revision of galphimia (malpighiaceae)

2007 C.ANDERSON:GALPHIMIA �

REVISION OF GalphImIa (malpIGhIacEaE)

ChristianeAndersonUniversityofMichiganHerbarium

3600VarsityDriveAnnArbor,Michigan48�08-2287

AbstrAct. Galphimia (Malpighiaceae, Byrsonimoideae, Galphimieae) comprises 26 species ofsuffrutescentherbs,shrubs,andtreelets.Fourspecies(G. amambayensis, G. australis, G.brasiliensis, G. platphylla)occurinSouthAmericasouthoftheAmazonbasin,andtheremainderinMexico;onespecies(G. angustifolia)extendsintoTexasandone(G. speciosa)intoCentralAmerica.Onespecies,G. gracilis,iscultivatedworldwideinwarmregions.Thegenusischaracterizedbybilaterallysymmetricalflowerswithyellowtoredpetals;ineightspeciesthepetalsarepersistent.Thesepalslackoilglands,butafewspeciesbearcalyxglandsthatresembletheleafglands.Thefruitisagloboseschizocarpbreakingapartintothreesmall�-seededcocci.Thebasechromosomenumberisx=6.Foreachspeciesafullsynonymy,detaileddescription,andrangemapareprovided;allbuttwospeciesareillustrated.Threenewspecies(G. calliantha, G. floribunda,G. speciosa)areproposed.

INTRODUCTION

The Neotropical genus Galphimia Cav. (Malpighiaceae, tribe Galphimieae)comprises26speciesoflargeherbs,shrubs,andtreeletsofmostlydrierhabitats.MostspeciesoccurinMexico,withoneextendingtoTexasandonerangingtonorthernNicaragua,but fourare restricted toSouthAmerica, southof theAmazonbasin.Galphimia ischaracterizedbyflowerswithyellowpetals(oftensuffusedwithred)borneinracemes,eithersolitaryorarrangedinpanicles;ineightspeciesthepetalsarepersistentinfruit.ThesepalslackthelargeoilglandstypicallyfoundinotherNewWorld Malpighiaceae, but in a few species the calyx has one or more smallglands that resemble the leaf glands.Ten fertile stamens surround a tricarpellateovarywiththreesubulatestyles.Theglobosefruitisaschizocarpthatbreaksintothree�-seededcocci.

ThehistoryofGalphimiaisoneofconfusion,causedinpartbymisinterpretationoftheearliestdescribedtaxa,competinggenericnames(GalphimiaCav.vs.Thryal-lisL.),andmisapplicationofvariousnamestoonecultivatedspecies(G. gracilis),but also by the superficial morphological similarity of many of the species. Theshowyinflorescencescatchtheeyeofcollectors,whooftenlabelallspecimensfromMexicoasG. glaucaorG. gracilis,andthosefromSouthAmericaasG. brasiliensis.Thisrevisionbuildsonpreliminarystudiesofthegenus(C.Anderson2003,2005a,b)andclarifiesthecircumscriptionsofthespeciesandassociatednomenclature.Itisintendedasabasisfordetailedfuturestudies.Manyspeciesarestillknownfromonlyfewcollections.Questionsremainconcerningthephylogeny,phytogeographichistory,pollinationanddispersalstrategies,ploidallevels,aswellasdevelopmentalaspects,suchasthenatureoftheunusualcalyxglands,theshifttopersistentpetals,andthesuppressionofthepeduncle.

TAXONOMICHISTORY

ThenomenclatureofGalphimiaandtheapplicationofnameshavebeenprob-lematical,partlyowingtouncertaintyaboutthestatusofthegenericnameThryallisL.andpartlytodoubtfulinterpretationoforiginaldescriptionswithoutaccesstotype

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Contr.Univ.MichiganHerb.25:�–82.2007.

2 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25

material.NowThryallisMart.hasbeenconservedoverThryallisL.,asynonymofGalphimiaCav.,andtheopportunitytostudytypesallowsforassignmentofnamesasintendedbytheoriginalauthors.

The taxonomic history of the genus now accepted as Galphimia begins withLinnaeus’s(�762)descriptionofhisnewgenusThryallis,containingonlyT. brasil-iensis.In�799CavanillespublishedthegenusGalphimia(ananagramofMalpighia)toaccommodatetwospeciesfromMexico(G. glauca, G. hirsuta)towhichheaddedG. glandulosain�80�.Lindley(�828)andMartius(�829)addednewspeciesfromBrazil toThryallis, andMartiusalsoprovidedagenericdescription reflectinghisconceptofThryallis.ItwasAdriendeJussieu(�833),thefirstmonographeroftheMalpighiaceae,whorealizedthatT. brasiliensis isnotcongenericwithLindley’sT. brachystachys andMartius’sT. longifoliaandT. latifolia.HetransferredT. brasiliensis toGalphimiaCav.,asG. brasiliensis(L.)Adr.Juss.,andrecognizedThryallissensuMartius,inwhichheincludedT. longifolia,T. latifolia,andT. brachystachys (Jussieu�840).

MostbotanistsfollowedJussieu’sleadinrecognizingGalphimiaandThryallissensuMartius(e.g.,Grisebach�858)untilKuntze(�89�)optedtoretainThryallisL.forallthespeciesthathadbeenplacedinGalphimiaandproposedthenewgenusHemsleyanatoreplaceThryallissensuMartius.Kuntze’sviewswereacceptedmostnotablybyRose(�909)andSmall(�9�0),buttheywererejectedbyNiedenzu(�9�4,�928),whoacceptedJussieu’sgenericclassification.ThecontroversywasnotsettleduntilaproposalbyMortonandCuatrecasas(�967)toconserveThryallissensuMar-tiusoverThryallisL.wasacceptedbytheNomenclatureSectionofthe��thInterna-tionalBotanicalCongressin�969.ThryallisMart.thusisnowincludedinthelistofconservednamesoftheInternational Code of Botanical Nomenclature(McNeilletal.2006).ItisaSouthAmericangenusoffivespeciesnotassociatedwiththeGalphi-mieae(C.Anderson�995).Yet,theconfusionconcerningthecorrectgenericnamepersists,particularlyinthehorticulturaltrade,andnamesinThryallisarestillusedoccasionallyforspeciesofGalphimia.

OnlytwoauthorsprovidedcomprehensiveaccountsofGalphimiaafterJussieu’s(�843)treatmentandRose’s(�909)briefsynopsis:Smallin�9�0(asThryallis)forthe series North American Flora (thus excluding the SouthAmerican taxa), andNiedenzu(asGalphimia)in�9�4asamonographandin�928aspartofhistreat-mentofMalpighiaceaeforDas Pflanzenreich.Regrettably,theworkofbothauthorswashinderedbyapaucityofgoodcollectionsandalackofopportunitytostudythemajorityof types.Asaresult, theirpublicationsareflawedand,asprimaryrefer-ences,contributedtothemanymisunderstandingsofspeciescircumscriptionsandnomenclaturethathaveplaguedallthoseconcernedwithGalphimiasincetheearlypartofthe20thcentury.

AtthetimethatJussieudefinedGalphimia,thegenusincludedonlythethreespeciesofCavanillesinadditiontoG. brasiliensis.TotheseBartling(�840)addedG. gracilis, G. grandiflora,andG. paniculata(hisnamesG. latifoliaandG. humboldtianaaresynonymsofG. paniculata)andJussieu(�843)describedG. multicaulis inhismonographofthefamily.ExplorationsofsouthernSouthAmericaandoftheAmeri-canWestandMexicointhelate�800sandearly�900sresultedinthediscoveryofadditionalspeciesfromthoseregions.Chodat(�890)describedG. australisandG. platyphyllafromParaguay.BythetimeRose(�909)publishedhisreviewofthegenus(asThryallis)fivemorespecieswereknownfromMexico:G. angustifolia(includ-ing G. linifolia;alsoinTexas),G. montana,G. sessilifolia,G. tuberculata(includingG. humilis),andG. vestita;however,theOaxacanendemicG. elegans,whichBaillon(�874)basedoncultivatedplants,wasoverlooked.Blake(�9�7)describedanother

2007 C.ANDERSON:GALPHIMIA 3

Mexican species, G. langlassei (as Thryallis langlassei). My studies (C.Anderson2003,2005a,b)addedonenewspeciesfromParaguay(G. amambayensis)andfivefromMexico(G. arenicola, G. mexiae, G. mirandae, G. oaxacana, G. radialis);threenewspecies,G. callianthafromGuerrero,G. floribundafromwesternMexico,andG. speciosafromsouthernMexicoandCentralAmerica,areproposedhere.Galphimianowincludes26species.

MORPHOLOGY

Habit and vesture.Most speciesofGalphimiaareshrubs thatmayreach4mandsomeformtreelets(to6m),butthosewithoutpedunclesaremostlylarge,oftensprawling,suffrutescentherbsarisingfromawoodybaseorsmallshrubsto�.5mhigh.

Inmostspeciesthevestureiscomposedofmedifixed,straighttowavyorcrisped,reddishbrown,sessileorsubsessilehairstoca.�mmlong.Inafewspeciesthemedi-fixedhairsareborneonpersistenttubercles,whichroughenthestructureafterthehairisshed(Fig.4c).Suchtuberclesmaybestronglyexpressed,asinG. tuberculata,orsubtlyandthenareseenonlywithmagnification,asinG. brasiliensisandthepeti-olesofG. gracilis.Basifixedhairsoccurontheovaryandfruitofsomespecieswithdeciduouspetals.Thehairsmaybeevenlydistributed(butsometimesabsentneartheapex)orrestrictedtothesutures.

In general, the stems and foliage are pubescent when very young but soonbecomeglabrescenttoglabrousatmaturity;theinflorescenceaxesarecommonlymoderatelytodenselypubescentbutalsoeventuallyglabrescentinage.GalphimiaglandulosaandG. floribunda lacksuchvesture;G. glandulosa isentirelyglabrous(exceptforapiloseovary),andG. floribundamayhaveonlysomescatteredhairsonpartsoftheinflorescenceorlackthem.Galphimiagracilisisunusualinthatthelaminashaveatuftofhairsattheapex;thistuftissloughedoffinageandseenbestinyoungerleaves.Threespeciesareeasilyrecognizedbythevestureofthevegeta-tive parts. In G. elegans, the laminas are abaxially pubescent with reddish brownhairs,whichmaybesloughedoff,andolderleavesmaybeonlypatchilypubescentoreventuallyglabrescent.Youngerleavesarealsoadaxiallysparselypubescent,butthehairsaresoonshed.Frequentlytheepidermisisraisedatthepointofattach-ment,whichisevidentafterthehairhasfallen.GalphimiavestitaandG. hirsutaareabundantlypubescentinallparts,includingtheabaxialsurfacesofthecalyxandthepetals.Galphimiahirsutaiscoveredwithhairsborneonpersistenttubercles.InG. vestitathedensetangledcoveringiscomposedofwhiteortranslucentT-shapedhairs,whichhaveatrabecula�–2mmlongborneonastalkca.0.5mmlong.SuchhairsdonotoccurinanyotherspeciesofGalphimia,althoughtheyarefoundinmanyothergeneraofMalpighiaceae.

Leaves.MostcollectionsofGalphimiaconsistoftheinflorescenceandsubtend-ingshoot,butonlyoccasionallyincludeleavesfromotherpartsoftheplant,whichareoftenlargerthanthoseneartheflowers.Thus,forsomespeciesthemeasurementsgiveninthedescriptionsmaynotreflectthefullrangeofleafdimensions.Theleavesareoppositeand,inmostspecies,petiolate,althoughthepetiolesmaybeveryshort;inspecieslackingpedunclesandalsoonespecieswithpersistentpetals,G. multicaulis,petiolesgenerallyaremuch less than�cm long. InG. sessilifolia thepetiolesareabsentorrudimentary(to�mmlong).Inthemajorityofspeciesthelaminasvaryfromellipticaltolanceolatetoovate,andinsometorhombic(e.g.,G. tuberculata).


Galphimiavestitahaslinearleaveslessthan�cmwide.Intwowidespreadspecies,G. angustifoliaandG. australis,laminasareexceptionallyvariableinshape,fromlineartoovate,sometimesonthesameindidual(Fig.�5a).

The margin is entire, but in species bearing tuberculate hairs the margin ofmatureleaves,atleastproximally,bearsthepersistenttubercles.ThelaminasofG. arenicolacommonlybearelongate,multicellular,epidermalprocessesinadditiontosuchtuberclesalongthemargin.Galphimia oaxacanaisdistinctiveinthatthemarginformsathickened,narrow,light-coloredband.

Allspecieshavecircularleafglands,whichareoftendiagnostic,especiallytheirsizeandposition,exceptinG. speciosa,inwhichtheplacementoftheleafglandsisvariable,evenalongthesameshoot(Fig.27c).Ingeneral,apairofglandsisborneonthelaminamargin,nearoratthebase,oronthepetiole,mostlyinthedistal�/2.Galphimiaglandulosaisdistinctiveinthattheleafglandsareusuallyborneintheproximal�/6–�/2ofthepetiole(Fig.7b,c).Theglandsmaybeflushwiththesurfaceorbarelyraised(e.g.,G. gracilis,Fig.�b),butmorecommonlyareprominent(e.g.,G. langlassei,Fig.9b)topeglike(e.g.,G. tuberculata,Fig.4b,e).Sometimes,oneormoresmalleradditionalglandsarefoundalongthemargin,distallyfromthemainpair,andsometimesoneorbothglandsareabsent.InG. sessilifoliaonlythefirstpairofleavesonashoothasoneortwoglands(Fig.�3b),thoughsometimesit,too,iseglandular;thisfirstpairalsodiffersfromotherleavesonthesameshootinhavingrelativelysmall,orbicularlaminas.

Thestipulesaredistinct,intrapetiolar,lineartonarrowlytriangular(e.g.,Fig.�b,20b),andpersistentonthestems.Inmostspecies,stipulesare�–3mmlong,butinG. radialistheyarecommonly3.5–6mmlong(Fig.�2c)andinG. vestita3.5–9mmlong(Fig.�7b).

Inflorescences.ThebasicpatterninGalphimia,asinmanygeneraofMalpighia-ceae,isasinglefloweronapedicel,whichisborneonapeduncle;thisunitrepresentsareducedinflorescence.Apairofbracteolessubtendsthepedicel;inmanygenerathebracteolesareborneatthebaseofthepedicel,butinGalphimiatheyarefoundonthepeduncle,neartheapex(e.g.,G. gracilis,Fig.�c)or,morecommonly,betweenthebaseandtheapexofthepeduncle(e.g.,G. grandiflora,Fig.34c).Abractalwayssubtendsthebaseofthepeduncle.Bractsandbracteolesarepersistent.Insixspeciesthepeduncleislostorrudimentary,andthepedicelisthensubtendedbythreestruc-tures,thebractandthepairofbracteoles(e.g.,G. angustifolia,Fig.�5f,j;G. vestita,Fig.�7h);inG. angustifoliaandG. brasiliensisthepeduncleismostlynotevidentbutsometimesexpressedinsomeinflorescences.Thebractsandbracteolesarepersistent;theyareeventuallysloughedoffinoldinflorescences.

The 5-merous, bilaterally symmetrical flowers are borne in terminal racemes,whicharesolitaryinmostspeciesorgroupedinternatepanicles,especiallyinrobustindividuals. In a few species, e.g., G. paniculata, the racemes are aggregated intolargepanicles,andmostoftheflowersarecrowdedalongtheaxes.Inspecieslack-ing peduncles the flowers are more widely spaced, often separated by a distanceapproximatingthelengthofthepedicel(e.g.,Figs.�5a,�8a);exceptinG. platyphylla, thesmallflowerscharacteristicofthesespeciesandtheiropeningonlyonetothreeatatimeaddtothesparserappearanceoftheracemewhencomparedtotheinflores-cencesoftheremainderofthegenus.Inspecieswithpersistentpetalstheflowersareusuallycloselyspacedalongtheaxes,andamongthesespeciesthepersistentpetalsaddtothedenseaspectoftheinflorescences(e.g.,Figs.33a,36a).

Theinflorescenceaxesaresparselytodenselypubescentduringanthesis,butthevesturemaybeabradedorsloughedoffinage.Thepedunclesandpedicelsarealso


usuallysomewhatpubescent,andwithinaninflorescencethepedicelsarecommonlymoredenselysothanthepeduncles.Frequently,thelineindicatingthearticulationbetweenthepedicelandpeduncleiscoveredawithdenseringofhairs.Inmanyspe-ciesthepedicelshavethehairsconcentratedinalineextendingfromthecalyxtothepeduncle.Twospeciesaredistinctiveintheirlackofvestureintheinflorescence:G. glandulosa,whichisentirelyglabrous(exceptforapiloseovary),andG. floribunda,inwhichtheracemesareglabrate(withafewscatteredhairs)toglabrous.

Calyx.Thecalyxiscomposedoffiveoblongtoellipticalsepals,whichmaybedistallyminutelydenticulateorminutelyeroseandsometimesciliate;inG. callianthathemarginisfringed.Thesepalsaremostlyglabrous,butpubescentinG. amambay-ensis, G. hirsuta,andG. vestita.Thespecieslackingpedunclesallhaveatuftofhairsattheapexofthesepal(e.g.,G. angustifolia,Fig.�5f);thistuftisseenbestinbudsandrecentlyopenedflowers.

Galphimia lackstheoilglandsthatarefoundonthecalyxofmostgeneraofNewWorldMalpighiaceae,andmostspecieshavelargeanthers,whichareassociatedwithpollinationsyndromes inwhichpollen is thereward.Yet, somespecieshaveglandsatthebaseofthesinusbetweenadjacentsepals(e.g.,Figs.3d,7d,20c),whichresembletheleafglandsinappearanceandnottheoilglandsfoundinothergenera.ThenatureoftheseglandsinG. australis(as“G. brasiliensis”)wasinvestigatedbyCastroetal.(200�).

Calyxglandsoccuronlyinninespecies,allwithdeciduouspetals(G. arenicola, G. glandulosa, G. hirsuta, G. montana, G. oaxacana, G. mirandae,andG. tuberculatainMexico,andG. australisandG. platyphylla inSouthAmerica).Theexpressionofcalyxglandsisoftenunstable,andonlyG. hirsutahasfivecalyxglandsineveryflower;however,G. hirsutaisknownfromonlythreecollections.InG. arenicola, G. glandulosa,andG. oaxacana thenumberofglandspercalyxvariesfromonetofive.InG. australis,G. montana, G. platyphylla,andG. tuberculatasomeorallflowersinaninflorescencemaylackcalyxglandsorbearonetofiveglands.OnlyonecalyxofonespecimenofG.mirandaeboreonegland,butforthisspecies,too,onlylimitedmaterialisavailable.

Corolla.The corolla is composed of five clawed petals, which are yellow, butoftenmarkedorsuffusedwithred,andglabrous,butwithhairsonandadjacenttotheabaxialmidribinG. hirsutaandG. vestita,twospeciescharacterizedbyabundantvesture,andinG. amambayensis.Inthemajorityofspecies,thepetalsaredeciduous.Eightspecies(G. calliantha, G. elegans, G. floribunda, G. glauca, G. grandiflora, G. multicaulis, G. paniculata, G. speciosa)havepersistentpetals,whichstiffenasthefruitmaturesandremainevenafterthecoccihavefallen(e.g.,Fig.3�d).Surprisingly,noneoftheauthorswhohavepublishedonGalphimiaremarkedonthishighlyunusualphenomenon,eventhoughsome(e.g.,Cavanilles�799;Bartling�840;Baillon�874)observedlivingplantscultivatedinbotanicalgardens.

Thebasiccorollapatternisfourequallateralpetalsandoneposteriorpetal(the“flag”petal)withalongerandstouterclawanddifferentlyshapedlimb.Thespecieslackingpedunclesallhavesubequalpetals,i.e.,theposteriorpetalislikethelateralonesthoughsometimesalittlebitlarger.ExceptforG. platyphylla,thesespecieshaveverysmallpetals,thelimbonlyto5mmlong.Onlyonespecieswithpeduncleshassubequalpetals,G. radialis.

Androecium.Theandroeciumiscomposedof tenglabrousstamens.Thefila-mentsareofunequallength.Thefilamentoppositetheanteriorsepalisthelongest,


thatoppositetheposteriorpetaltheshortest;thelateralfilamentsareofintermediatelength.OnlyG.brasiliensisandG. vestitadepartfromthispattern.InG. brasiliensisthefilamentsofstamensoppositethelateralsepalsareequaltoeachotherandonlyslightlyshorterthanthatoppositetheanteriorsepal,andthefilamentsofstamensoppositeallpetalsareequal.InG.vestitathefilamentsareallequallylong.Adjacentfilamentsareusuallyverybrieflyconnateatbase.

Theanthersaresubequalwithinanandroeciumandopenbylongitudinalslits.Inmostspeciestheyrangefrom2.5to3.5mmlong(G. elegans:�.8–2mm;G. glandulosa:3.3–4.4mm),butinspecieslackingpedunclestheyaremuchshorter(G. angustifolia, G. brasiliensis:0.7–0.9mmlong;G. amambayensis, G. australis, G. platyphylla, G. vestita:�–�.5mmlong).

