovateorevensuborbicular,areonly�.2–2.3timesaslongaswide,andoftenhaveanobtuseapex.Comparedto

�2 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME24

G. australis,theflowersandfruitsarelarger,andtheplantsareapparentlyshorter(to0.5mvs.to�mtall)andmorerobust,i.e.,theaxestendtobestouter.Futurefieldworkmayuncoverecologicaldifferencesbetweenthesetwospecies.

excluded nAme

Galphimia pubescens(Adr.Juss.)Herter,Fl.illustr.Uruguay�4(��):49�.�956.—Thecombinationisnotvalidlypublished,becauseHerterdidnotcitethebasionym(Article33.3).

ACKNOWLEDGMENTS

I thankWilliam R.Anderson and David Johnson for their advice and helpful comments on themanuscript.KarinDouthitdrewthehandsomeillustrations.Thecuratorsofthefollowingherbariakindlypermittedaccesstotheircollectionsand/orprovidedloans:A,BM,BR,C,CAS,CEPEC,CTES,ENCB,F,G,GH,GOET,HB,K,MICH,MBM,MO,NY,P,RB,S,SP,UB,US,W,WIS.

LITERATURECITED

Castro,M.A.,A.S.Vega,andM.E.Múlgura.2001.StructureandultrastructureofleafandcalyxglandsinGalphimia brasiliensis(Malpighiaceae).Amer.J.Bot.88:�935–�944.

Chodat,R.�892.Malpighiacées.Mém.Soc.Phys.Genève3�(2:3):�–27.Jussieu,Adr.de.�833[“�832”].Malpighiaceae.InFlora brasiliensis meridionalisbyA.deSaint-Hilaire,3:

5–86,t.�6�–�80.Paris:A.Belin.Niedenzu,F.�9�4.Galphimia. ArbeitenBot.Inst.Königl.LyceumHosianumBraunsberg5:2�–28.———.�928.Malpighiaceae.InDas Pflanzenreich,ed.A.Engler,IV.�4�:�–870.Leipzig:WilhelmEngel-

mann.