Gynoecium.Theovaryistricarpellateandbearsthreesubequal,subulatestyleswithaterminalminutestigma.Twocarpelsareanteriorandoneposterior,ontheplaneofsymmetry.Theovaryisalwaysglabrousinspecieswithpersistentpetals.Tenspecieshaveapiloseovary;inseventhehairsareevenlydistributed(G. amambayen-sis, G. angustifolia, G. brasiliensis, G. glandulosa, G. langlassei, G. mexiae, G. vestita),butinthreethehairsareconcentratedalongthesutures(G. oaxacana, G. radialis, G. tuberculata).OvarypubescenceisvariableinG. montana;insomespecimenstheovariesbearsomehairsalongthesuturesandinotherstheyareglabrous.Thehairsarebasifixedandinmostspeciesca.0.�–0.3mmlongandthuseasilyoverlooked,especiallyinspecieswithhairsmostlyonthesutures;inG. radialisthehairsareupto0.8mmlong.

Fruits.Thefruit isaschizocarpthatbreaks intothreecocci.Eachpebble-likecoccuscontainsagloboseseedwithadarkbrown,smoothseedcoat.Theglobosetoellipsoidembryoisfoldedinhalf,thelargeroutercotyledonenfoldingthesmallerinner one. Cocci generally are from 3 to 5 mm long, but only 2.5 mm long in G. glauca.Thecocciaresmoothtosomewhatrugosewhendry,andglabrousorminutelypilose(liketheovaries),andlackanyobviousdispersalmechanism.Theyfallintothe“detritus fruit” category, i.e., small pebble-like units blown about by wind ordispersedbywater.Inherbariumspecimenswithmaturefruits,thecocciareoftenslightlydehiscentalongthedorsalsuture,althoughtheopeningisnotsufficienttoreleasetheseed;itisnotknownwhetherthisdorsalsplittingisnaturaloranartifactofdrying.Thebaseofthecoccusandtheareoleoftenhavesomeaerenchyma,whichmayprovidesomebuoyancy.

DISTRIBUTION

Galphimia, in general, occurs in dry habitats and has an unusual range.Themajority of species are found in Mexico, with one (G. speciosa) extending intoCentralAmericaandanother(G. angustifolia)intoTexas.TheMexicanspeciesarerecordedmostlyfromdrysituationsinpine-oakforestsandshrublands(“matorral”)andalsodisturbedareas.OnlyG. grandifloraoccursinmesicsitesinforestsdomi-natedbyoaks,pines,andfirs.GalphimiaarenicolahasbeenfoundonlyondunesandincoastaldeciduousforestonthePacificcoastoftheIsthmusofTehuantepec.SeetheappendixforalistofspeciesfoundinMexicoandsortedbystate.

FourspeciesarefoundinSouthAmerica,southoftheAmazonbasin(Figs.�9,2�),allinopenhabitats(openwoodlandsandgrasslands,cerrado,caatinga,campolimpo,camposujo,matorral,andsecondarysites).Galphimia australis,awidespreadandvariable species found fromeasternBoliviaand southernBrazil tonorthern


Argentina,issympatricwithG. amambayensis(knownonlyfromthetypefromeast-ernParaguay)andG. platphylla(easternParaguayandadjacentBrazil).GalphimiabrasiliensisisrestrictedtoeasternBrazil,fromParaíbatoBahiaandadjacentPiauí.The SouthAmerican species are morphologically most similar to G. angustifoliaandG. vestitaofnorthernMexico,whichgrowindryanddeserthabitats,fromthornscrubandtropicaldecidousforesttosandywashesandarroyos;G. angustifoliaisalsoreportedfromlimestone,caliche,andgypsumsubstrates.GalphimiaangustifoliaisacommonspeciesfoundfromcentralTexastoTamaulipasandwesttoCoahuila(andsomecollections from theDurangoborder)aswell as ineasternBajaCaliforniaSurandwesternmostSonoraandadjacentSinaloa(Fig.�6).TherangeofG.vestitaextendsfromSonoratoNayarit,withonecollectionfromsouthwesternChihuahua(Fig.�0).

GENERICANDINFRAGENERICAFFINITIES

ThephylogenyofMalpighiaceaeandtherelationshipsofthegeneraarecur-rentlythefocusofastudybyW.R.AndersonandC.C.Davis;earlierresultswerereportedbyCameronetal.(200�)andDavisetal.(200�).Galphimia isplacedinthetribeGalphimieaeofsubfamilyByrsonimoideae,withLophantheraAdr.Juss.,SpacheaAdr.Juss.,andVerruculariaAdr.Juss.(W.R.Anderson�978).InadditiontothesharedcharacteristicsnotedbyAnderson(�978), thepresenceof latexhasbeenreportedforallfourgenera(Lophanthera, Spachea:W.R.Anderson�98�,200�;Galphimia,Verrucularia:Vegaetal.2002).InareviewofchromosomenumbersofNeotropicalMalpighiaceae,W.R.Anderson(�993)notedthatn=6isthelowestnum-berreportedforthefamilyandthatn=6oramultipleof6istypicalformembersofsubfamilyByrsonimoideae.CountsreportedforLophantheraandVerruculariaaren=6;noneareknownforSpachea.FewcountsareavailableforGalphimia,buttheyindicatethatalsoinGalphimiax=6.W.R.Anderson(�993)predictedthatthelownumberandmorphologicalevidenceplacetheGalphimieaenearthebaseofthefam-ilyphylogeny,apositionconfirmedbyCameronetal.(200�)andDavisetal.(200�).

TwovoucheredcountsofG. gracilisaren=�2.Oftheseonlyonewasobtainedfrombudscollected in thefield(Fryxell & Anderson 3483;W.R.Anderson�993)andtheonlyoneforwhichIsawthevoucher.MacBryde(�970)alsocitedavoucherforhis cultivated source (MacBryde & Herrera-MacBryde 63).Literature reportsofcountsn=�2fromcultivatedplantsofGalphimiamostlikelyapplytoG. gracilis;however,gardenspecimensofMalpighiaceaeareoftenmislabeledeventogenus,andintheabsenceofvoucherstheidentityofsourcesremainsuncertain.LombelloandForni-Martins(2002)report2n=24for“G. brasiliensis.”Thevoucher(Lombello 50,UEC,notseen)wascollectedfromplantscultivatedinCampinasandlikelyrepre-sentsG. gracilisorperhapsG. australis.MacBryde(�970)andSemple(�970)listedin thesamesummaryofchromosomereportsdifferingcounts forG. angustifolia.MacBrydecitedn=�2(voucher:Lynch 710,MO!)andSemplelistedn=20(voucher:MacBryde 72,notseen).Bothvoucherswerepreparedfromtransplantedindividu-alscollectedinTexas.Threevoucheredcountsfor“G. glauca,”alln=6,havebeenpublishedfortwospecieswithpersistentpetals:G. floribunda(voucher:Breedlove 19114,Seavey�975)andG. speciosa(voucher:Breedlove 7072,Kyhos�966;voucher:Anderson 13555,W.R.Anderson�993).Averypreliminarymolecularsurveyofspe-ciesofGalphimia(C.C.Davis,pers.comm.)indicatesthatG. gracilismaybebasaltotherestofthegenus.Itisintriguingthatspecieswithpersistentpetals,whicharepresumedtobeamongthemorerecentlyderivedinthegenus,havealowerchromo-somenumberthanthetwospecieswithdeciduouspetals.


AreliablephylogenyofGalphimiaisnotyetavailable;therefore,thespeciesofGalphimiaarehereassignedtothreeinformalgroupsandwithinthesegroupsareplacedaccordingtomorphologicalsimilarity.GroupsIIandIIImayprovetoformderivedclades.ItispossiblethatthetwoNorthAmericanspeciesandthefourSouthAmericanspeciesofGroupIIrepresenttwoseparatemigrations;possiblyG. brasil-iensis,whichdiffersnotablyfromtheotherthreeSouthAmericantaxa,representsathirdevent.

GroupIcomprises�2speciesfromMexicoandisthemostdiverse.Allspecieshavedeciduouspetalsandwell-developedpedicelsandpeduncles.InallbutG. radia-listheposteriorpetaldiffersfromthelateralones.Somespecieshaveanentirelyorpartiallypiloseovaryandfruit,andsomehaveaglandularcalyx.Fourspecies(G. are-nicola, G. gracilis, G. hirsuta, G. tuberculata)havethevesturecomposedofhairsborneonpersistenttubercles;thisconditionisnotasstronglyexpressedinG. gracilisasintheotherthree.Threespecieshaverelativelylong,elongatelaminaswiththeglandsborneonthemargin(neveronthepetiole);ofthese,G. langlasseiandG. mexiaehavepiloseovaries,butinG. radialissuchhairsarefoundonlyonthesutures.

Group II includes the four SouthAmerican species, and two from northernMexico(G. angustifoliaintoTexas, G. vestita).Inall,thepeduncleislostorrudimen-tary,andthewand-like,elongateracemesbearwidelyspacedflowerswithsubequalpetals;usuallyonlyonetothreeflowersareopenatthesametime(uptoseveninG. platyphylla).Thesepalsbearanapicaltuftofhairs,whichiseventuallysloughedoff.Theflowers,exceptinG. platyphylla,aresmall,withpetalsonlyto5mmlong,andallhavesmallanthers,to�.5mmlong.TheovaryandfruitareglabrousonlyinG. australisandG. platyphylla,theonlyspecieswithcalyxglands.Galphimia vestitahaslinearleaves,ashapefoundalsointhevariableG. angustifoliaandG. australisbutinnootherspeciesinthegenus.Galphimiabrasiliensisistheonlyspeciesofthisgroupwithhairsborneonpersistenttubercles.

GroupIIIcompriseseightspeciesthathavepersistentpetals.Theflowersarecloselyspacedalongtheinflorescenceaxis,andthecrowdedaspectoftheracemesisenhancedbythelargepetals,whichareretainedbeyondmaturationofthefruit.Allspecieshavewell-developedpeduncles,aneglandularcalyx,andaglabrousovaryandfruit.Thelateralpetalsareofadifferentshapeandsizethantheposteriorone.Overall,theflowersarethelargestinthegenus.Thelimbofthepetalsismostlylon-gerthan7mmandinG. grandifloraitmeasuresto�5.5mmlong.Theseeightspeciesmaybethemostrecentlyderived.Theyshareanoverallsimilaraspectandarethetaxamostoftenmisdetermined;manycollectorslabelanyspecimenwithpersistentpetals“G. glauca.”

TAXONOMY

Note:MeasurementsoffloralpartsweretakenfromflowersrehydratedwithPohl’ssolution(Pohl�965).

GalphimiaCav.,Icon.5:6�.�799.Malpighia[subgenus]Galphimia(Cav.)Persoon,Syn. pl. �: 506. �805.—Lectotype, designated by O’Donell and Lourteig,�943:Galphimia glaucaCav.

ThryallisL.,Sp.pl.(ed.2)554.�762,nom.rej.,nonThryallisMartius,�829,nom.conserv.VorstiaAdanson,Fam.pl.2:(23).�763,nom.superfl.Galphimiasect.MicrogalphimiaNied.,ArbeitenBot.Inst.Königl.LyceumHosianumBraun-sberg5:2�.�9�4,nom.superfl.—type:Thryallis brasiliensisL.[=Galphimia brasiliensis(L.)Adr.Juss.].


Suffrutescent herbs arising from a woody base, subshrubs, shrubs, or treelets.Leavesopposite,petioledorsubsessiletosessile;laminalinear,lanceolate,elliptical,ovate,rhombic,obovate,oroccasionallysuborbicular,apexacuteorapiculate,baseacute,attenuate,cuneate,ortruncate,narrowlydecurrentornot,glabrous(pubescentinG. hirsutaandG. vestita,sparselysoinG. elegans),marginentire(withtuberclesinG. arenicola,G. gracilis,G. hirsuta, andG. tuberculata,withelongatemulticellularoutgrowthsinG. arenicola),costaprominentabaxially,secondaryveinsprominentorprominulousornotevident;stipulesdistinct,intrapetiolar,linearornarrowlytrian-gular,persistentonstems.Leafglandsusually2andpaired,disklikeandprominentorflushwiththesurface,orpeglikeandstalked,borneintheproximalportionofthelaminaonthemarginnearoratthebase,orborneonthepetiole,thepairofglandssuboppositeorseparated,sometimeswithadditionalglandsonthelaminarmarginorwithonly�gland,orsometimesglandsabsent.Inflorescenceaterminalracemeorapanicle;flowersborneonapedicel,pedunclespresentandsubtendedbyabractandbearingapairofbracteoles,thearticulationcommonlymarkedwitharingofhairs,orpeduncleabsentorrudimentaryandthepedicelsubtendedbyabractandapairofbracteoles.Flowersbilaterallysymmetrical.Sepals5,oblong,elliptical,orlanceolate,eglandularorwithaglandatthebaseofthesinusbetweenallorsomeadjacentsepals.Petals5,clawed,subequalortheposteriorpetal(the“flag”)differ-ingfromthelateralones,glabrous(pubescentonthemidveinabaxiallyinG. hirsutaandG. vestita,alsosometimesinG.amambayensis),yellowandoftensuffusedwithred,deciduousorpersistent,thelimbellipticaltoovateortriangular.Stamens�0,glabrous,filamentsdifferinginlength(equalinG. vestita),thatoppositetheanteriorsepalthelongest,thoseoppositetheposteriorandposterior-lateralpetalstheshort-est,antherssubequal,dehiscingbylongitudinalslits.Ovaryspherical,tricarpellate(2carpelsanterior,�carpelposterior),trilocular(all3loculesfertile),glabrousorfinelypilosewithbasifixedhairs;styles3,subulate,glabrous;stigmaminute,terminal.Fruitaschizocarp,splittinginto3cocci,theseglabrousorpiloseliketheovary,carpophoreabsent;seedcoatshiny(dark)brown;embryogloboseorsubglobose,radiclestraight,cotyledonsoblongandfoldedattheproximal2/3–2/5,theoutercotyledonlargerthanandenclosingtheinnerone.Basechromosomenumber:x=6.

Key to the species of GALphimiA

�. PlantsofSouthAmerica.2. Sepalsabaxiallytomentoseorpatchilysoinfruit;ovaryandfruitpilosealongthesuturesand

ontheproximal2/3ofthesurface,thedistal�/3glabrous;Paraguay(Amambay). �7.G. amambayensis

2. Sepalsglabrousorwithscatteredhairsnearthebaseand/orwithatuftofhairsattheapex;ovaryandfruitglabrousoruniformlypilose.3. Ovaryandfruitpilose;petioles3–�0mmlong,tuberculate-strigose,orinolderleavesonly

the persistent tubercles remaining; Brazil (Bahia, Paraíba, Pernambuco, and adjacentPiauí). �5.G. brasiliensis

3. Ovaryandfruitglabrous;petioles�.5–6mmlong,glabrous.4. Laminas linear to linear-lanceolate to lanceolate to narrowly elliptical, (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;commoninsouthernBrazil,easternBolivia,Paraguay,northeasternArgentinaandadjacentUruguay. �6.G. australis

4. Laminaselliptical,broadlylanceolate,ovatetobroadlyovate,occasionallysuborbicular,(�.5–)2–4(–5)cmwide,length/widthratio�.2–2.3(–3);pedicelsglabrous;sepals4–5.7mmlong,2–2.5mmwide;petallimb7–8.5mmlong;cocci4–5mmlong;Paraguay(Amambay,Caaguazú,Canendiyú)andadjacentBrazil(MatoGrossodoSul).

�8.G. platyphylla

�0 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25

�. PlantsoftheU.S.A.(Texas;G. angustifolia),Mexico,andCentralAmerica(G. speciosa).5. Inflorescenceaterminalracemewithonly�–2(–3)flowersopenatonetime,flowerswidely

spacedalongtheaxis,usuallythedistancebetweenflowersaboutaslongorlongerthanthelengthofthepedicels;pedunclesabsentorrudimentary;petalsdeciduous,subequal;anthers0.7–�.5mmlong.6. Leavesglabrousorwithafewscatteredhairs;sepalsglabrousexceptforanapicaltuftof

hairs(bestseeninbudandyoungflowers);U.S.A.(Texas)andMexico(BajaCaliforniaSur,Chihuahua,Coahuila,Durango,NuevoLeón,SanLuisPotosí,Sinaloa,Sonora,Tamaulipas).

�3.G. angustifolia6. Leaves densely pubescent; calyx densely pubescent; southwestern Chihuahua, Nayarit,

Sinaloa,Sonora. �4.G. vestita5. Inflorescenceaterminalracemeorapanicle,morethan3flowersopenatonetime,flowers

closelyspacedalongtheaxis,thedistancebetweenflowersvariableandshorterthanthelengthofthepedicels;pedunclespresent;petalsdeciduousorpersistent,theposteriorpetaldifferingfromthelateralpetals(petalssubequalinG. radialis);anthers�.8–4.4mmlong.7. Petiolesabsentorrudimentary(upto�mmlong);laminabaseshallowlycordateortrun-

cate;leafglandspresentonlyonbasalpairofleavesofashoot,theseleavesmuchsmallerthanthedistalonesandsuborbicular;Oaxaca. ��.G. sessilifolia

7. Petiolespresent,atleast3mmlong;laminabaseacuteorcuneateordecurrent(sometimestruncateinG. montana);leafglandsnotrestrictedtobasalmostpairofleavesofashoot.8. Laminasabaxiallypubescent,atleastthoseclosetotheinflorescence.

9. Laminasallevenlypubescentonbothsurfaces;petalsdeciduous,oftenabaxiallypubescent;calyxabaxiallypubescentandglandular;Guerrero. 4.G. hirsuta

9. Laminas,atleastthoseclosetotheinflorescence,abaxiallypubescentorpatchilysoinolderleaves,adaxiallyonlysparselypubescentandglabrescent;petalspersistent,glabrous;calyxwithscatteredhairsandeglandular;Oaxaca. 20.G. elegans

8. Laminasglabrousorwithafewscatteredhairs.�0. Inflorescenceaxes,peduncles,andpedicelsglabrousorwithafewscatteredhairs.

��. Petalsdeciduous,commonlysuffusedwithred;calyxusuallyglandular;ovaryandfruitpilose;leafglandsborneonthepetioleintheproximal�/6–�/2(rarelydistally);Guerrero,Jalisco,México,Michoacán,Sinaloa. 7.G. glandulosa

��. Petalspersistent,yellow;calyxeglandular;ovaryandfruitglabrous;leafglandsborneonthemarginofthelaminaorneartheapexofthepetiole;Chihuahua,Durango,Jalisco,Sinaloa,Sonora,Nayarit. 26.G. floribunda.

�0. Inflorescenceaxes,peduncles,andpedicelsmoderatelytodenselypubescent.�2. Petalspersistentandspreadingbelowthe fruit, retainedevenafter thecocci

havefallen,limboflateralpetals7.5–�5.5mmlong;calyxglandsabsent;ovaryandfruitglabrous.�3. Petiolesrudimentaryorveryshort,eveninthelargestleavesonly0.3–0.8

(–�)cmlong;Guanajuato,Jalisco,MichoacánandadjacentMéxico,disjuncttoOaxaca. 22.G. multicaulis

�3. Petioleswelldeveloped,(0.8–)�–5cmlong(shorteronlyinsmallestleavesofG. glauca).�4. Leafglandsborneataboutthemiddleofthepetiole(sometimesmore

distallyinthesmallestleavesneartheinflorescence);Guerrero,Jalisco,México,Michoacán,Morelos,Nayarit,westernOaxaca,Puebla,Zacatecas.

23.G. paniculata�4. Leafglandsborneonthemarginofthelamina,oratorneartheapexof

thepetiole.�5. Limboflateralpetals�0–�5.5mmlong,6–��mmwide;sepals4–5.5

mmlong;bracts3–6.7mmlong.�6. Sepals with a fringed margin; leaf glands a pair borne well

above the base of the lamina, the glands sessile, ca. 0.5 mmin diameter, often with additional smaller glands distally;Guerrero. 24.G. calliantha

�6. Sepals with a glabrous margin; leaf glands a pair borne atthebaseofthelamina,oftenatthetipofabasaltooth,oratthe apex of the petiole, each gland 0.6–�.3 mm in diameter,prominent topeglike (and thenup to�.5mm long);Colima,Jalisco,México,Michoacán,Morelos. 25.G. grandiflora

�5. Limboflateralpetals7–9.5(–�0)mmlong,5–8.5mmwide;sepals2.5–4mmlong;bracts�.5–3(–4)mmlong.

2007 C.ANDERSON:GALPHIMIA ��

�7. Leaf glands borne above the base of the lamina, often atthe apex of a tooth; limb of lateral petals elliptical to lan-ceolate, limb of posterior petal broadly triangular; cocci ca.2.5 mm long, ca. 2 mm in diameter; inflorescence a terminalraceme; Aguascalientes, Guanajuato, Hidalgo, Jalisco, NuevoLeón, Querétaro, San Luis Potosí, Tamaulipas, Zacatecas.

�9.G. glauca�7. Leaf glands borne at the base of the lamina and/or the

distal portion of the petiole, commonly variable amongthe leaves of a branchlet; limb of lateral petals triangu-lar to ovate, limb of posterior petal broadly triangularto suborbicular; cocci 3.5–4 mm long, ca. 3 mm in diam-eter;inflorescenceaterminalracemeorleafyternatepanicle;Mexico (Chiapas, Oaxaca, Puebla, Veracruz), Guatemala,Honduras,andadjacentNicaragua. 2�.G. speciosa

�2. Petalsdeciduous,limboflateralpetals4.5–8.5mmlong;calyxglandspresentorabsent;ovaryandfruitpiloseorglabrous.�8. Petalssubequal,theposteriorpetalonlyslightlylargerthanthelateralpet-

als;ovaryandfruitwithhairsonthesuturesandintheproximal�/4–�/2,thehairsupto0.8mmlong;stipulesupto0.6mmlong;Guerrero. �0.G. radialis

�8. Petalsunequal,theposteriorpetaldifferingfromthelateralpetalsinsizeandshape;ovaryandfruitglabrousorwithhairsupto0.3mmlong;stipulesto0.3mmlong.�9. Ovaryandfruitevenlypiloseovertheentiresurface.

20. Anthers3–4mmlong;filamentofstamenoppositetheanteriorse-pal3.5–4.5mmlong,filamentofstamenoppositetheposteriorpetalshorter thanthatof stamensopposite theposterior-lateral sepals(�.6–2.6mmvs.3.5–4.6mm); limbof lateralpetals6–7mm long,4.5–5.5mmwide;peduncles0.6–�.2(–�.6)timesaslongaspedicels;Colima,Guerrero,Jalisco,Michoacán. 8.G. langlassei

20. Anthers2.3–2.5 (–2.8)mmlong;filamentof stamenopposite theanterior sepal 2.2–3 mm long, filaments of stamens opposite theposteriorpetalandposterior-lateralsepalssubequal[�.5–�.7(–2)mm vs. �.5–�.8 (–2.2) mm]; limb of lateral petals 4.5–6 mm long,3.7–4.7mmwide;peduncles0.3–0.5(–0.7)timesaslongaspedicels;northwesternJaliscoandadjacentNayarit. 9.G. mexiae

�9. Ovaryandfruitglabrous,orwithhairsonlyonthesutures(inG. tuber-culataalsointhebasal2/3).2�. Secondaryveinsof laminanotorbarelyevidentabaxially;ovary

glabrous.22. Pedicels�4.5–�8.5mmlong;peduncles0.2–0.3timesaslongas

pedicels;sepals3.5–4.2mmlong;laminasellipticaltobroadlyso, obovate, rhombic, ovate, or suborbicular, the base oftendecurrent;Guerrero,Puebla. �2.G. mirandae

22. Pedicels2–�0mmlong;peduncles0.5–3timesaslongaspedi-cels;sepals2.5–2.8mmlong;laminasnarrowlylanceolatetonar-rowlyellipticaltoelliptical,thebasenotdecurrent(orslightlysoinG. arenicola).23. Marginoflaminaentire;bracteolesborneatorjustbelow

theapexofthepeduncle;pedicels5–�0mmlong,peduncles0.5–�timesaslongaspedicels;limbofposteriorpetal7–8.5mmlong;Tamaulipas,SanLuisPotosí,andVeracruz,andadjacentregionsofHidalgo,Puebla,andQuerétaro.

�.G. gracilis23. Margin of lamina (at least near the base) with scattered

tubercles and/or somewhat longer epidermal processes;bracteolesborneintheproximal�/4–�/2ofthepeduncle;pedicels2–5mmlong,peduncles�.2–3timesaslongasthepedicels; limb of posterior petal ca. 4.5 mm long; coastalOaxaca. 6.G. arenicola

2�. Secondaryveinsoflaminaevidentabaxially,prominuloustopromi-nent;ovarywithhairsonsutures(orglabrousinG. montana).


24. Vegetativepartswithhairsborneonpersistenttuberclesand/oronlywiththetubercles(thehairsalreadyshed);branchletscom-monlyroughenedbypersistenttubercles;peduncles�–2.2timesaslongaspedicels;Colima,Guerrero,Jalisco,Nayarit,Sinaloa.

5.G. tuberculata24. Vegetative parts with sessile hairs or glabrous; branchlets

smoothorthebarklightlyfissured;peduncles0.2–0.8timesaslongaspedicels.25. Laminasthick-textured(coriaceouswhendried),abaxially

papilloseortheepidermisatleastblistered,themarginathickened light-colored band to 0.2 mm wide; secondaryveins prominent abaxially; ovary and fruit with hairs onsuturesPacificslopeofOaxaca. 3.G. oaxacana

25. Laminas thin-textured (papery when dried), abaxiallysmooth,themarginnotthickened;secondaryveinsevidentto prominulous abaxially; ovary and fruit with hairs onsutures or glabrous; Colima, Guerrero, Jalisco, Micho-acán, Nayarit, Sinaloa, and southwestern Durango andZacatecas. 2.G. montana

Group i (petalsdeciduous;pedunclespresent)

1. Galphimia gracilisBartling,Linnaea�3:552.�840[“�839”].Thryallis gracilis (Bar-tling)Kuntze,Rev.gen.pl.�:89.�89�.—type:basedonplantscultivatedinthe botanical garden of Göttingen (holotype: unknown).—Neotype, des-ignatedbyCuatrecasasandCroat,�98�:aspecimenpreparedfromplantscultivatedatthebotanicalgardeninParis(P-JU��5�0,microfiche:MICH!,photo:MICH!).

Galphimia gracilisvar.gracillimaHieronymus,Bot.Jahrb.Syst.20,Beibl.49:36.�895.—type:coLombiA.Cauca:specimenscultivatedinCerrito,Lehmann 7439(lectotype,designatedbyCuatrecasasandCroat,�98�:F;isolectotypes:GH!K!S!US!).

Fig.�.

Shruborsubshrubto4m;stemsstrigosetotomentulose,becomingglabratetoglabrous inage.Vestureofallvegetativepartsofstraighttowavy,reddishbrownhairs0.3–0.6(–0.9)mmlong,subsessileoreachhairborneonapersistenttubercleto0.05mmhigh,hairsonthepetioleborneonapersistenttubercleupto0.�mmhigh.Laminasofthelargerleaves2.5–7cmlong,2–3cmwide,ellipticalornarrowlyelliptical,apexapiculateandwithatuftofhairsinyoungleaves(sometimesretainedinmature leaves),baseacute,glabrousorwitha fewhairsscatteredonthecostaabaxially,secondaryveinsnotorbarelyevident,marginentire;petioles0.7–�.5cmlong,tomentulosetoglabrate,irregularlytuberculate;glandsusuallyapairborneonthemarginofthelamina2–�0mmabovethebaseorsometimeswith3–4glands,eachgland0.3–0.4mmindiameter,disklikeandprominentorflushwiththemargin,orsometimespeglikeandca.0.�mmlong;stipules�.3–3.2mmlong,0.4–0.7mmwide,linear,withscatteredhairsattheapexandalongthemargin.Inflorescenceaterminalraceme,theaxestomentuloseorstrigose;peduncles3–6.3mmlong,pedicels5–�0mmlong,bothtomentuloseorstrigose,peduncles0.5–�timesaslongaspedicels;bracts2.5–3mmlong,0.6–0.8mmwide,linear,bracteoles�.4–2mmlong,0.3–0.5mmwide,linear,bractsandbracteolesglabrousorwithscatteredhairsalongthemargin;brac-teolessuboppositeorupto0.5mmapart,usuallyborneatorjustbelowtheapexof

2007 C.ANDERSON:GALPHIMIA �3

FIG. �. Galphimia gracilis. a. Flowering and fruiting branch. b. Base of leaf and stipules, adaxialview.c.Flower,sideview,posteriorpetalatupperright.d.Posteriorpetal(left)andlateralpetal(right),adaxialviews.e.Stamenfromoppositesepal,abaxialview.f.Stamenfromoppositepetal,adaxialview.g.Gynoecium.h.Fruit.i.Coccus,adaxialview.Scalebar:a,3cm;b,c,6mm;d,h,4.2mm;e–g,i,3mm.(BasedonFryxell & Anderson 3484,MICH.)


thepeduncle.Sepals2.5–3.2mmlong,�.2–�.5mmwide,ellipticaloroblong,margindenticulate-erose indistal�/2,glabrous;glandsabsent.Petalsdeciduous,unequal,yellowortheclawssuffusedwithred,glabrous,marginerosetoirregularlydenticu-lateandwithscatteredfimbriae;lateralpetals:claw2–2.5mmlong,0.5–0.7mmwide,limb6–7(–8.5)mmlong,4–4.6(–5)mmwide,narrowlyellipticaltonarrowlyovate,apexobtuse,baseacute;posteriorpetal:claw4–5mmlong,�–�.5mmwide,limb7–8.5mmlong,6–7(–8)mmwide,broadlytriangular,apexbroadlyobtuse,basetruncate.Stamensofunequallength;filamentoppositeanteriorsepal(4.2–)4.5–5mmlong,filamentsoppositeanterior-lateralpetals3–4mmlong,filamentsoppositeanterior-lateralsepals4–5.2mmlong,filamentsoppositeposterior-lateralpetals2–3mmlong,filamentsoppositeposterior-lateralsepals3.5–4mmlong,filamentoppositeposteriorpetal2–2.3mmlong;anthers2.3–3mmlong.Ovaryglabrous;styles(5–)6–7mmlong.Coccus4.5–5mmlong,3–3.3mmindiameter,glabrous;areole3.5–3.7mmlong,2.8–3mmwide;outercotyledon3–4mmlong,innercotyledon2.4–3mmlong.Chromo-somenumber:n=�2(W.R.Anderson�993).

Phenology.Collectedinflowerthroughouttheyear(exceptMarchandOctober),infruitinJanuaryandFebruaryandfromAprilthroughSeptember.

Distribution (Fig. 2). Mexico (Hidalgo, Puebla, Querétaro, San Luis Potosí,Tamaulipas,Veracruz);deciduouslowlandforest,acahuales,roadsides,ofteninwetsituations, such as streambanks and ditches, commonly cultivated and adventiveoutsidethenaturalrange;20–900m.

represeNtAtive specimeNs.mexico.hidALGo:Mpio.Huejutla,Tehuetlán,Hernández M. et al. 6385(CAS,ENCB,MEXU,MO,XAL);Mpio.Huejutla,nearHuejutla,Seler & Seler 660(GH,US).—puebLA:Mpio.Metlaltoyuca,Metlaltoyuca,Turra 2337(ENCB).—QuerétAro:Jalpan,alorientedeTanchanaquito,El Sabinito, López 366 (IEB, MICH).—sAN Luis potosí: Mpio. SanAntonio,Tanjasnec, Alcorn 1836(MEXU,TEX);SofVillaJuárez,Clark 6837(MO);Mpio.CiudadValles,2miWofChontal,onRanchoPagoPago,Fryxell & Anderson 3484(CAS,CHAPA,DUKE,ENCB,MBM,MEXU,MICH,NY,TEX);2miEofTamazunchale,Nsideofriver,Hitchcock & Stanford 7318(DS,UC,US);2miNofTanquian,Johnston & Crutchfield 5675(LL,MEXU,MICH,TEX);Tamazunchale,Lundell&Lundell 7145(MICH,US);Valles,Lundell & Lundell 7275(LL,MEXU,MICH,NY,US);SanDieguito,Palmer 104in�904(F,GH,K,MO,NY,UC,US);RasconStation,Pringle 3099(A,BM,BR,ENCB,F,G,GH,GOET,LL,MEL,MEXU,MO,NY,S,TEX,UC,W);Huichihuayán,Rzedowski 7819(ENCB,MEXU,TEX).—tAmAuLipAs:nearLimón,73miSofCiudadVictoria,Frye & Frye 2665 (DS,GH,MICH,MO,NY,UC,US);3kmalEdeNuevoMorelos,González Medrano 12187 (MEXU,MO);Chancal,OcampoRd,Kenoyer & Crum 3579 (GH,MICH);�0kmalSdeLlera,Puig 5212 (ENCB,MICH).—verAcruz:Tantoyuca,Berlandier 2149(G,GH,NY);Mpio.Tempoal,ElMirador,presaPasodePiedras,2�°3�'N98°07'W,Calzada et al. 6217(ENCB,F,IEB,MEXU,XAL);Mpio.Tepetzintla,SanJoséCopaltitla,7kmalNEdeTepetzintla,Castillo C. & Benavides 2294(F,MEXU,XAL);Mpio.Ozuluama,Castillo C. et al. 270(F,IBUG,MICH,XAL,WIS);alongHwy�80betweenTampicoandPozarica,�2miNofOzuluama,38kmNofNaranjos,Croat 66092(CAS,MICH,MO);Mpio.Pánuco,�.7kmWNWofHwyMex-70,�0kmEofEbano,22°�2'N,98°�7'W,Diggs & Nee 2572(F,MICH,NY,XAL);Mpio.Chicontepec,carr.aBenitoJuárez,3kmantesdeBenitoJuárez,20°54'N,98°�3'W,Durán E. et al. 264(XAL);Wartenberg,nearTantoyuca,prov.Huasteca,Ervendberg 130(G,GH,GOET);�3miSofTampico,Hwy�80toTuxpan,Lasseigne 4897(ENCB,MEXU,MICH,MO,NY,WIS);36miNofTantoyucaonRte�20,Ozment et al. 388(WIS);roadfromTamiahuatoTuxpan,about6miSofTamiahua,Tucker 2060(DUKE,ENCB);Mpio.AltoLucero,carr.SantaAna–LosAtlixos,�9°52'N,96°32'W,Vázquez B. 1818(XAL).

Galphimia gracilis isanattractiveshrubnativetoeasternMexico.Theshootsappearreddishowing to thevesture,composedof straight towavyhairs thataresubsessileorborneonpersistenttuberclesca.0.05mmhigh,butto0.�mmhighonthepetioles.Thenarrowlyellipticaltoellipticallaminashaveanapicaltuftofhairs(sloughedoffinage)andglandsonthemarginplacedwellabovethebase.Thebrac-teolesareusuallyborneatorneartheapexofthepeduncle.


FIG.2.DistributionofGalphimia gracilisandG. montana.

The names Galphimia gracilis (and Thryallis gracilis) have been widely mis-appliedtomanyotherspeciesofGalphimiaofMexico,particularly tothosewithdeciduouspetals.Galphimia gracilisalsoisoftenconfusedwiththepartlysympatricG. glauca,whichiseasilyseparatedbyitspersistentpetalsandsmallerfruits(2.5mmlongvs.4.5–5mmlonginG. gracilis).Galphimiaglaucaalsodiffersinitsbracteoles(placedwellbelowtheapexofthepeduncle)andlackoftubercles;thelaminasvaryfromellipticaltolanceolateorovatetorarelysuborbicular,whereasthelaminasofG. gracilisareonlyellipticalornarrowlyso.Small(�9�0)includedG. graciliswithinhisverybroadlycircumscribedThryallis glauca.

Galphimia gracilis is widely cultivated in warm regions, as“goldshower” and“shower-of-gold”(andequivalentsinlocallanguages),andalsoas“thryallis.”It isfrequentlyadventiveandoccasionallylocallynaturalized.Thespecieswasintroducedtogardensas“Galphimiaglauca”(e.g.,Maund�837).Inhorticulturalpublications,inthenurserytrade,andonwebsites,G. gracilisisoftenlistedandpicturedas“Gal-phimiaglauca,”“Galphimiabrasiliensis,”“Thryallisglauca,”and“Thryallis gracilis.”SeealsothediscussionunderG. glauca.

Bartling(�840)basedG. gracilisonlivingplantsgrownatthebotanicalgardeninGöttingen,andnoauthenticmaterialisknown.Giventhemanywell-preparedcollec-tionsavailableforthisspecies,itisunfortunatethatafragmentarygardenspecimenintheJussieuherbariumwaschosenasneotype.


2. Galphimia montana(Rose)Nied.inEngler,PflanzenreichIV.�4�:60�.�928.Thry-allis montanaRose,Contr.U.S.Natl.Herb.�2:28�.�909.—type: mexico.Durango:SierraMadrenearHuazamota(fideMcVaugh,�972),�5Aug�897,Rose 2309(holotype:US!;isotypes:MEXU!NY!).

Galphimia glaucaf.parvifoliaNied.,ArbeitenBot.Inst.Königl.LyceumHosia-numBraunsberg5:25.�9�4.—type: mexico. Jalisco:slopesofthebarrancaofGuadalajara,4500ft,8Aug�902,Pringle 9698(holotype:B,destroyed;lectotype,designatedbyCuatrecasasandCroat,�98�:NY!;isotypes:F,GH!K!L!MEXU!MO!NY!US!).

Shrubortreeletto3.5m;stemsandbranchletssparselytomentulosewhenveryyoung,soonglabrous.Vestureofallvegetativepartscomposedofsessile,straightorwavytocrisped,reddishbrownhairs0.2–0.6mmlong.Laminasofthelargerleaves2–5(–5.5)cmlong,�–3cmwide,lanceolatetoovatetoorbicular,apexapiculate,baseacuteorsometimestruncate,glabrous,secondaryveinsalwaysevidentandusuallyprominulousabaxially,marginentire;petioles0.4–�.5cmlong,glabrateorglabrous;glandsusuallyapairborneonthemarginatornearthebaseofthelaminaorto�0mmabovethebaseorsometimes�orbothglandsborneonthepetiole,sometimeswithonly�glandorglandsabsent,eachgland0.4–0.6mmindiameter,disklikeandprominent, or peglike and up to 0.7 mm long; stipules �–2.5 mm long, 0.3–� mmwide, linear tonarrowlytriangular,glabrous.Inflorescencea terminalraceme, theaxessparselytomentulosetoglabrate;peduncles2–5mmlong,sparselytomentulose,pedicels5.5–�0.5mmlong,glabrateorsparselytomentulose,peduncles0.2–0.6timesaslongaspedicels;bracts�.5–3mmlong,0.4–0.5mmwide,linear,bracteoles�–�.5(–2)mmlong,0.2–0.4mmwide, linear,bractsandbracteolesglabrous;bracteolessuboppositeorto2mmapart,borneataboutthemiddleofthepeduncle,oronly�bracteoleborneatthemiddleandtheothernearthebaseorintheproximal�/4–�/3.Sepals2.4–3mmlong,�–�.4mmwide,oblong,glabrous,margindenticulate(-ciliate)inthedistal�/2;glands0–3(–5?)percalyx,0.2–0.5mmindiameter,sessileto0.3mmlong.Petalsdeciduous,unequal,glabrous,yellow,theclawssuffusedwithred,marginirregularly denticulate/fimbriate; lateral petals: claw �.2–2.2 mm long, ca. 0.5 mmwide,limb5.5–6.8(–7.5)mmlong,3.5–4.2mmwide,ellipticalornarrowlylanceolate,apexobtuse,baseattenuate;posteriorpetal:claw3–4mmlong,�–�.5mmwide,limb5–6.5mmlong,4.5–6mmwide,broadlytriangularorsometimessuborbicular,apexbroadlyrounded,basetruncateorslightlycordate.Stamensofunequallength;fila-mentoppositeanteriorsepal3–3.7mmlong,filamentsoppositeanterior-lateralpetals2.6–3(–3.5)mmlong,filamentsoppositeanterior-lateralsepals2.5–3.�(–3.5)mmlong,filamentsoppositeposterior-lateralpetals�.5–2mmlong,filamentsoppositeposterior-lateralsepals2.7–3.2(–3.5)mmlong,filamentoppositeposteriorpetal�.5–2mmlong;anthers(2.2–)2.5–3mmlong.Ovaryglabrousorpiloseonlyonthesutures;styles4–5mmlong.Coccus3–3.5mmlong,2.6–2.8mmindiameter,glabrousorpiloseonlyonthesutures;areoleca.3mmlong,ca.2.3mmwide;outercotyledon2–2.3mmlong,innercotyledon�.5–�.7mmlong.Chromosomenumberunknown.

Phenology.CollectedinflowerandfruitfromJulythroughDecember.Distribution (Fig. 2). Mexico (Colima, southern Durango, Guerrero, Jalisco,

Michoacán,Nayarit,southernSinaloa,southernZacatecas);tropicallowdeciduousforest(“selvabajacaducifolia,”“bosquetropicaldeciduo”);�50–�200m.

AdditioNAL specimeNs exAmiNed.mexico.coLimA:��–�2mibyrdSofColima,rdtoManzanillo,�9°05'N,�03°47'W,Webster & Breckon 16116 (MEXU).—Guerrero: Mpio.Ahuacuotzingo, cercadeTrapicheViejo,40kmalNEdeChilapa,Acosta & López 73 (MICH);Guayameo,Dtto.Minas,Hinton


et al. 9372 (DS,F,GH,MEXU,MICH,NY,US);Mpio.Tepecoacuilco,33kmalSdeIgualaTerraceríaaCoacoyula,Koch & Fryxell 8246 (CAS,CHAPA,ENCB,MEXU,MICH,NY);RíoBalsas,Orcutt 4183 (BM,DS,F,GH,MEXU,MO,US);hillsidesnearBalsasStation,Pringle 10063 (BM,C,CAS,F,G,GH,GOET,L,MEXU,MICH,MO,NY,S,UC,US,W).—JALisco: Mpio.SanCristóbaldelaBarranca,Km37adelantedelpueblodeSanCristóbalBarranca,Ornelas U. & Cházaro B. 767 (CHAPA,ENCB,IBUG);barrancanearGuadalajara,Palmer 97in�886(BM,G,GH,K,MEXU,MICH,MO,NY,US);betweenBolañosandGuadalajara,Rose 3050 (NY,US);Mpio.Tuxcacuesco,5–6kmWSWdeTuxcacuesco,CerrodelPalacio,�9°40'40"N,�04°0�'�0"W,Santana M. & Rosales 6494 (MICH,WIS);Mpio.Amatitán,BarrancaSantaRosa,orillaRíoSantiago,Villarreal 6658 (IBUG).—michoAcáN: Dtto.Zitácuaro,Hinton 13220 (BM,DS,ENCB,F,GH,MEXU,MICH,NY,S,UC,US);Mpio.Huetamo,MalPaso,Hinton et al. 8039 (A,BM,F).—NAyArit: Mpio.Nayar,JesúsMaría,caminoaLaMesa,cercadelarroyoElFraile,Colunga & Zizumbo 32 (CAS,MEXU);valleyoftheRíoJesúsMaríanearvillageofJesúsMaría,Feddema 1243 (MICH);Mpio.Nayar,3.9kmalNEdeJesúsMaría,caminoaHuejuquilla,22°�6'N,�04°30'W,Flores F. 1059 (MEXU,MICH,MO);CañóndeJesúsMaría,Tepic,Goldsmith 159 (F,GH,MO,UC,US);Mpio.Acaponeta,nearJesúsMaría,Norris & Taranto 13961 (IEB,MICH);Mpio.Nayar,7kmalWdeJesúsMaría,carr.alaMesadelNayar,22°�5'N,�04°38'W,Ramírez R. 442 (IEB,MICH,MEXU,MO);Mpio.Nayar,ElPuente,�2kmalNWdeJesúsMaría,carr.aLaMesadelNayar,22°�5'N,�04°33'W,Tenorio L. 16570 (MEXU,MICH,MO).—siNALoA: Mpio.San Ignacio,SanAgustín,González Ortega 526 p.p. (MEXU);rdtoMicroondaslaPalma,�.3miEofHwy�5,thisjunction26miSofHwy40junctionSofMazatlánand�.5miNofRosario,Reveal 4021 (K,MICH,MO);Mpio.Concordia,CerrodelOcote,Trejo & Dehesa 1085 (F,US).—zAcAtecAs: Mpio.Moyahua,SantaRosa,CerroLaGarruñera,Enríquez E. 136 (MEXU);Mpio.Moyahua,SanLorenzo,CerrodeLasAnonas,Enríquez E. 48 (IBUG,MEXU).

Galphimia montanaisagracefulshrubortreeletofthedeciduouswoodlandsofwesternMexico,whichshowsconsiderablevariationthroughoutitsrange.Theslen-dershootsbearthin-texturedlanceolatetoovatetoorbicularleaves;theglandsmaybeborneonthemarginofthelaminaorat/neartheapexofthepetiole.Thecalyxmaybeeglandularorhave�ormoreglands.Theovaryandfruitisusuallyglabrous,butoccasionallyeachcarpel/coccusbearsarowofhairsalongthedorsalsuture;G. montanaistheonlyspeciesinthegenusthatexhibitssuchvariabilityinovaryvesture.InitshabitG. montanaresemblesG. sessilifolia,endemictoOaxacaandnamedforthedistinctivecoriaceoussessileleaves.

CollectionsofG. montanaareoftenconfusedwithG. gracilis,whichisnativetoeasternMexicobutwidelycultivatedandoftenadventive.Bothhavedeciduouspetals,butG. gracilisdiffersinitslargerflowerslackingcalyxglands,largerglabrousfruits(4.5–5mmlong,3–3.3mmindiameter),ellipticallaminasthatbearatuftofhairsattheapex,andbracteolesusuallyborneatorneartheapexofthepeduncle.InG. montanahairsaresessile,whereasinG. gracilistheyareoftenattachedtoatinypersistenttubercle;thepetiolesareusuallyroughenedbysuchtubercles.Specimensof G. montana have also been misdetermined as G. glauca, which has persistentpetals;Niedenzu(�9�4)evendescribedacollectionofG. montanaasaformofG. glauca(f.parvifolia).TwootherspecieswithdeciduouspetalsfromwesternMexicowithwhichG. montanamightbeconfusedareG. langlasseiandG. mexiae.Bothhavedenselypiloseovariesandfruits,andlackcalyxglands.

3. Galphimia oaxacana C.Anderson,Contr.Univ.MichiganHerb.24:�5.2005.—type: mexico.Oaxaca:Dtto.Tehuantepec,RanchoRicardo,alNdeBuenosAires,entrandoporHierbaSanta,�2Sep�985,Torres C. 7306(holotype:MICH!;isotypes:F!MEXU!). Fig.3.

Shrubortreeletto6m;stemsandbranchletssparselypubescentwhenyoung,soon glabrous.Vesture of all vegetative parts of sessile, straight to wavy, reddishbrownhairs0.3–0.8mmlong.Laminasof the larger leaves3.5–6cmlong,�.5–3.5cmwide,ellipticalorovatetolanceolate,apexapiculateorsometimesacute,baseacute,glabrous,abaxiallypapilloseorsometimesonlyslightlyso(theepidermisat


FIG.3.Galphimia oaxacana.a.Floweringbranch.b,c.Proximalportionoftwoleaves,abaxialview.d.Flowerbud,opening,borneonpedicelandpeduncle,withportionof inflorescenceaxis;notecalyxglands.e.Lateralpetal.f.Posteriorpetal.g.Stamens,adaxialview,oppositeposteriorpetal(right)andposterior-lateralsepal(left).h.Gynoecium.i.Cocci,lateralview(above)andadaxialview(below).Scalebar:a,4cm;b–d,8mm;e,f,5mm;g,4mm;h,2.7mm;i,3.3mm.(Basedon:a,b,g,i,Martínez R. 55,MICH;c–f,h,Torres C. 122,MEXU.)

leastblistered),thickandcoriaceouswhendried,secondaryveins(atleastthefirsttwopairs)prominentabaxially (usuallyappearingwhite indrymaterial);marginentire, thickened;petioles0.8–�.5cmlong,glabrousorwitha fewscatteredhairs;glandsusuallyapairborneonthemarginofthelamina8–�5mmabovethebaseorsometimeswith3–6glands,eachgland0.4–0.7mmindiameter,disklikeandpromi-nent,orsometimesflushwiththemargin;stipules�.5–2.5mmlong,0.8–�mmwide,narrowlytriangular,glabrous. [Sometimesshrubswithonlytheterminalbranchesbearingaflushofsmallleavesalongshortinternodes(toca.�cmlong)andashortinflorescence,theleaves�–2.5cmlong,0.5–0.9cmwide;seediscussion.]Inflorescenceaterminalraceme,theaxessparselytomentuloseorstrigose;peduncles3.5–6.5mmlong,pedicels(7–)�0–�5mmlong,bothsparselytomentulose,peduncles0.3–0.5timesaslongaspedicels;bracts(�.5–)2–2.8mmlong,0.5–0.9mmwide,linear,bracteoles0.8–�.7mmlong,0.3–0.6mmwide,linear,bractsandbracteolesglabrous;bracteolesborneintheproximal�/5–�/2ofthepeduncle,subopposite.Sepals2.5–3mmlong,�.3–�.5mmwide,oblong,glabrous,margindenticulate-ciliateinthedistal�/4;glands�–3(–5)percalyx,0.3–0.5mmindiameter,to0.2mmlong.Petalsdeciduous,unequal,yellow,theclawssuffusedwithred,glabrous,margindenticulate;lateralpetals:claw(�.5–)2–2.5mmlong,0.5mmwide,limb(5–)6.5–8mmlong,(3.8–)4–4.5mmwide,ellipticaltonarrowlylanceolate,apexobtuse,baseacute;posteriorpetal:claw(3–)


4–5mmlong,(�–)�.2–�.5mmwide,limb(4.5–)5–6.5mmlong,(4–)5.5–6mmwide,triangular,apexbroadlyobtuse,basecordate.Stamensofunequallength;filamentopposite anterior sepal 4–4.5 mm long, filaments opposite anterior-lateral petals2.5–3mmlong,filamentsoppositeanterior-lateralsepals3.6–4.2mmlong,filamentsoppositeposterior-lateralpetals�.5–2mmlong,filamentsoppositeposterior-lateralsepals3.3–3.5mm long,filamentoppositeposteriorpetal�.5–2mm long;anthers(2.5–)2.8–3.5mmlong.Ovarypiloseonthesutures,otherwiseglabrous,hairsca.0.�mmlong;styles(4.7–)5–5.3mmlong.Coccusca.3.5mmlong,ca.2.7mmindiam-eter,piloseonthesutures,otherwiseglabrous;areoleca.3mmlong,ca.2.5mmwide;matureseednotseen.Chromosomenumberunknown.

Phenology.CollectedinflowerfromAugusttoOctober,infruitfromSeptembertoDecember.

Distribution(Fig.�0).Mexico(Oaxaca,PacificslopesoftheIsthmusofTehu-antepec);indeciduousforest(“selvabajacaducifolia”)andtransitiontopine-oakforest;500–��00m.

AdditioNAL specimeNs exAmiNed.mexico.oAxAcA:alrededoresdelCerroGuiengola,a�0kmaprox.alNWdeTehuantepec,Cabrera 7413(MEXU,MO);Dtto.SantoDomingoTehuantepec,Mpio.Mixte-quilla,a�8kmdeMixtequilla,carreteraaPasoEscondido,�6°27'N,95°�9'W,Calzada 19255(MEXU);Mpio.SantiagoLaollaga, recorridoporelaguajeCoyol,alWdeLaollaga,�6°34'N,95°�4'W,Campos V. 4039(F,MEXU,MO);Mpio.SantiagoLaollaga,recorridohaciayporelArroyodeHierbaSanta,alEdeGuichixu,brechaLaollaga–GueveadeHumboldt,�6°4�'N,95°�6'W,Campos V. 4129 (F,MEXU,MO);Dtto.Tehuantepec,Mpio.BuenosAires,BuenosAires,rumboaElCerroArenal,Martínez R. 55(IEB,MICH,MO);Dtto.Tehuantepec, ruinasdelCerroGuiengola,Torres C. 40 (MEXU,MO);Dtto.Tehuantepec,“LasPalmitas,”laderaorientedelCerroGuiengola,Torres C. 122(IEB,MEXU,MO);Dtto.Tehuantepec,��kmalWdelaChiviza,haciaLachiguiri,Torres C. 5717(MEXU,MICH,XAL);Dtto.Tehuantepec,�2.2kmalWdelaChiviza,haciaLachiguiri,Torres C. 5720(F,MEXU);Dtto.Tehuantepec,��.3kmalNdeLaChiviza,Torres C. & Martínez 5880(MEXU,MICH).

Galphimia oaxacanaisknownonlyfromdryhabitatsonthePacificsideoftheIsthmusofTehuantepec.Theleavesarecoriaceousandhaveathickenedmargin.Theabaxialepidermisisgenerallypapillose;thecellsareatleastslightlyraisedtogiveablisteredaspect.Thecostaandmajorsecondaryveinsareabaxiallyveryprominentandappearwhiteindriedspecimens.ThecollectionsTorres C. 122, Torres C. 5717, andTorres C. & Martínez 5880 areunusualinthattheyconsistofterminalbranchesbearingaflushofverysmallleavesalongshortinternodes(toca.�cmlong)andashortinflorescence;theleavesare�–2.5cmlong,0.5–0.9cmwide.Exceptforsize,thesecollectionsmatch G. oaxacanainallaspects,andthisgrowthformmayreflectparticularlydrygrowingconditions.TheonlyotherspeciesfoundintheIsthmusofTehuantepecisthecoastalG. arenicola,whichhasmembranousleaves,lackscalyxglands,andhasaglabrousovaryandfruit;itdiffersmostnotablyinthepresenceofelongateepidermalprocessesonthelaminarmarginandhairsborneonpersistenttubercles.

4. Galphimia hirsutaCav.,Icon.5:62.May�799.Malpighia hirsuta(Cav.)Persoon,Syn.pl.�:506.�805[combinationalsoproposedbyPoiretinLamarck,Ency-cl.,Suppl.4:6.�8�6].Galphimiahirsutaf.rufaK.Koch,BerlinerAllg.Gar-tenzeitung�857(50):394.�857,nom.superfl.Thryallis hirsuta(Cav.)Kuntze,Rev.gen.pl.�:89.�89�.—type:mexico. Guerrero:interChilpancingoetRíoAzul,Neé s.n.(holotype:MA-475693,microfiche:MICH!,photo:MO!;pos-sibleisotypes:BM!MA-475694,microfiche:MICH!).


Shrubto�m;stemspubescentwhenyoung,becomingglabrousbutroughenedbytubercles.Vestureofallvegetativepartsofstraightorcrisped,reddishbrownhairs0.5–�.8mmlong,eachhairborneonapersistenttubercleupto0.�mmhigh.Laminasofthelargerleaves5–7.5cmlong,2–3cmwide,lanceolatetoellipticaltorhombic,apex apiculate, base cuneate-decurrent, pubescent, secondary veins prominulousabaxially,marginentire,evenlypubescentonbothsurfaces;petioles0.2–�cmlong,pubescent, inolderleavessomeorallhairsabradedandonlythetuberclespersis-tent;glandsapairborneonthemarginatornearthebaseofthelamina,eachgland0.2–0.4mm indiameter,peglike,0.4–0.6mm long; stipules2.5–3mm long,0.5–0.7mmwide,linear,abaxiallywithscatteredhairs.Inflorescenceaterminalraceme,theaxespubescent;peduncles5–7.3mmlong,pedicels4.5–6.2mmlong,bothpubescent,peduncles0.9–�.3timesaslongaspedicels;bracts2.2–3mmlong,0.5–0.7mmwide,linear,bracteoles�.4–�.6mmlong,0.4mmwide, linear,bractsandbracteoleswithscatteredhairsabaxiallyandalongthemargin;bracteolesborneataboutthemiddleofthepeduncle,suboppositeorupto�mmapart.Sepalsca.2.5mmlong,ca.�mmwide, oblong, pubescent, margin ciliate; glands 5 per calyx, 0.4–0.5 in diameter, to0.3mmlong.Petalsdeciduous,unequal,yellow,withscatteredhairsontheclawandmidveinabaxially,marginfinelydenticulate;lateralpetals:claw2–2.2mmlong,0.4–0.5mmwide,limb6.5–7mmlong,4–4.5mmwide,narrowlytriangular,apexobtuse,basecordate;posteriorpetal:clawca.3.5mmlong,ca.�.3mmwide,limb5.5–6mmlong,ca.6mmwide,broadlytriangular,apexobtuse,basetruncate.Stamensofunequallength;filamentoppositeanteriorsepalca.3.8mmlong,filamentsoppositeanterior-lateralpetalsca.3.6mmlong,filamentsoppositeanterior-lateralsepalsca.3.2mmlong,fila-mentsoppositeposterior-lateralpetalsca.2.2mmlong,filamentsoppositeposterior-lateralsepalsca.3mmlong,filamentoppositeposteriorpetalca.2.5mmlong;anthers3.3–3.5mmlong.Ovaryglabrous;stylesca.5.5mmlong.Maturefruitnotseen.

Phenology.CollectedinflowerandimmaturefruitinJulyandSeptember.Distribution(Fig.2).Mexico(Guerrero);pine-oakforest;�00–�000m.

AdditioNAL specimeNs exAmiNed.mexico.Guerrero:Dtto.Galeana,Atoyac,Hinton 14533(GH,MICH,NY,US);ca.20miNofSanLuisdelaLoma,Rowell 3144(MICH).

Galphimia hirsutadiffersfromallbutoneotherspeciesofGalphimiainthatitispubescentinnearlyallitsparts(onlytheandroeciumandgynoeciumareglabrous).Thevestureiscomposedofhairsborneontubercles,whichpersistandgivearough-enedaspecttoolderparts.Thelaminasvaryfromlanceolatetoellipticaltorhombicandbearapairofpeglikeglandsonthemarginatornearthecuneate-decurrentbase.TheonlyotherspeciesinthegenusthatisentirelypubescentisG. vestitaofnorthernMexico,but thehairsarenotborneontubercles; italsodiffers fromG. hirsuta in its linear laminas, lackofpeduncles,muchsmaller subequalpetals,andpiloseovariesandfruits.Galphimia hirsutamightbeconfusedwiththemorewidelydistributedandperhapssympatricG. tuberculata,whichisalsocharacterizedbyper-sistenttuberclesandcalyxglands,butthatspeciesiseasilyseparatedbyitsessentiallyglabrousleavesandcalyx,andthepubescentovaryandfruit.

ThelikelyisotypeatBMconsistsofafloweringshootandbearsthenote“misitamicis.Zea”;thespecimenapparentlybelongedtotheRoemerherbarium,whichcametotheBMholdingsaspartoftheShuttleworthherbarium.

EarlierstudentsofGalphimia(e.g.,Bartling�840;Jussieu�843;Rose�909;Nie-denzu�9�4,�928)misappliedthenameG. hirsutatospecimensofG. elegans,aOaxa-canendemicwithpersistentpetalsandabaxiallysparselypubescentlaminas,whichhadbeenintroducedtobotanicalgardensinthe�800s;seethatspecies(no.20).

2007 C.ANDERSON:GALPHIMIA 2�

5. Galphimia tuberculata(Rose)Nied. inEngler,PflanzenreichIV,�4�:602.�928.Thryallis tuberculata Rose, Contr. U.S. Natl. Herb. �2: 28�. �909.—type: mexico. Sinaloa: between Rosario and Colomas, �2 Jul �897, Rose 1607(holotype:US!,photos:A!F!NY!;isotypes:NY-fragment!GH!).

Thryallis humilisRose,Contr.U.S.Natl.Herb.�2:280.�909.Galphimia humilis(Rose) Nied. in Engler, Pflanzenreich IV, �4�: 60�. �928.—type: mexico.Nayarit:roadbetweenConcepciónandAcaponeta,TerritoriodeTepic,29Jul�897,Rose 1907(holotype:US!,photos:A!F!NY!;isotype:NY!).

Fig.4.

Shruborsubshrubto2m;stemstuberculate-strigosewhenyoung,inageonlythetuberclesremaining.Vestureofallvegetativepartsofwavytocrisped,reddishbrownhairs0.3–�.9mmlong,borneonapersistenttubercleupto0.2(–0.5)mmhigh.Laminasofthelargerleaves3.8–7.5cmlong,�.4–3cmwide,ellipticalorlanceolateorrhombic,apexacuteorapiculateorsometimesobtuse-apiculate,basecuneate-decurrent,glabrousorwithscatteredhairsonthemidribabaxially,inolderleavesthemidribandalsothemarginsnearthebasewithtuberclesonly,secondaryveinsprominentorprominulousabaxially;petioles0.2–�.2cmlong,tuberculate-strigoseoronlythetuberclesremaining;glandsborneonthemargin0.5–2mmabovethebaseofthelaminaorrarelyatthebase,usuallyapairorsometimesonly�gland,eachgland0.2–0.5mmindiameter,peglike,(0.2–)0.5–�.2(–�.6)mmlong,ortheglandsabsent;stipules(�.5–)2–3.6mmlong,0.5–�.2mmwide, linear,glabrousorwithscatteredhairsalongthemargin.Inflorescenceaterminalraceme,theaxestuberculate-strigoselikethestems;peduncles4.5–��mmlong,pedicels2.5–7mmlong,bothtuberculate-strigose,peduncles�–2.2timesaslongaspedicels;bracts2.5–4.5mmlong,0.5–0.8(–�)mmwide,linear,bracteoles�.�–2.6mmlong,0.3–0.5mmwide,linear,bractsandbracteolesglabrousorwithscatteredhairsattheapexandalongthemargin;bracte-olesborneintheproximal�/3–�/2ofthepeduncle,suboppositeorupto3mmapart.Sepals3–3.7mmlong,�.4–�.7mmwide,oblongtonarrowlyelliptical,glabrous,theapexdenticulate(-ciliate)orentire;glands0–5percalyx,0.2–0.4mmindiameter,usu-ally0.2–0.3mmlongorrarelyonaspurupto�.2mmlong.Petalsdeciduous,unequal,yelloworyellowmarkedwithred,glabrous,marginsubentireorfinelydenticulate;lateralpetals:claw2.2–3mmlong,0.5–0.7mmwide,limb6–7.3mmlong,4.5–6mmwide,narrowlytriangular(-ovate),apexobtuse,basetruncateorcordate;posteriorpetal:claw3.4–4.5mmlong,�.4–2mmwide, limb6–6.5mmlong,6–7.2mmwide,broadlytriangular,apexobtuse,basetruncatetoslightlycordate.Stamensofunequallength;filamentoppositeanteriorsepal(2.5–)3.2–4.5mmlong,filamentsoppositeanterior-lateralpetals3–4mmlong,filamentsoppositeanterior-lateralsepals3–4mmlong,filamentsoppositeposterior-lateralpetals�.8–2.2mmlong,filamentsoppositeposterior-lateralsepals3–3.5mmlong,filamentoppositeposteriorpetal(�.6–)�.8–2.2mmlong;anthers(2.5–)3–3.5mmlong.Ovarypilose,inthedistal�/3onlyonthesutures,hairsca.0.�mmlong;styles(5.2–)5.5–6.2mmlong.Coccus3.5–4.5mmhigh,ca.3mmindiameter,pilose,inthedistal�/3onlyonthesutures;areoleca.4mmlong,ca.3mmwide;matureseednotseen.Chromosomenumberunknown.

Phenology.CollectedinflowerinMarchandfromMaythroughSeptember,andinfruitinMarch,May,July,andAugust.

Distribution (Fig. 5). Mexico (Colima, Guerrero, Jalisco, Nayarit, Sinaloa);deciduousforest,matorral,andinsecondaryvegetation;sealevelto400m.

represeNtAtive specimeNs.mexico.coLimA:coastallowlandsca.�5miSEofTecomán,rdtoCoa-huayana,Mich.,ca.2mifromRíoCoahuayana,McVaugh 16070(MEXU,MICH).—Guerrero:Tecpan,


FIG.4.Galphimia tuberculata.a.Floweringbranch.b.Baseoflamina,abaxialview.c.Branchenlargedtoshowtubercles.d.Detachedlargeleaf,adaxialview.e.Baseoflaminain“d,”adaxialview.f.Flowerwithsubtendingpedicelandpeduncleandportionofinflorescenceaxis.g.Lateralpetal.h.Posteriorpetal.i.Androeciumlaidout,abaxialview,thestamenatrightoppositeposteriorpetal.j.Gynoecium.k.Cocci,adaxialview(left)andlateralview(right).l.Seed.Scalebar:a,d,4cm;b,c,e,i–l,4mm;f–h,8mm.(Basedon:a–c,f–j,McVaugh 16070,MICH;d,e,Ortega 6757,F;k,l,Lott 3741,MICH.)

torredemicroondascercadePapanoa,54kmalNWdeTecpan,Koch & Fryxell 82221(CHAPA,ENCB,MEXU,MICH,NY);LaUnión,Langlassé 255(F,G,GH,P,US).—JALisco:PuntaFarallón,Bullock 2053(CAS);Mpio.LaHuerta,caminoantiguosur,Est.BiologíaChamela,�9°30'N,�05°03'W,Lott 1739(CAS,MICH,TEX,WIS);Mpio.LaHuerta,RanchoCuixmala,Cumbres�,rdthroughdeforestedarea,�9°3�'N,�04°56'W,Lott 3741 (CAS,MICH).—NAyArit:Mpio.Tepic,ViveroForestalElCorte,Benítez-Paredes 3166(MEXU);Mpio.Ruiz,�–3kmWofElVenado,rdfromRuiztoJesúsMaría,Breedlove 45278(CAS,MICH);Mpio.Huajícori,8kmalNdeHuajícori,carr.aQuiviquinte,22°43'N,�05°�9'W,Ramírez 656(IEB,MICH,MEXU,MO);Mpio.Acaponeta,Rose 1450(NY,US).—SiNALoA:Mpio.SanAgustín,González Ortega 4001(MEXU,US);Mazatlán,González Ortega 7271(CAS,F,G,K,MEXU,US).


FIG.5.DistributionofGalphimia elegans, G. mirandae, andG. tuberculata.

Galphimia tuberculata isnamed for thepersistent tubercles that roughen theshootsandleavesafterthehairstheybearareshed.SuchtuberclesarealsofoundinG. brasiliensis(easternBrazil),G. gracilis(easternMexico),andG. hirsuta(Guer-rero),buttheyaremostpronouncedinG.tuberculataandtheOaxacanG. arenicola,bothspeciesofPacificcoastal lowlands.Galphimia arenicoladiffers in its smallerflowers,eglandularcalyx,glabrousovaryandfruit,andtheunusualmulticellularpro-cessesthatarefoundalongthemarginintheproximalpartofthelamina.Galphimia hirsutaisknownonlyfromthelowlandsofGuerreroand,likeG.tuberculata,hasaglandularcalyx.Itisnamedfortheabundantvesturethatcoversnearlyallpartsoftheplant,whereasinG.tuberculataonlytheyoungestpartsandinflorescenceaxesarepubescent.

6. Galphimia arenicolaC.Anderson,Syst.Bot.28:2�7.2003.—type: mexico.Oaxaca:Mpio.Tehuantepec, de Rincón Bamba a Garrapatero, Rincón Bamba seencuentraa44kmalWdeSalinaCruz,carr.aPochutla,�5°59'N,95°27'W,3�Aug�988,Martínez 1817(holotype:MO!;isotypes:IEB!MEXU!).

Fig.6.

Shrubto4m;stemspubescentwhenyoung,becomingglabrousbutroughenedbytubercles.Vestureofallvegetativepartsofstraighttowavy,reddishbrownhairs,0.5–�mmlong,eachhairborneonapersistenttubercleupto0.�mmhigh.Laminas


ofthelargerleaves2.5–4.8cmlong,�–2cmwide,narrowlylanceolatetonarrowlyelliptical,apexacute,baseacuteorslightlydecurrent,adaxiallyglabrous,abaxiallywithscatteredhairs(especiallyonthecosta)whenyoungbutsoonglabrousbutwithpersistenttubercles,secondaryveinsonlyfaintlyevidentabaxially,marginwithir-regularlyspaced,elongate,multicellular,epidermalprocesses,similartothetuberclesbutatleasttwiceaslong;petioles0.4–�cmlong,verysparselystrigosetoglabrous,roughenedbypersistenttubercles;glandsapair,borneonthemargin2–5mmabovethebaseofthelamina,rarelywith4or5glands,eachgland0.4–0.5mmindiameter,disklikeandprominent,orpeglikeandupto0.8mmlong,stipules�.5–2.5mmlong,0.5–0.7mmwide,narrowlytriangulartosubulate,abaxiallywithscatteredhairsandthemarginciliate.Inflorescenceaterminalraceme,theaxestuberculateandstrigose,peduncles4–7.8mmlong,pedicels2–5mmlong,bothtomentulose,peduncles�.2–3timesaslongaspedicels;bracts2.3–3mmlong,0.5–0.6mmwide,linear,bracteoles�.3–�.7mmlong,0.4mmwide,linear,bractsandbracteolesglabrousorwithscatteredhairsalong themargin;bracteolesborne in theproximal�/4–�/2of thepeduncle,suboppositeorupto�mmapart.Sepals2.5-2.8mmlong,�.4–�.6mmwide,elliptical,glabrousorwithscatteredhairsnearthebase,margindenticulate-eroseinthedistal

FIG.6.Galphimia arenicola.a.Floweringbranch.b.Leaf.c.Baseoflamina,showingapairofmar-ginalleafglandsandmarginalprocesses.d.Fruitborneonpedicelandpeduncle,withportionofinflores-cenceaxis.e.Lateralpetal.f.Posteriorpetal.g.Stamens,adaxialview,oppositeposteriorpetal(right)andposterior-lateralsepal(left).h.Gynoecium.i.Coccus,adaxialview.j.Embryo.Scalebar:a,4cm;b,2cm;c,d,8mm;e,f,5.7mm;g–i,4mm;j,2.7mm.(Basedon:a–c,Martínez 1817,MO;d–h,Castillo C. et al. 9642,MEXU;i,j,Castillo C. et al. 9498,XAL.)


�/2;glandsabsent.Petalsdeciduous,unequal,yellowandsuffusedwithred,glabrous,margindenticulate;lateralpetals:claw2–2.3mmlong,0.5–0.6mmwide,limb5–5.5mmlong,3.3–4mmwide,ovatetoelliptical,apexobtuse,basetruncateorslightlycordate;posteriorpetal:clawca.3.5mmlong,�.6mmwide,limbca.4.5mmlong,ca.4.5–5mmwide,triangular,apexbroadlyobtuse,basecordatetoauriculate.Stamensofunequallength;filamentoppositeanteriorsepal3–3.5mmlong,filamentsoppositeanterior-lateralpetals2.5–3mmlong,filamentsoppositeanterior-lateralsepals3–3.�mmlong,filamentsoppositeposterior-lateralpetals�.5–�.8mmlong,filamentsoppo-siteposterior-lateralsepals2.5–2.6mmlong,filamentoppositeposteriorpetal�.6–2mmlong;anthers2.4–2.8mmlong.Ovaryglabrous;stylesca.3.6mmlong.Coccusca.5mmhigh,ca.4mmindiameter,glabrous;areole4–4.8mmlong,3.3–4mmwide;outercotyledon2.7–2.8mmlong, innercotyledon2.3–2.5mmlong.Chromosomenumberunknown.

Phenology.CollectedinflowerandfruitinAugustandSeptember.Distribution (Fig. 5). Mexico (Oaxaca); on sandy soils, in dunes and coastal

deciduousforest;sealevelto70m.

AdditioNAL specimeNs exAmiNed.mexico.oAxAcA:Mpio.Huatulco,5kmde lacarr.nac.por labrechaalasplayasdeCacaluta,�5°45'�0"N,96°�0'�0"W,Castillo C. et al. 9498 (XAL);Mpio.Huatulco,PlayadeSanAgustín,�5°40'30"N,96°�4'20"W,Castillo C. et al. 9642 (MEXU,XAL);Dtto.Tehuantepec,Mpio.SantiagoAstata,LasPeñas,500malsuroestedelalagunaqueencuentraa2kmalsurdeBarradelaCruz,�5°49'25.4"N,95°57'58.5"W,Elorsa C. 5030(MICH);PlayaCoyote,Liebmann 8675(C);SanAgustín,Liebmann 8676, 8682(C);Dtto.Tehuantepec,Mpio.SanPedroHuamelula,CerroPiedradelAire,200malsurestedelRanchoElParaíso,�5°52'7.8"N,95°50'�4.3"W,Salas M. 4122(MICH).

Galphimia arenicolaisfoundondunesandindeciduousforestalongthecoastofOaxaca.Itisuniqueinthegenusinthatthemarginsofthelaminasbearelongate,multicellular,epidermalprocesses.LikeG. tuberculata,alowlandspeciesfoundfromSinaloatoGuerrero,G. arenicolaismarkedbypersistenttuberclesonitsshootsandleaves,butdiffersinitssmallerflowerswithaneglandularcalyxandaglabrousovary.Galphimia oaxacana,whichisknownonlyfromtheIsthmusofTehuantepec,lackstubercles;ithasapiloseovaryand(usually)calyxglands.

7. Galphimia glandulosaCav.,AnalesHist.Nat.�:37.Oct�799,nonGalphimia glan-dulosaRose,�897.Malpighia glandulosa(Cav.)Persoon,Syn.pl.�:506.�805,nonMalpighia glandulosaCav.,�789.Malpighia biglandulosaPoir.inLam.,Encycl.,Suppl.4:7.�8�6.Galphimia glandulosavar.lanceolataDC.,Prodr.�:582.�824,nom.superfl.Thryallis glandulosa(Cav.)Kuntze,Rev.gen.pl.�:89.�89�.—type: mexico.Guerrero:“Acapulco,Acambaro,&c.,”Née s.n.(holotype: MA-29470, excluding the inflorescence mounted in middle ofsheetandthebranchletatlowerright,whichareG. paniculataBartl.;micro-fiche:MICH!,photos:F!MICH!MO!)

Galphimia glandulosaRose,Contr.U.S.Natl.Herb.5:�37.�897,nonGalphimia glandulosaCav.,�799.Thryallis palmeriRose,Contr.U.S.Natl.Herb.�2:28�.�909.Galphimia paniculatavar.glandulosaNied.inEngler,PflanzenreichIV,�4�:599.�928.—type: mexico.Guerrero:nearAcapulco, Palmer 474in�895(holotype:US!,photos:F!NY!;isotype:NY-fragment!).

Thryallis dasycarpaSmall,N.Amer.Fl.25:�5�.�9�0.—type: mexico.Sinaloa:Rosario,Jan�895,Lamb 470(holotype:NY!;isotypes:DS!F,G!GH!NY-fragment!UC-fragment!US!).

Fig.7


FIG. 7. Galphimia glandulosa. a. Flowering branch. b. Large leaf. c. Base of petiole with glands.d.Oldflowerwithmaturingfruit,borneonpedicelandpedunclewithportionofinflorescenceaxis;notecalyxglands.e.Lateralpetal.f.Posteriorpetal.g.Stamens,adaxialview,oppositeposteriorpetal(left)andposterior-lateralsepal(right).h.Gynoecium.i.Cocci,adaxialview(above)andlateralview(below).j.Embryo.Scalebar:a,b,4cm;c,8mm;d–h,5mm;i,j,4mm.(Basedon:a,d,González Ortega 5141,US;b,c,Pérez 1562,MEXU;e–h,McVaugh 22903,MICH;i,j,Anderson & Anderson 5849,MICH.

Shruborsuffrutescentherbto4m;floweringshootsusuallyleaflessorbearingonlyyoungleaves,sometimeswithmatureleavespresent.Vestureabsentexceptforpiloseovaryandfruit.Laminasofthelargerleaves4.5–�4cmlong,2–6cmwide,lan-ceolatetoellipticaltoovatetorhombic,apexapiculate,basecuneate,secondaryveinsprominulousabaxially;petioles�–4cmlong;glandsapairusuallyborneonthepeti-oleinthebasal�/6–�/2,oppositeorupto4mmapart,rarelyalsoatthebaseofthebladeorabsent,eachgland0.8–�.5mmindiameter,disklikeandprominent;stipules�.2–�.8mmlong,0.7–0.8mmwide,linearorverynarrowlytriangular.Inflorescenceaterminalracemeorapanicle,theaxeslaxandcommonlywhip-like;peduncles2.5–6.5(–�0.5)mmlong,pedicels6.2–�2.3mmlong,peduncles0.3–0.7(–�)timesaslongaspedicels;bracts�.7–3.2mmlong,0.4–0.7mmwide,linear,bracteoles�.2–�.7(–2)mm


long,0.3–0.5mmwide,linear;bracteolesoppositeorsubopposite,borneintheproxi-mal�/2ofthepeduncletonearitsbase.Sepals(2–)3–3.5mmlong,�.2–�.7mmwide,lineartooblong,marginentireorfinelydenticulateinthedistal�/2;glandsusually5percalyx,sometimesfewerornone,0.3–0.5(–0.7)mmindiameter,prominentorpeglikeandupto0.5mmlong,eachborneatthebaseofthesinusbetweenadjacentsepals.Petalsdeciduous,unequal,yellowbutusuallysuffusedwithred,marginden-ticulate;lateralpetals:claw�.3–2mmlong,0.5–0.6mmwide,limb6–7.5mmlong,3.5–4.5mmwide,elliptical,apexobtuse,baseattenuate;posteriorpetal:claw2.5–3.3mmlong,�.5–2mmwide,limb5.5–7mmlong,4–5(–5.8)mmwide,ovate-triangular,apexbroadlyrounded,basegraduallyattenuate.Stamensofunequallength;filamentoppositeanteriorsepal3–3.2mmlong,filamentsoppositeanterior-lateralpetals2–2.5mmlong,filamentsoppositeanterior-lateralsepals2.7–3mmlong,filamentsoppositeposterior-lateralpetals�.5–�.6mmlong,filamentsoppositeposterior-lateralsepals2.8–3.5mmlong,filamentoppositeposteriorpetal�.5–�.9mmlong;anthers3.3–4.4mmlong.Ovarypilosethroughoutorsometimesthedistal�/3glabrous,hairsto0.�mmlong;styles(3.7–)4–5.5mmlong.Coccus4.5–5mmhigh,3.2–3.5mmindiameter,pilosethroughoutorsometimesthedistal�/3glabrous;areole4.5–5mmlong,ca.3mmwide;outercotyledonca.3mmlong,innercotyledonca.2.2mmlong.Chromo-somenumberunknown.

Phenology.CollectedinflowerandfruitfromDecembertoMay,onecollectioninflowerinOctober(González M. 4010).

Distribution(Fig.8).Mexico(Guerrero,Jalisco,México,Michoacán,Sinaloa;onecollectionfromVeracruz);inoakforestanddeciduousforest;20–��50(–2450)m.

FIG.8.DistributionofGalphimia glandulosaandG. sessilifolia.


represeNtAtive specimeNs.mexico. Guerrero:Temisco,Bruff 1308(MEXU);trailEofTemisco,NofRíoBalsas,Dtto.Adama,Mexia 8948a(UC);arribaIcacosAcapulco,Miranda 4346(MEXU);Achotla,Reko 4919(US).—JALisco:Mpio.LaHuerta,ArroyoChamela,0.3kmEofHwy200(BarradeNavidad–PuertoVallarta),Km63.5,environsofEst.Biol.Chamela(UNAM),Bullock 1114(MICH);steephills3–�5kmbyrdSofJilotlán,McVaugh 22903(ENCB,MICH);Mpio.LaHuerta,Est.Biol.Chamela,Pérez J. 1562(MEXU);Mpio.LaHuerta,Chamela,rdalongRíoChamelaneargroundsofthe“InstitutodeBiología”ofUNAM,�9°32'N,�05°05'W,van Rooden 726(MEXU,MICH).—méxico:ElPalmarChico,Tejupilco,González M. 4010 (MEXU);piedelaloma,Tejupilco,González M. et al. 5538(MEXU,XAL);Platanal,Temascaltepec,Hinton et al. 5749(A,F,G,NY);Mpio.Temascaltepec,Nanchititla,Hinton et al. 7354(BM,ENCB,F,GH,LL,MEXU,NY,S,US).—michoAcáN:20kmbyrdSofUruapanonrdtoCuatroCaminos,Anderson & Anderson 5849(MICH);rdtoApatzingán,ca.2kmNAguililla,Burch 2016(MO);Mpio.Pómaro,alNdeMaruata,caminoaPómaro,Guevara F. 1620(CHAPA,ENCB,IBUG);Mpio.Tiquicheo,entreHuetamoyTiquicheo,Medrano 4124(MEXU).—siNALoA:Mpio.SanIgnacio,Balboa,SanJavier,González Ortega 5138(MEXU,US);Balboa,González Ortega 5141(US);Mpio.Mazatlán,Guaracha,ElQuelite(?),González Ortega 5419(ENCB,MEXU,US);Mpio.Mazatlán,Mazatlán,González Ortega 5631(ENCB,K,MEXU,US);vic.ofCuliacán,González Ortega 6693(US);3�miNofMazatlánoncoasthwy,Kimnach 1918(CAS,MEXU);vicinityofRosario,Rose 14567(A,C,F,GH,MO,NY,P,S,US);vicinityofGuadalupe,Rose 14776(GH,NY,US).—verAcruz:Córdoba,Reko 5133(US).

Galphimia glandulosaisreadilyseparatedfromallotherspeciesofbyitslackofvesture(exceptforthepiloseovaryandfruit),itspetioleswithapairofglandsintheproximal�/6–�/2,andthewhip-likeinflorescences.Theflowershavea(commonly)glandular calyx and petals mostly entirely suffused with red.This species flowersbeforeorwhiletheleavesdevelopandthusisrarelycollectedwithmatureleaves.ThepartlysympatricG. floribundaisalsocommonlyglabrousinallitsparts,butithasaneglandularcalyx,persistentyellowpetals,andaglabrousovaryandfruit,andbearstheleafglandsatthebaseofthelamina.TherangeofG. glandulosaextendsfromcentralSinaloatosouthernMéxicoandGuerrero;therecordfromVeracruzissurprisingandthelocalityperhapsofdoubtfulaccuracy.

SeveralstudentsofGalphimiawereuncertainabouttheapplicationofthenameG. glandulosaCav.,andthusthisspecieshasbeenknownunderseveralsynonyms.Rosedescribeditin�897asanoveltywiththesameepithet.Later(�909)heacceptedKuntze’s(�89�)transferofallspeciesofGalphimiatoThryallisL.,andprovidedthenomennovumThryallis palmeriforG. glandulosaRose,becausethecombinationThryallis glandulosa(Cav.)Kuntzehadalreadybeenpublished;RoselistedThryallis glandulosa(Cav.)Kuntzeamongthe“uncertainspecies.”Small(�9�0)acceptedT. palmeri,fromwhichheexcludedG. glandulosaCav.(butdidnotaccommodatetheCavanillesnameelsewhereinhistreatment),andalsopublishedThryallis dasycarpa,whichhedifferentiatedbyitseglandularcalyxandsmallerleaves.ThetypecollectionofThryallis dasycarpaconsistsoffloweringbranchletsandonlythesmallerleavesassociatedwiththeinflorescences;thecalycesoftheholotypeareindeedeglandularbutafewintheisotypesdohaveaglandortwo.

8. Galphimia langlassei(S.F.Blake)C.Anderson,Syst.Bot.28:7�5.2003.Thryallis langlasseiS.F.Blake,Contr.GrayHerb.52:7�.�9�7.—type: mexico.Guer-rero(fideMcVaugh,�95�):SanAndrés,700m,22Mar�899,Langlassé 955(holotype:GH!;isotypes:F,G!K!P!US!;photoofPisotype:MICH!).

Fig.9.

Shruborsubshrubto3m;stemssparselytomentulosewhenyoung,soongla-brous.Vestureofallvegetativepartscomposedofsessile,straightorwavytocrisped,reddishbrownhairs0.2–0.8mmlong.Laminasof the larger leaves5–�4cmlong,�.5–4.8cmwide,ellipticalornarrowlysotolanceolate,apexacuteorapiculate,base


FIG.9.Galphimia langlassei.a.Leaf.b.Baseoflamina,showingpairofmarginalglands.c.Flowerbudborneonpedicelandpeduncle,withportionofinflorescenceaxis.d.Lateralpetal.e.Posteriorpetal.f.Stamens,abaxialview,oppositeposteriorpetal(right)andposterior-lateralsepal(left).g.Gynoecium.h.Cocci,lateralview(left)andadaxialview(right).Scalebar:a,4cm;b,f–h,4mm;c,�cm;d,e,5mm.(Basedon:a,b,Cuevas & Rosales 1828,IEB;c,Daniel & Bartholomew 4849,MICH;d,e,Cochrane et al. 11687,WIS;f,g,Santana M. et al. 4807,WIS;h,Pepita 86,ZEA.)

attenuate and somewhat decurrent, glabrous or sometimes with a few scatteredhairsabaxiallyonthemidrib,costaandsecondaryveinsprominentabaxially,marginentire;petioles�–2.5cmlong,glabrousorwithafewscatteredhairs;glandsusuallyapair(rarelyonly�gland)borneonthemarginofthelamina0.�–�cmabovethebase,eachgland0.5–�.�mmindiameter,disklikeandprominent,orpeglikeandupto�.2mmlong;stipules�.2–2.8(–3.5)mmlong,0.4–�mmwide,linearornarrowlytriangular,margindistallyciliate.Inflorescenceaterminalracemeorpanicle,theaxestomentuloseorsparselyso;peduncles4.5–�2.5mmlong,pedicels4.5–�3mmlong,bothtomentulose,peduncles0.6–�.2(–�.6)timesaslongaspedicels;bracts�.3–2.2(–3.4) mm long, 0.5–0.8 mm wide, linear, bracteoles �–2 (–2.5) mm long, 0.3–0.5(–0.7)mmwide,linear,bractsandbracteolesglabrousorwithafewscatteredhairsalongthemargin;bracteolessuboppositeorupto2mmapart,borneinthedistal�/4–�/2ofthepeduncle.Sepals2.7–3.5mmlong,�.2–2mmwide,oblongtoelliptical,glabrous,margindistallydenticulate-ciliate;glandsusuallyabsent[inHinton et al. 11683somecalyceswith�–3glands,eachgland0.3–0.5mmindiameter,prominent].Petalsdeciduous,unequal,yellow,oftensuffusedwithred,especiallyalongtheclawandmidribofthelimb,glabrous;lateralpetals:claw2.3–2.8(–3)mmlong,0.5–0.6mmwide,limb6–7mmlong,4.5–5.5mmwide,ovate,apexrounded,basetruncateorsubacute;posteriorpetal:claw3.5–4mmlong,�.3–�.5mmwide,limb6.5–7.7(–8)mmlong,6–7.7mmwide,broadlytriangular,apexbroadlyrounded,basetruncatetocordate.Stamensofunequallength;filamentoppositeanteriorsepal3.5–4.5mmlong,filamentsoppositeanterior-lateralpetals2.8–3.5(–3.7)mmlong,filamentsoppositeanterior-lateralsepals3–4(–4.5)mmlong,filamentsoppositeposterior-lateralpetals�.6–2.5(–2.8)mmlong,filamentsoppositeposterior-lateralsepals3.5–4.6mmlong,


filamentoppositeposteriorpetal�.6–2.6mmlong;anthers(2.8–)3–3.6(–4)mmlong.Ovarypilose,hairsto0.3mmlong;styles5–6.5(–6.8)mmlong.Coccus4–4.5mmhigh,3–3.8mmindiameter,pilosetoglabrescentwhenmature;areole3.5–4.2mmlong,2.5–3.8mmwide;matureembryonotseen.Chromosomenumberunknown.

Phenology.CollectedinflowerfromNovemberthroughJune,infruitfromFeb-ruarythroughJune.

Distribution(Fig.�0).Mexico(Colima,Guerrero,Jalisco,Michoacán);oakforestsandpine-oakforests;�00–�7�0m.

represeNtAtive specimeNs. mexico. coLimA: 9–�0 km E or SE by winding rd from Minatitlán,McVaugh 26226 (MICH).—Guerrero:Mpio.MontesdeOca,SanAntonio,Hinton et al. 11683 (GH,US).—JALisco: Mpio. Casimiro Castillo, 3–4 km SE of Casimiro Castillo;ArroyTacubaya, at base ofCerroLaPetaca,ca.20kmSSWofAutlán,�9°34'50–54"N,�04°24'29–45"W,Cochrane et al. 11687(WIS); Mpio.CasimiroCastillo,�–2kmalEdeCasimiroCastillo,Cuevas & Rosales 1828(IEB,WIS,ZEA);rdtoMicroondasLosMazosnearsummitofpass(PuertoLosMazos)betweenAutlánandLaHuerta,Daniel & Bartholomew 4849(CAS,MICH);Mpio.Tecalitlán,carr.Cd.Guzmán–Pihuamo,Km46,terraceríaLla-nitos–Mexiquilloa�6km,Fuentes O. 47(CHAPA,ENCB,IBUG,MICH);Mpio.Autlán,�2–�3kmalSSEdeAutlán,500–�000malSSEdeAhuacapán,�9°39'50"N,�04°�9'07"W,Guzmán & Cuevas 879(IBUG,MEXU,WIS,ZEA);Mpio.TalpadeAllende,brechadeTalpaaLaCuesta,Machuca N. et al. 7063(MICH);alonghwySWofAutlántowardManzanillo,onupperslopesofbarrancaabovethepass,McVaugh 10234(MEXU,MICH);precipitousmountainsides�0–�2kmabove(Nof)LaCuesta,belowpasstoTalpadeAllende,McVaugh 23357(ENCB,MICH);3kmSporbrechaaLasJoyas–Ahuacapán,Pepita86(ZEA);Mpio.Tecalitlán,Km6,brechaaJilotlándeDolores,Pérez de la Rosa 7(IBUG,MICH);Mpio.Cuautitlán,entreAyotitlányChancol,�9°28'N,�04°�0'W,Robles et al. 712(IEB,WIS,ZEA);Mpio.Cuautitlán,4–5kmalNWdeMinatitlán,2–3kmalNNEdePeñaColorada,LasPesadas,�9°24'N,�04°04'W,Santana M. et al. 4807(WIS,ZEA).—michoAcáN:Aquila,Dtto.Coalcomán,Hinton et al. 15867(NY,US).

FIG.�0.DistributionofGalphimia langlassei, G. oaxacana,andG. vestita.


Galphimia langlasseiisacommonshruborsubshrubofthepine-oakforestsoflowlandwesternMexico.Ithaselongateleavesbearingglandsonthemarginofthelamina,andtheovaryandfruitisabundantlypilose;onlyonecollectionexamined,Hinton et al. 11683,hadcalyxglandsonafewflowers.TheotherspeciesinitsrangewithpiloseovariesandfruitsareG. glandulosa, G. mexiae,andG. radialis.Galphimia glandulosaisreadilydistinguishedbyitselongate,whip-likeinflorescencesbearingflowerswithnearlyall-redpetalsandaglandularcalyx.Inaddition,theleafglandsareborneinthebasal�/6–�/2ofthepetiole.Galphimia radialisdiffersfromG. lan-glasseiinthatitspetalsaresubequalandthehairsontheovaryandfruitareto0.8mmlong;itisknownonlyfromcentralGuerrero.SeethediscussionofG. mexiae(no.9)foracomparisonwithG. langlassei.

9. Galphimia mexiaeC.Anderson,Syst.Bot.28:7�7.2003.—type: mexico.Jalisco:SanSebastián[delOeste]toHaciendadeOtotal,�425m,2Mar�927,Mexia 1778(holotype:CAS!;isotypes:A!GH!MO!US!). Fig.��.

Herborsubshrubto3m;stemssparselytomentulosewhenyoung,soonglabrous.Vestureofallvegetativepartsofsessile,wavytocrisped,reddishbrownhairs0.2–0.5mmlong.Laminasofthelargerleaves4–8.5(–�2)cmlong,�.8–4cmwide,lanceolatetoellipticaltonarrowlyrhombic,apexacuteorapiculate,basecuneateandsomewhatdecurrent,glabrous,secondaryveinsprominentorprominulousabaxially;petioles�–2(–3.5)cmlong,glabrous;glandsapairborneonthemarginofthelaminacom-monly ca. � cm above the base, each gland 0.4–0.8 mm in diameter, disklike andprominent,orrarelypeglikeandupto0.5mmlong;stipules2–3mmlong,0.5–�.8mmwide,lineartonarrowlytriangular,glabrousanddistallyciliate.Inflorescenceatermi-nalracemeorsmallpanicle,theaxestomentulose;peduncles3.5–7mmlong,pedicels8.5–�2mmlong,bothsparselytomentulosetoglabrate,peduncles0.3–0.5(–0.7)timesaslongaspedicels;bracts2–4mmlong,0.6–�.2mmwide,linear,bracteoles�.2–2mmlong[2.5–3.7mmlonginNelson 4060],0.4–0.6mmwide,linear,bractsandbracteolesglabrousorwithscatteredhairsalong themargindistally;bracteolessuboppositeorupto2mmapart,borneinthedistal�/4–�/2ofthepeduncleorsometimesjustbelowtheapex.Sepals2.5–3mmlong,�.4–2.3mmwide,oblongtonarrowlyellip-tical,glabrous,marginentiretodistallydenticulate(-ciliate);glandsabsent.Petalsdeciduous,unequal,yellowandsuffusedwithredalongtheclawandmidribofthelimb,glabrous,marginfinelydenticulate;lateralpetals:claw2–2.8mmlong,0.5–0.6mm wide, limb 4.5–6 mm long, 3.7–4.7 mm wide, ovate-triangular, apex subacutetoobtuse,basetruncateorslightlyattenuate;posteriorpetal:claw3.3–4mmlong,�.4–�.8mmwide,limb5–6.5mmlong,5–6.5mmwide,broadlytriangularorovate-triangular,apexbroadlyrounded,basetruncateoracute.Stamensofunequallength;filamentoppositeanteriorsepal(2.2–)2.6–3mmlong,filamentsoppositeanterior-lateralpetals(2.2–)2.4–3mmlong,filamentsoppositeanterior-lateralsepals2–2.8mmlong,filamentsoppositeposterior-lateralpetals�.8–2.3(–3)mmlong,filamentsoppositeposterior-lateralsepals�.5–�.8(–2.2)mmlong,filamentoppositeposteriorpetal�.5–�.7(–2)mmlong;anthers2.3–2.5(–2.8)mmlong.Ovarypilose,hairsto0.3mmlong;styles4–5mmlong.Coccus4.5–5.5mmhigh,3.5–4mmindiameter,pilosetoglabrescentinage;areole4–4.5mmlong,3–3.5mmwide;outercotyledon2.5–2.8mmlong,innercotyledon�.8–2.�mmlong.Chromosomenumberunknown.

Phenology.CollectedinflowerandfruitfromFebruarythroughMay.Distribution(Fig.25).Mexico(northwesternJaliscoandadjacentNayarit);pine-

oakforest;630–�880m.


FIG.��.Galphimia mexiae.a.Floweringbranch.b.Baseoflamina,showingmarginalglands.c.Flowerwithposteriorpetaluppermost,borneonpedicelandpeduncle,withportionofinflorescenceaxis.d.Lat-eralpetal.e.Posteriorpetal.f.Stamens,adaxialview,oppositeposteriorpetal(left)andposterior-lateralsepal(right).g.Gynoecium.h.Cocci,lateralview(right)andadaxialview(left).i.Embryo.Scalebar:a,4cm;b,8mm;c,�cm;d,e,5mm;f–h,4mm;i,2.7mm.(Basedon:a–c,Anderson & Anderson 6122,MICH;d–g,Nelson 4060,US;h,i,Almeda 2543,MICH.)

AdditioNAL specimeNs exAmiNed.mexico.JALisco:27.8miSSEofPuertoVallarta, Almeda 2543(CAS,MICH);lowhillsca.35km(strtline)EofCaboCorrientes,onrdfromPuertoVallartatoElTuito, Anderson & Anderson 6122(MICH);Mpio.Talpa,entreCumbredelTejamanilyCuale, González T. 102(MICH);Mpio.CaboCorrientes,steepmountainsides3–�0kmgenerallyEonrdtoMinadelCuale,fromjunction5kmNWofElTuito, McVaugh 26411A(MICH);trailSanSebastiántoLasMesitas, Mexia 1863-a(F,US);rdbetweenMascotaandSanSebastián, Nelson 4060(GH,US).—NAyArit:�7.7kmalWdeTepic,7kmalWdeCarranza, Cowan 4773(MICH,TEX);Mpio.Tepic,7kmalSdelaentradaalcaminodelCuarenteño,2�°28'N,�05°00'W, Flores F. 2531(MEXU).


Galphimia mexiaeisknownonlyfromasmallregioninnorthwesternJaliscoandadjacentNayarit.Itisdistinctiveinitsandroecium,composedofveryshortfilamentsandanthers.Allpetalshaveabroadlytriangularlimb;thatoftheposteriorpetalnar-rowsonlyslightlytoanexceptionallybroadclaw.ThesetraitsreadilyseparateitfromthesympatricG. langlassei,whichalsohasusuallynarrowlaminaswithglandsonthemarginnearthebaseandapiloseovaryandfruit.CollectionsofG. mexiae havebeenmisdeterminedasG. glandulosa(as“Thryallis dasycarpa”)andG. tuberculata,whichalsoshareapiloseovaryandfruit,butnotthecharactersoftheandroeciumandcorolla.Galphimiaglandulosaisentirelyglabrous,bearstheleafglandsintheproximal�/6–�/2ofthepetiole,andhaswhip-likeinflorescences;theflowershavenearlyall-redpetalsandaglandularcalyx.Galphimia tuberculataisnamedforthepersistenttuberclesfoundontheaxes,petioles,andlaminarmargins;G. mexiaelackssuchtubercles.

10. Galphimia radialisC.Anderson,Syst.Bot.28:7�9.2003.—type: mexico.Gue-rrero:Mpio.Chichihualco,29kmalWdeFilodeCaballo,carr.aPto.delGallo,2�00m,�8Aug�982, Tenorio L. 1471(holotype:MICH!). Fig.�2

Shrub to 3 m; stems tomentulose when young, soon glabrous.Vesture of allvegetativepartsofsubsessile,wavytocrisped,reddishbrownhairs0.2–0.7mmlong.Laminasofthelargerleaves7–�2cmlong,2–5cmwide,lanceolatetoelliptical,apexapiculate, base cuneate-decurrent, glabrous, secondary veins prominulous, marginentire;petioles�.5–2.3cmlong,glabrous;glandsapairborneonthemarginofthelaminacommonly2–2.5cmabovethebase,eachgland0.4–0.5mmindiameter,flushwiththemargin;stipules3.5–6mmlong,�.2–�.5mmwide,linear,glabrousordistallyciliate.Inflorescenceaterminalraceme,theaxestomentulose;peduncles4–9.5mmlong,pedicels7–�3.5mmlong,bothtomentulose,peduncles0.5–0.8timesaslongaspedicels;bracts3–6mmlong,0.5–0.7mmwide,linear,abaxiallywithscatteredhairsespeciallyalongthemargin,bracteoles�.6–3mmlong,0.4–0.6mmwide,linear,gla-brous;bracteolessuboppositeorupto2.2mmapart,theproximalbracteoleinsertedbelowtoca.themiddleofthepeduncle,thedistaloneataboutthemiddletothedistal�/4ofthepeduncle.Sepals3.5–4.5mmlong,�.6–2mmwide,lineartooblong,glabrousorwithafewscatteredhairsabaxially,margindenticulate-ciliateoronlyinthedistal�/4;glandsabsent.Petalsdeciduous,subequal,theposteriorpetalslightlylargerthanthelateralones,glabrous,yellowandsuffusedwithredalongtheclawandmidribofthelimb,margindenticulate-fimbriate;claw�.8–2.8mmlong,0.6–0.7mmwide,limb6.2–7.7mmlong,5.5–6.5mmwide,broadlyovate,apexbroadlyrounded,basesubtruncate.Stamensofunequallength;filamentoppositeanteriorsepal4.5–4.8mmlong,filamentsoppositeanterior-lateralpetals4.2–4.5mmlong,filamentsoppo-siteanterior-lateralsepals4.4–4.5mmlong,filamentsoppositeposterior-lateralpetals3.2–4mmlong,filamentsoppositeposterior-lateralsepals4–4.3mmlong,filamentoppositeposteriorpetal3–3.5mmlong;anthers2.2–2.5mmlong.Ovarypilosealongthesuturesandinthebasal�/3–�/2,hairsto0.8mmlong;styles4.6–5.2mmlong.Coc-cus4.7–5mmlong,3.8–4mmindiameter,pilosealongthesuturesandinthebasal�/3–�/2;areoleca.4mmlong,2.5–3mmwide,matureseednotseen.Chromosomenumberunknown.

Phenology.CollectedinflowerinJanuary,May,August,andNovember,infruitinMarchandAugust.

Distribution(Fig.25).Mexico(Guerrero);pine-oakforest;�700–2�00m.


FIG.�2.Galphimia radialis.a.Leaf.b.Baseoflamina,showingmarginalglands.c.Node,showinglongstipules.d.Flowerbud,borneonpedicelandpeduncle,withportionofinflorescenceaxis.e.Lateralpetal.f.Posteriorpetal.g.Stamens,abaxialview,oppositeposteriorpetal(right)andposterior-lateralsepal(left).h.Gynoecium.i.Cocci,lateralview(left)andadaxialview(right).Scalebar:a,4cm;b,c,�cm;d,�.3cm;e,f,5mm;g–i,4mm.(Basedon:a,Estrada R. 119,IEB;b–d,Fonseca 1001,MEXU;e–h,Tenorio L. 1471,MICH;i,Paray 2014,ENCB.)

AdditioNAL specimeNs exAmiNed. mexico.Guerrero:Mpio.AtoyacdeAlvarez,trayectoentreLosArrayanesyElDescanso,Estrada R. 119(IEB);Mpio.Chichihualco,cercadeHierbaBuena,Fonseca 1001 (MEXU);Mpio.Tlacotepec, faldasdelCerroTlacotepec,Paray 2014 (ENCB);CerroTeotepecyalrededores,Paray 4065(MEXU);Mpio.Malinaltepec,Malinaltepec,�7°�4'N,98°40'W,Wagenbreth 807(MO).

Galphimiaradialisistheonlyspeciesamongthosewithwell-developedpedun-clesinwhichthepetalsaresubequal.Italsohaslargerstipules(3.5–6mmlong)thanmostspecies(usually�–3mmlong),andunusuallylonghairsontheovary(to0.8mmlongvs.0.�–0.3mmlong).ItmaybeconfusedwithG. langlassei,whichhassimilarleaves,butunequalpetalsandlongeranthers(3–4mm).

11. Galphimia sessilifoliaRose,Contr.U.S.Natl.Herb.3: 3�3.�895.Thryallis ses-silifolia(Rose)Rose,Contr.U.S.Natl.Herb.�2:28�.�909.—type: mexico.Oaxaca:LasSedas,6000ft,�Aug�984,Pringle 4901 (holotype:US!;isotypes:A!BM!BR!CM,F,G!GH!GOET!K!MEXU!MO!NY!P!S!UC!US!W!;photoofPisotype:MICH!). Fig.�3.

Shrub toca.�m; stemssparsely strigosewhenyoung, soonglabrous.Vestureofallvegetativepartsofsubsessiletosessile,straighttowavy,reddishbrownhairs0.2–0.7mmlong.Laminasofthelargerleaves�.5–4cmlong,�–2.5cmwide,ovateornarrowlyso,apexapiculate,basetruncateorshallowlycordate,glabrous,coriaceous,


FIG.�3.Galphimia sessilifolia. a.Floweringbranch.b.Baseofleafwithglands,abaxialview.c.Nodewithstipules.d.Detachedlargeleaf.e.Posteriorpetal(left)andlateralpetal(right).f.Androeciumlaidout,abaxialview,thestamenatleftoppositeposteriorpetal.g.Gynoecium.h.Fruitborneonpedicelandpeduncle,withportionofinflorescenceaxis.i.Coccus,adaxialview.j.Coccus,lateralview.Scalebar:a,d,4cm;b,c,f,g,i,j,4mm;e,h,8mm.(Basedon:a,b,f,g,Lorence 4626,MEXU;c,Torres C. 7151,MICH;d,Pringle 4901, MEXU;e,Pringle 7458,MEXU;h,Lorence 4626,MO;i,j,Salinas T. 6607,MICH.)


secondaryveinsprominulousabaxially;marginentire;leavessubsessile,petiolesto�mmlong,glabrous;glandsabsent,orpresentonlyonthefirstpairofleavesatthebaseofashoot,theselaminasthesmallestandsuborbicularandwith�or2glandsborneonthemarginatornearthebase,glands0.3–0.4mmindiameter,disklikeandsessile,orpeglikeandupto0.7mmlong;stipules�–2mmlong,ca.0.5mmwide,lin-ear,glabrous.Inflorescenceasolitaryterminalraceme,theaxesglabrate;peduncles�–5mmlong,pedicels7.5–�2mmlong,bothsparselytomentulose,peduncles0.2–0.6(–0.9) timesas longaspedicels;bracts �.5–2.5mm long, 0.7–0.8mm wide, linear,bracteoles �–2 mm long, ca. 0.5 mm wide, linear, bracts and bracteoles glabrous;bracteolesbornevariouslyon thepeduncle fromnear thebase to justbelow theapex,suboppositeorupto0.5mmapart.Sepals2.5–3mmlong,�.2–�.5mmwide,oblong,glabrous,margindenticulate-ciliateinthedistal�/4–�/3;glandsabsent.Pet-alsdeciduous,unequal,yellowandtingedwithredattheapexandoftenwitharedstripeabaxiallyalongthemedianvein,glabrous,margindenticulate;lateralpetals:claw2–2.5mmlong,ca.0.5mmwide,limb6.5–7mmlong,3.5–4.5mmwide,oblongornarrowlyovate,apexobtuse,baseattenuatetosubtruncate;posteriorpetal:claw3–4mmlong,�.2–�.4mmwide,limb6.5–7mmlong,5–6mmwide,triangular,apexbroadlyrounded,basecordate.Stamensofunequallength;filamentoppositeante-riorsepal5–5.3mmlong,filamentsoppositeanterior-lateralpetals4–4.5mmlong,filamentsoppositeanterior-lateralsepals4.5–5mmlong,filamentsoppositeposte-rior-lateralpetals3–3.2mmlong,filamentsoppositeposterior-lateralsepalsca.4.5mmlong,filamentoppositeposteriorpetalca.3mmlong;anthers2.5–4mmlong.Ovaryglabrous;styles5.5–7mmlong.Coccusca.3.5mmlong,ca.2.5mmindiameter,smooth,glabrous;areoleca.2.7mmlong,ca.2.3mmwide;matureseednotseen.Chromosomenumberunknown.

Phenology.CollectedinflowerinJulyandAugust,infruitinJuly,August,andDecember.

Distribution(Fig.8).Mexico(Oaxaca);inaridscrub,matorral,drysitesinpine-oakanddeciduousforest;�980–2400m.

AdditioNAL specimeNs exAmiNed. mexico. oAxAcA: near Penan, Conzatti s.n. (US); along hwybetweenOaxacaandTehuacán,26.2miNNWjct.Hwy�90,Daniel & Baker 3758(MICH);Mpio.Huitzo,Dtto.Etla,ca.0.5kmalEdeLasSedas,Lorence 4626(F,MEXU,MICH,MO,UAMIZ);�5kmbyrdSEofMiahuatlánonrdtoPuertoAngelinhighmtnsofSierraMadredelSur,�6°�2'N,96°30'W,Marcks & Marcks 1003(LL,MICH,TEX,WIS);LasSedas,Pringle 7458(F,MEXU);Mpio.SanJerónimoSosola,ElParián,Dtto.Etla,�7°23'N,97°00'W, Salinas T. 6607(CAS,MICH);Dtto.Tlaxiaco,3.7kmalNEdelOjite,cercadeTlaxiaco,Torres C. 7151(F,MEXU,MICH,UAMIZ).

Galphimia sessilifolia,knownfromafewcollectionsfromOaxaca,isreadilyrec-ognizedbyitssubsessile,ovate,coriaceousleaves;leafglandsdeveloponlyonthefirstpairofleavesofashoot,whichhasnotablysmallerandorbicularlaminas.Galphimia multicaulis,whichoccursinOaxaca,istheonlyotherspeciesinthegenusinwhichthepetiolesarerudimentaryorveryshort.UnlikeG.sessilifolia, ithaspersistentpetalsandglandspresentonthemarginofalllaminas.

12. Galphimia mirandaeC.Anderson,Contr.Univ.MichiganHerb.24:�3.2005.—type: mexico.Puebla:Mpio.Tehuitzingo,�2kmalNWofElPitayo,carr.aIzúcardeMatamorosPuebla,�350m,28Jul�983,Torres C. 3296(holotype:MEXU;isotype:MBM!MO!). Fig.�4.


Shrubortreeletto4m;stemstomentulose,soonglabrescenttoglabrous.Ves-tureofallvegetativepartsofsessile,wavytocrisped,reddishbrownhairs0.4–�mmlong.Laminasofthelargerleaves3–6.5cmlong,2–4cmwide,ellipticaltobroadlyso, obovate, rhombic, ovate, or suborbicular, apex obtuse to acute, base acute todecurrent(especiallyinlargerleaves),glabrous,secondaryveinsprominulousornotevident;marginentire;petioles0.5–�cmlong,glabrous;glandscommonlyapaironthemarginofthelaminawellaboveoratthebase,ifbasedecurrentthensuperficiallyappearingplacedonthepetiole,sometimes�ormoreadditionalglandsborneonthemarginofthelamina,eachgland0.5–�mmindiameter,disklikeandwithathickrim;stipules2–4mmlong,0.7–�.2mmwide,narrowlytriangular,glabrous.[Sometimeswithonlytheterminalbranchesbearingaflushofsmallleavesalongshortinternodes,thelaminas�.5–2.5cmlong,�–�.5cmwide.]Inflorescenceaterminalraceme,theaxes,peduncles,andpedicelstomentulosebutglabrescentinolderparts;peduncles3–6mmlong,pedicels�4.5–�8.5mmlong,peduncles0.2–0.3timesaslongaspedi-cels; bracts 2.5–3.2 mm long, 0.5–0.8 mm wide, linear, bracteoles �.5–2 mm long,0.3–0.5mmwide,linear,bractsandbracteolesglabrousorwithafewscatteredhairsabaxially;bracteolesborneataboutthemiddleofthepeduncle,suboppositeorupto0.3mmapart.Sepals3.5–4.2mmlong,�–�.5mmwide,narrowlyovatetooblong,glabrous,marginentireorfinelydenticulateattheapex;glandsabsent,orrarelywithonegland0.5mmindiameter.Petalsdeciduous,unequal,yellow,glabrous,marginfinelydenticulate;lateralpetals:claw2.5–2.8mmlong,0.5mmwide,limb7–8.5mmlong,4.5–5.5mmwide,ellipticalornarrowlylanceolate,apexsubacute,basegradu-allyattenuate;posteriorpetal:claw4–4.2mmlong,�–�.2mmwide,limb6.2–7.5mmlong,6–7mmwide,triangularorsometimessuborbicular,apexobtuse,baseslightlycordateorslightlyso.Stamensofunequallength;filamentoppositeanteriorsepal

FIG.�4.Galphimia mirandae.a.Leafwithovatelamina(detailofbaseoflaminashowingmarginalgland).b.Leafwithellipticallamina.c.Flowerborneonpedicelandpeduncle,withportionofinflorescenceaxis;notecalyxgland.d.Lateralpetal.e.Posteriorpetal.f.Sixstamenslaidout,abaxialview,stamenatleftoppositeposteriorpetal.g.Gynoecium.Scalebar:a,4cm(4mm);b,4cm;c,�cm;d,e,5.7mm;f,g,4mm.(Basedon:a,Miranda 2147,MEXU;b–g,Torres C. 3296,MEXU.)


4–4.5mmlong,filamentsoppositeanterior-lateralpetals3–3.2mmlong,filamentsoppositeanterior-lateralsepals3.7–4mmlong,filamentsoppositeposterior-lateralpetals 2–2.2 mm long, filaments opposite posterior-lateral sepals 3.6–4 mm long,filamentoppositeposteriorpetal2–2.3mmlong;anthers3.3–3.7mmlong.Ovaryglabrousorwithscatteredhairsonthesutures,hairsto0.�mmlong;styles5–6.6mmlong.Maturecoccusnotseen.Chromosomenumberunknown.

Phenology.CollectedinflowerandimmaturefruitinJuly.Distribution(Fig.5).Mexico(Guerrero,Puebla);tropicaldeciduousforest;�350m.

AdditioNAL specimeN exAmiNed.mexico.Guerrero:8kmalNWdeChilpancingo,sobrelacarreteraaChichihualco,�350m,7Jul�966,Asteinza(ENCB),Cabrera s.n.(ENCB),Rzedowski 22755(DS,ENCB,MICH,TEX).—puebLA:[Izúcarde]Matamoros,24Jul�942,Miranda 2147(MEXU).

Galphimiamirandaehassucculentlaminaswithlarge,thick-rimmedglandsonthemargin(Fig.�4a);ifthebaseisdecurrenttheglandsmayappeartobeborneonthepetiole,but carefulexamination shows themembedded in the laminar tissue.Thespeciesisalsodistinctiveinthatthepedicelsareverylong(�4.5–�8.5mm)andgreatlyexceedthepeduncles.Galphimiamirandaeisknownfromonlyfivegather-ings;ofthesethetwoPueblacollectionsarefromnearbylocalities,andlabeldataoftheGuerrerocollectionsindicatethattheywereprobablyobtainedfromthesamepopulation.Additionalcollectionsareneeded todetermine thevariabilityof leafshape,stylelength,andovaryandfruitpubescence.

Group ii(petalsdeciduous;pedunclesabsentorrudimentary)

13. Galphimia angustifoliaBenth.,Bot.voy.Sulphur,9,pl.5.�844. Thryallis angusti-folia(Benth.)Kuntze,Rev.gen.pl.�:89.�89�[combinationalsoproposedbyRose,Contr.U.S.Natl.Herb.�2:280.�909].—type:mexico.BajaCaliforniaSur:CapeSt.Lucas,�84�,Hinds s.n.(holotype:K!,photo:MICH!).

Galphimia linifolia A.Gray,Gen.amer.bor.2:�96,pl.�73.�849.Thryallis linifo-lia(A.Gray)Kuntze,Rev.gen.pl.�:89.�89�.—type: u.s.A.[Texas:]banksoftheMedinaRiver,Jun�849,Wright 93(lectotype,heredesignated:GH!;isolectotypes:BM!G!MEL!NY!P!US!).

Galphimia linifoliavar.oblongifoliaA.Gray,Pl.Wright.�:36.�852.Galphimia angustifoliavar.oblongifolia(A.Gray)S.Watson,Proc.Amer.Acad.Arts.24:42.�889.Thryallis angustifoliavar.oblongifolia(A.Gray)Vail,Bull.TorreyBot.Club22:228.�895.—type: u.s.A.[Texas:]banksoftheMedinaRiver,Jun�849,Wright 94(holotype:GH!;isotypes:BM!DS!G!K!NY!US!).

Galphimia angustifolia var. ovalifolia Arènes, Notul. Syst. (Paris) �2: �89.�946.—type:mexico.Sonora:Guaymas,Palmer 217in�887(lectoype,heredesignated:P!,photo:MICH!;isolectotypes:BM!C!GH!K!NY!UC!US!). [Cited erroneously byArènes as“Scott Elliot, Madagascar, sans num. niloc.”]

Fig.�5.

Suffrutescentherbandsmallshrubto�m;stemsdenselypubescentwhenyoung,glabrescentinage.Vestureofallvegetativepartsofstraighttowavy,whitetored-dishbrown,subsessilehairs0.6–�.5mmlong.Laminasofthelargerleaves2.5–4cmlong,0.3–2cmwide,lineartonarrowlyellipticalornarrowlylanceolateornarrowlyovatetoellipticalorovate,apexacuteorapiculate,baseacute,glabrousorabaxially


FIG.�5.Galphimia angustifolia.a.Floweringhabit.b.Wideleaf,adaxialview.c.Baseoflaminain“b,”abaxialview.d.Narrowleaf,abaxialview.e.Baseoflaminain“d,”abaxialview.f.Flowerbud,withsessilepedicelsubtendedbybractandtwobracteoles.g.Petal.h.Partialandroeciumlaidout,abaxialview,thestamenatrightoppositeposteriorpetal.i.Gynoecium.j.Fruit.k.Twococci,adaxialview(left)andlateralview(right).l.Twococci,adaxialviews.m.Embryo.Scalebar:a,4cm;b,d,2cm;c,e,h,i,m,2mm;f,j,4mm;g,k,l,2.7mm.(Basedon:a,Pringle 1940,GH;b,c,Reverchon 1505,NY;d,e,Correll 29602,LL;f,Miller et al. 5665,MICH;g–i,McGregor 16749,US;j,l,m,Daniel 6895,CAS;k,Butterwick & Lott 3728,TEX.)


withsomescatteredhairsontheproximalportionofthecosta,secondaryveinsnotevidentor slightlyprominulousabaxially;marginentire;petioles0.�–0.9cm long,pubescentbutglabrescentwithage;glandsusuallyapairborneonthemarginatthebaseofthelaminato5mmabovethebase,orsometimeswithonly�glandorrarelytheglandsabsent,eachgland0.�–0.3mmindiameter,sessileorprominent,orpeglikeandupto0.4mmlong,sometimeswith�orrarely2additionalsmallerglandsborneonthemarginintheproximalhalfofthelamina;stipules2–3.5mmlong,0.3–0.5mmwide,linear,glabrousorwithscatteredhairsonthemargin.Inflorescenceaterminalraceme, theaxes sparsely todensely pubescentbut glabrescent inage,pedunclesusuallyabsentorrudimentary(rarelyto3.5mmlong),pedicels2–�0mmlong,pubes-centorsparselyso;bracts2–3.7mmlong,0.5–0.7mmwide,linear,bracteoles�.2–�.5(–�.8)mmlong,0.2–0.3 (–0.5)mmwide, linear,bractsandbracteolesglabrousorwithscatteredhairsalongthemarginoronlyatapex.Sepals3.2–3.5mmlong,�.3–�.5mmwide,narrowlylanceolate,glabrousbutwithatuftofhairsattheapex;glandsabsent.Petalsdeciduous,subequal,theposteriorpetalsometimesalittlelarger,yel-low,becomingorangeorredinage,glabrous,marginslightlyerose-undulate;claw�.5–2mmlong,0.4mmwide,limb3.5–4.5mmlong,2.3–3mmwide,ovatetonarrowlytriangular,apexobtuse,basesubtruncatetoacute.Stamensofunequallength;fila-mentoppositeanteriorsepal3.7–4mmlong,filamentsoppositeanterior-lateralpetals2.6–3.5mmlong,filamentsoppositeanterior-lateralsepals3.2–3.7mmlong,filamentsoppositeposterior-lateralpetals2.7–3.3mmlong,filamentsoppositeposterior-lateralsepals3.2–3.7mmlong,filamentoppositeposteriorpetal2.5–3.3mmlong;anthers0.7–0.9mmlong.Ovarypilose,hairsca.0.�mmlong;styles3.5–4.8mmlong.Coccus3–3.5mmlong,2–2.2mmindiameter,pilose,glabrescentinage;areole2–3mmlong,�.5–2mmwide;outercotyledon2.3–2.7mmlong,innercotyledon�.8–2.2mmlong.Chromosomenumberunknown.

Phenology. Collected in flower and fruit from April through November(–December),inSonoraalsoinJanuaryandFebruary,inBajaCaliforniaSurfromAugustthroughMay.

Distribution(Fig.�6).U.S.A.(Texas)andMexico(BajaCaliforniaSur,Coahuila,Durango,NuevoLeón,SanLuisPotosí,Sinaloa,Sonora,Tamaulipas);onlimestone,caliche,andgypsumsubstratesindryanddeserthabitats,inwashesandarroyos,inopentropicaldeciduousforest,matorral,andthornscrub,ondunes;sealevelto�730m.

represeNtAtive specimeNs. U.S.a.texAs.BanderaCo.:MedinaLake,Palmer 12269 (A,CAS,GH,MO).BeeCo.:NofBeeville,Albers 46332 (F,TEX).BexarCo.:nearBracken,Groth 64 (CAS,F,GH,US).BlancoCo.:PedernalesFallsStatePark,studyplotno.�,Oefinger 324 (TEX).BrewsterCo.:abovebedofSanFranciscoCanyon,ca.3miupfromtheRioGrande,ca.�8miSofSanderson,Butterwick & Lott 3728 (TEX).ComalCo.:SmithsonValley,�5miNWofNewBraunfels,Cutler 3248 (F,GH,MO,NY).DuvalCo.:7miSofSanDiegotowardsBenavides,Correll & Johnston 25504 (S,TEX).EdwardsCo.:3/4miSofdamonHackberryCreek,WsidebelowDeadman’sHollow,Smith & Butterwick 224 (LL).FayetteCo.:withoutlocality,Matthes 241 (G,W).GoliadCo.:McNamaraRanch,0.2miNEofFM-�35�onSsideofHwy59,Hill 5462 (ENCB).HaysCo.:�2miWofSanMarcosonrdtoWimberly,EdwardsPlateau,Johnson 266 (TEX);SanMarcosandvicinity,Stanfield s.n. (NY).JimHoggCo.:withoutlocality, Lehmann & Davis 6 (F).KarnesCo.:0.2miWofCoyCity,Farm-to-MarketRd8�,Johnson 1305 (TEX).KendallCo.:�3miNWofBoerne,Barneby 14496 (CAS,NY).KerrCo.:Heller 1737 (BR,C,F,G,GH,K,MICH,MO).KimbleCo.:NofCastroville,Reverchon 1505 (DS,F,MO,NY,S).KinneyCo.:AnacachoMtsonAnacachoRanch,Correll 30433 (GH,MO,TEX).LiveOakCo.:rdtoFiestaMarinajustEofRte534,4.6miNofRte359,Hill 10628 (ENCB,GH,MICH,MO,NY).MedinaCo.:alongstatefarmrd�283,4miEofMedinaDam,29°23'N,99°24'W,Miller et al. 5841 (MICH,MO).MendocinoCo.:S.GeronimoValley,Reverchon 26 (GH).RealCo.:lookoutoverFrioRiver,alongRte83severalmiNEofLeakey,Correll 29602 (GH,LL).SanPatricioCo.:ca.2miNWofMathisalongedgeofFrelscalichepit,Turner 80-86M (TEX).StarrCo.:2kmSofSantaMargarita,betweenFalconandRoma,Butterwick & Strong 1335 (TEX).


TravisCo.:MountBonnell,Albers et al. 46460 (F,MO,NY,S,TEX).UvaldeCo.:2miNofBlewett,Muller 8013 (LL,MICH).ValVerdeCo.:abovedamatfootofDevil’sLake,ca.20miNNWofDelRio,McVaugh 10589 (BM,DUKE,ENCB,G,GH,MICH,NY,LL);lookoutoverFrioRiver,alongRte83severalmiNEofLeakey,29°28'N,�00°58'W,Miller et al. 5665(MICH,MO,TEX).Countyunknown:UpperGuadelupe,Lindheimer 361 (BM,G,GH,K,MO,NY-fragment,P,US,W).

mexico.bAJA cALiforNiA sur: EofCaboSanLucas, Carter 2259(DS,MEXU,UC,US);ArroyodelSalto,EofLaPaz,24°�2'N,��0°7.5'W, Carter 2586(DS,K,UC,US);CañóndelosPotrerillos,ca.5kmNWofLoreto,26°0�'N,��°22'W, Carter 4590(BM,MEXU,MICH,UC);CalienteManantialsalongRíoAguaCaliente,�3kmSWofSantiago,foothillsofEslopeofSierradelaLaguna,23°26'N,�09°49'W, Daniel 6895(CAS,MICH);LasCuevitasbelowComondu, Gentry 4223(DS,GH,MO,UC,US);BahíaSanNicolás, Johnston 3736(CAS,GH,MO,NY,UC,US);Miraflores, Jones 24169(A,CAS,F,MO,UC);WsideofSantaCruzIsland,25°�7'N,��0°44'W, Moran 3825(DS);hillSofMulegé,26°52'N,��°59'W, Moran 9037(MICH,SD,UC);DanzanteIsland, Moran 9264(CAS,SD);aboveEnsenadaBallena,EspírituSantoIsland,29°29'N,��0°20'W, Moran 9651(SD);�0miSofMulegé, Shreve 7085(DS,F,MICH,US);5miNofSantiago, Shreve 7288a(DS,F,MICH,US);IslaSta.Catalina,ladoSWdelaisla,a�0mdelaplaya,25°35'N,��0°47'W, Sousa, P. 92(CAS,ENCB,IBUG,IEB,MEXU);CapeRegion,ArroyoSantaAnitanearCasaVieja,ca.5miSWofLaPalma,8miNWfromSantaAnita, Thomas 7740(CAS,DS,ENCB,GH,MEXU,UC,US);SanJosédeMagdalena, Wiggins 11380(CAS,DS,ENCB,GH,MO,NY,SD,UC,US);�.5miNWofSanBártolo, Wiggins 14755(CAS,DS,GH,K,MEXU,TEX,UC);CapeSt.Lucas, Xantus 15(GH,K,NY,US).—coAhuiLA:nearRanchoMargaritaheadquarters,SerraníasdelBurroMts,65miNWofSabinas,Gould 10649(ENCB,MICH,TEX);32airmiNEofSanPedro,�miSWofLasDelicias,26°�4'N,�02°49'W,Henrickson 6061(TEX);�3kmNofLasMargaritas,26°34'30"N,�02°50–5�'W,Johnston et al. 10357E(CAS,LL,MEXU,NY);mts24miNEbyNfromMonclova,Palmer 128in�880(F,G,GH,NY,US);5kmNEofJimulco,25°N,�03°W,Stanford et al. 134(DS,GH,MEXU,MO,NY,UC);canyonsinSierradelSobacoafewkmWofLasDelicias,Stewart 2802(GH);�2.6kmNNEofLasMar-garitas,26°34'N,�02°5�'W,Wendt et al. 9491(CAS,LL,MEXU,NY);Mpio.Múzquiz,HaciendaMariposa,

FIG.�6.DistributionofGalphimia angustifolia.


nearPuertoSantaAna,Wynd & Mueller 282(A,K,MICH,MO,NY,S,US);Mpio.Torreón,SierradeJimulco,�50kmalEdeLaMinadeSanJosé,25°06'N,�03°�3'W,Villarreal 5520(TEX).—durANGo:30kmSWofGómezPalacios,25°24'30"N,�03°43'W,Johnston et al. 10400C(CAS,F,LL,MEXU,MO,NY);20kmNWofEstaciónChocolate,25°25'N,�03°43'W,Johnston et al. 12199(CAS,F,LL,MEXU,MO,NY,SD).—Nuevo LeóN:5miSofSabinasHidalgo,alongroadtoMonterreynearKm�074,Chute M-260(MEXU,MICH);Linares,Clark 6809(MO,NY);Mpio.SabinasHidalgo,�2.8miSofSabinasHidalgo,Cowan 5354(NY,TEX);ca.25kmNWofMonterrey,alongrdtoNuevoLaredo,Dieterle 3677(ENCB,MEXU,MICH);Mpio.SabinasHidalgo,�2miSofSabinasHidalgo,26°20'N,�00°07'W,Dorr 2484(CAS,CHAPA,MEXU,NY,TEX);betweenMonterreyandMontemorelos,Dziekanowski et al. 1736(ENCB,NY);��miNEofSabinasHidalgo,5miSWofVallecillos,Johnston & Graham 4607(MEXU,MICH,TEX);Obispado,LosAlamillos,Lacás 408(F);29miNofSabinasHidalgo,McGregor 16749(LL,US);Mpio.Villaldama,SierraGomas,inCanyonElAlamo,26°2�'N,�00°27'W,Patterson 6671(TEX);Monter-rey,Pringle 1940(BM,F,G,GH,L,NY,S,UC,US,W);hillsnearMonterrey,Pringle 2708(F,MEXU,MO,NY,UC);40miSofLaredo,Schery 16(MICH,MO);7.3miSofSabinasHidalgo,Weaver 2024(DUKE,MEXU,MO).—sAN Luis potosí:3miSofTamaulipaslineonAntiguoMorelos–Valleshwy,Johnston & Graham 4498(MEXU,MICH,TEX);withoutlocality,Parry 94(GH,K,MO,NY,P,USF).—siNALoA:Cer-rosdelFuerte,�8–24miNofLosMochis,Gentry 14291(LL,MEXU,MICH,US);CerrosdeNavachisteaboutBahíaTopolobampo,Gentry 14365(US);hillsnearYachtHotel,Topolobampo,Hastings & Turner 64-112(DS,SD);Topolobampo,Palmer 184in�897(C,DS,MICH,P,S,UC,US).—soNorA: CañónNaca-puli,ca.4kmNofBahíaSanCarlos,Felger 85-865(MICH);CañónlasBarajitas,SierraElAguaje,ca.�8kmNWofSanCarlos,28°02'32.0"N,��°�2'40.�"W,Felger 95-128(CAS);EnsenadaGrande(=SanPedroBay),Felger et al. 11570(SD);3.6mibyroadNEofNewKino,Felger 15297(ENCB,LL,SD);Navojoa,Gentry 1661(F,GH,K,MO);EnsenadadePerros,ladoEdelaIslaTiburón,28°46'N,��2°�6'W,Tenorio L. 9503(TEX);SanAgustínBay,28°03'N,��°04'W,Turner 79-26(MEXU,UC);RíoMayoregion;baseofCerro(Mesa)Masiacaonrdtomicrowavetower,�.3miEofHwy�5(SEofNavajoa),26°46'35"N,�09°�8'30"W,Van Devender 95-1090(MICH).—tAmAuLipAs:CerroTinaja,vicinityofSanJosé,SierradeSanCarlos,Bartlett 10319(LL,MEXU,MICH,US);LaTamaulipeca,vic.ofSanMiguel,SierradeSanCarlos,Bartlett 10582(ENCB,F,LL,MEXU,MICH,NY,US);Mpio.CiudadVictoria,�3.6kmfromVilladeCasastowardSotoLaMarina,Cowan 5211(CAS,NY,TEX);alonghwybetweenGonzálesandEst.Zaragoza,5.4miSEofturnofftoEst.Zaragoza,23°�0'N,98°44'W,Daniel 811(MICH);alongrdtoSanFrancisco,�.0miEHwy85SofCd.Victoria,Daniel & Baker 3705(MICH);4.7miWofjunctionofHwy80&85onHwy80,alonggradeWofAntiguoMorelos,Dunn et al. 17574(ENCB);22miSofVillagran,Johnston & Crutchfield 4310A(TEX);vicinityofVictoria,Palmer 483in�907(F,GH,NY,US);rdbetweenLaPescaandSotoLaMarina,8.�miEofSotoLaMarina,23°5�'N,98°00'W,Patterson 7272(TEX);Jau-mave,Viereck 665(US).

Galphimia angustifolia is a commonspeciesof southernTexasandnorthernMexico; it is the only species of Galphimia found in the United States. It formssprawlingsuffrutescentherbsorsmallshrubscomposedofmanyascendingshootsarising from a woody base and terminating in lax, small-flowered inflorescences.Leafshapevariesgreatlyfromlineartoovate,andsomeregionalpatternscanbediscerned.InmostcollectionsfromBajaCaliforniaSurandSonorathelaminasareellipticaltoovate,yetinsome,includingthetypeofG. angustifolia,andthosefromSinaloa,thelaminasarevarynarrow.Collectionsfromtheremainderoftherangehave mostly linear to narrowly lanceolate/elliptical laminas, but specimens withbroaderlaminasarenotuncommon.Onoccasion,aspecimenbearsbroaderlaminasnearthebaseofashootandnarroweronesdistally(Fig.�5a).SuperficiallyG. angus-tifolia ismostsimilartotheequallywidespreadandvariableSouthAmericanG. australis,fromwhichitisreadilyseparatedbyitspiloseovaryandfruitandeglandu-larcalyx.InMexicoG. angustifoliaiseasilydifferentiatedfrommostotherspeciesofGalphimiainitsrangebythebroom-likehabit,lackofpeduncles,andsmallflowers;theonlyspeciessharingthesecharactersisthedenselypubescentG. vestita.

14. Galphimia vestitaS.Watson,Proc.Amer.Acad.Arts2�:42�.�886.Thryallis vestita(S.Watson)Rose,Contr.U.S.Natl.Herb.�2:28�.�909.—type:mexico. Chi-huahua:HaciendaSanMiguel,nearBatopilas,Palmer 187in�885(holotype:GH!,photos:A!F!NY!;isotype:US!). Fig.�7.


FIG. �7. Galphimia vestita. a. Habit. b. Node with stipules (and detached marginal leaf gland).c.Abaxialleafsurface.d.Flowerbud,anteriorsepalbentdown.e.Flower.f.Petal,abaxialview.g.Anthers,adaxialview(left)andabaxialview(right).h.Fruit.i.Cocci,abaxialview(above)andadaxialview(below).j.Coccuswithhalfofabaxialwallremoved.Scalebar:a,4cm;b,4mm(2mm);c,h–j,4mm;d,5.7mm;e,f,5mm;g,2mm.(Basedon:a–d,White 3574,GH;e–j,Wiggins 7471,DS.)


Herborsmallshrubfromwoodyrootstock,to0.5m;stems, leaves,andinflo-rescencedenselypubescent.Vestureofallpartsofwhiteor translucentT-shapedhairs,thetrabecula�–2mmlong,straighttowavyorcrisped,thestalkca.0.5mmlong.Laminasofthelargerleaves2.5–5cmlong,0.2–0.8cmwide,linear,apexacutetoacuminate,baseacute,secondaryveinsnotevident;marginentire;petioles�–3cmlong;glandsusuallyapairborneonthemarginto2mmabovethebaseofthelaminaorsometimesatthebase,eachglandca.0.�mmindiameter,disklikeandprominent,orpeglikeandupto0.5mmlong,sometimesthelaminartissueextendedintoatoothandtheglandborneattheapex;stipules3.5–9mmlong,0.5–0.8mmwide,linear.Inflorescenceaterminalraceme;pedunclesabsent,pedicels4.5–9(–��)mmlong;bracts4–�4mmlong(oftenexceedingthepedicel),0.5–�mmwide,linear,bracteoles�.5–2.5mmlong,0.2–0.3mmwide,linear.Sepals4.5–5.3mmlong,�.5–�.7mmwide,narrowly triangular tonarrowly lanceolate,pubescentabaxially;glandsabsent.Petalsdeciduous,subequal,yelloworyellowsuffusedwithredespeciallyonthemidrib,becomingredinage,pubescentonthemidribabaxially,marginshallowlyerose-undulate;clawca.�.5mmlong,0.3–0.4mmwide,limb3.5–5mmlong,3mmwide,narrowlyovateornarrowlyelliptical,apexobtuseorapiculate,basegraduallyacuteorsubtruncate.Stamenssubequal;filaments4–5.5mmlong;anthers�.3–�.5mmlong.Ovarypilose,hairsca.0.�mmlong;styles6.8–7.8mmlong.Coccus3.5–4mmlong,2.3–2.4mmindiameter,pilose,inagethehairssloughedoffthesurfacebutretainedalongthesutures;areole2.8–3mmlong,ca.2mmwide;outercotyledonca.2.5mmlong,innercotyledonca.2mmlong.Chromosomenumberunknown.

Phenology.CollectedinflowerfromJulythroughDecember,andinfruitfromAugustthroughOctober.

Distribution (Fig. �0). Mexico (southwestern Chihuahua, Nayarit, Sinaloa,Sonora);inthornforestandsandywashes;400–600m.

AdditioNAL specimeNs exAmiNed. mexico.NAyArit:Mpio.ElNayar,JesúsMaría,caminoalapistadeaterrizaje, Colunga & Zizumbo 42(CAS,MEXU);Mpio.Acaponeta,nearCancialalongtrailWofJesúsMaría, Norris & Taranto 14063(MICH), 14085(MICH).—siNALoA:Cofradia,vic.ofCuliacán, 20Oct�904,Brandegee s.n.(UC);Cofradia, 27Oct�904,Brandegee s.n.(C,UC);Cienegita,SEofBadiraguato, Gentry 5939(ARIZ);Choix, Gentry 6779(GH,MICH,NY).—soNorA:ElCoyote,inE-centralSonora, Felger 3616(ARIZ,ENCB,SD);�9.8mibyrdEofmainhwy(Mex.Hwy�5)onrdtoUres, Felger 3912(MEXU);ArroyoGuajaray,RíoMayo, Gentry 1137(F,GH,K,MEXU,MO,S,UC,WIS);SanBernardo,RíoMayo, Gentry 2282(GH,MO);Mpio.Yecora,Curea,28°�8'42"N,�09°�6'42"W,Reina G. et al. 98-1261(MICH);nearintersectionofHwy�5androadthroughSuhuraltowardSanAgustín, Stevens 1518(ENCB);CañóndelasBellotas,regionofRíodeBavispe,NESonora, White 3574(ARIZ,BR,GH,MEXU,MICH);�3miSofDivisaderos, Wiggins 7464D(DS,US);�4miSofDivisaderos, Wiggins 7471(DS,MICH,US).

Galphimia vestitaiseasilyrecognizedbyitslinearanddenselypubescentleaves.TheonlyotherspecieswithabundantvestureonnearlyallitspartsisG. hirsutaofGuerrero,whichhasbroadlaminas;itsmuchlargerflowersareborneonpedicelssubtendedbypedunclesandhaveunequalpetals.TheandroeciumofG. vestita isuniqueinthegenusinthatthestamensaresubequal;inallotherspeciesatleastsomestamenshavefilamentsdifferinginlength.Galphimia vestitaissometimesmisidenti-fiedasG. angustifolia,whichalsooccursinnorthernMexicoandinsomepopulationshaslinearlaminas,butitsleavesareglabrousorbearonlyafewscatteredhairs.

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


Galphimia brasiliensis var. pubescens f. ovata Nied., Arbeiten Bot. Inst. Königl.Lyceum Hosianum Braunsberg 5: 22. �9�4.—type: brAziL. Bahia: withoutlocality,�830,Salzmann s.n.(lectotype,designatedbyC.Anderson,2005a:G!;isolectotypes:MO!P!).

Fig.�8.

Subshrubandshrubto�.5m,oftensprawlingandscrambling;stemspubescentwhenyoung,becomingglabrousbutoftenroughenedbytubercles.Vestureofallveg-etativepartsofstraighttowavy,lightbrowntowhitehairs,0.3–0.8mmlong,eachhairborneonapersistenttubercleto0.25mmhighortheepidermisonlyslightlyraisedatpointofattachment.Laminasofthelargerleaves2–5cmlong,�–3.8cmwide,ovateorelliptical,apexapiculateoracute,baseacutetotruncate,glabrousbutoftenwithafewscatteredhairsontheabaxialcostaandalongthemarginnearthebase,second-aryveinsprominulousorbarelysoabaxially;marginentireandcommonlywithafewpersistenttuberclesnearthebase;petioles0.3–�cmlong,tuberculate-strigoseoronlythetuberclesremaining;leafglandsusuallyapairborneonthemarginto0.6mmabovethebaseofthelamina,sometimeswithonly�glandortheglandsabsent,eachgland0.�–0.3mmindiameter,disklikeandprominentorsometimesdrawnoutintoatoothto0.3(–0.5)mmlong;stipules�.5–2.7mmlong,0.3–0.6wide,linearorverynarrowlytriangular,withscatteredhairsalongthemargin.Inflorescenceaterminalraceme,theaxespubescentandtuberculate;pedunclesabsentorrudimentaryandupto2(–3)mmlong,glabrate,pedicels2.5–5.5mmlong,pubescent;bracts�.2–2mm

FIG.�8.Galphimia brasiliensis.a.Branchwithinflorescence.b.Detailoflaminabase,showingmar-ginalgland.c.Leaf.d.Petal.e.Stamensoppositeposterior-lateralsepal(left)andposteriorpetal(right),abaxialview.f.Gynoecium.g.Fruit,subtendedbycalyxandfilaments,borneonpedicelandpedunclewithportionofinflorescenceaxis.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,MICH;b,c,Harley 16312,MICH;h,i,Blanchet 3904,G.)


long,0.4–0.5mmwide,linear,bracteoles0.5–0.8mmlong,0.3–0.4mmwide,linear,bracts and bracteoles glabrous. Sepals ca. 2.5 mm long, �–�.� mm wide, narrowlyelliptical,glabrousorwithscatteredhairsatthebase,usuallywithatuftofhairsattheapex;glandsabsent.Petalsdeciduous,subequalbuttheposteriorpetalsometimeswithaslightlywiderclaw,yellow,oftenmarkedwithred,becomingpink/redinage,glabrous,marginfinelydenticulate-erose;claw0.5–�mmlong,0.4–0.5mmwide,limbca.4mmlong,ca.3mmwide,triangular-ovate,apexobtuse,basegraduallyacute.Sta-mensofunequallength;filamentofstamenoppositeanteriorsepalca.3.5mmlong,filamentsofstamensoppositelateralsepals3.2–3.3mmlong,filamentsofstamensoppositepetalssubequal,ca.2mmlong;anthers0.7–0.8mmlong.Ovaryfinelypilose,thehairsca.0.05(–0.�)mmlong;styles3.3–4.3mmlong.Coccusca.3.6mmlong,ca.2mmindiameter,finelypilose;areoleca.2.6mmlong,ca.�.7mmwide;matureseednotseen.Chromosomenumberunknown.

Phenology.Collectedinflowerandfruitthroughouttheyear.Distribution (Fig. �9). Eastern Brazil (Bahia, Paraíba, Pernambuco, Piauí); in

caatinga,cerrado,atedgesofwoodsandgalleryforests,inthornscrub;280–��50m.

AdditioNAL specimeNs exAmiNed. Brazil.bAhiA:Mpio.Jussiape,estradaJussiape/Abaíra,ca.5kmdeJussiape,Amorim et al. 2833(MO);�0kmWofSerrinhaonrdtoConceiçãodoCoité,��°38'S,39°W,Anderson 11737 (CAS, MBM, MICH, NY);Valente, 6 km N ofValente on rd to Santaluz, 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);Blanchet 1046 (G);Blanchet 2184 (BM,G);SerradeJacobina,Blanchet 2674 (BM,BR,C,G,MO,NY,P,W);Blanchet 3904 (BM,F,G);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);LagoadeEugenia,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);Milagres,arre-dores,Hatschbach 42452 (MBM,MICH);Itaberaba,Fda.Morros,Hatschbach 48206 (BR,CEPEC,MBM,MICH,MO);MorrodaGaraffa,�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 3070 (CEPEC,MICH);beiRemonso,Ule 7180 (G,L);SantaLuz,30kmWofQueimadas,�5kmNWofSantaLuz,��°09'S,39°28'W,Webster 25670 (CAS,MICH).—pArAíbA:withoutlocality,Coêlho de Moraes 2116 (A,NY,S).—perNAmbuco:Sertânía,FazendaCoxi,Alencar 3 (MICH);Area-projetoSuape,CabodeSto.Agostinho,parteanterior(EstaçãoD),Andrade-Lima & Medeiros-Costa 48 (F,MBM);PraiadoGuaibo,35kmSofRecife,Deguchi et al. B-1739(MO);nearPernambuco,Gardner 944 (BM,GH,NY,S,W);20kmdePetrolinaemdireçãoaAfrânio,Heringer et al. 185 (UB);20kmdePetrolinaaAfrânio,Heringer et al. 975 (RB);Houllet s.n. (BR);Olinda,Pickel 676 (SP);Ridley et al. s.n. (BM);Alagoinha,SWvonCaruarú,Vogel 129 (MICH).—piAuí:LagoaComprida,Gardner 2077 (BM,GH).

Galphimia brasiliensis is characterized by its pubescence composed of hairsborneontinypersistenttubercles,mostlyovateleaves,andafinelypiloseovary.Theandroeciumisuniqueinthegenusinthatthefilamentsofstamensoppositethelater-alsepalsaresubequal,asarethoseofstamensoppositeallpetals.Inallotherspecies,exceptG. vestita,anterior-lateralfilamentsdifferfromtheposterior-lateralones.

Galphimia brasiliensis is restricted to eastern Brazil (Paraíba to Bahia andadjacentPiauí),buttraditionallythenamehasbeenappliedtoanyspecimenofGal-phimiafromSouthAmerica,especiallytothewidespreadG. australis,whichlackstuberclesandhasaglabrousovaryandfruit,a(usually)glandularcalyx,andamoresouthernrange.OccasionallyplantsofG. graciliscultivatedinSouthAmericaare


FIG.�9.DistributionofGalphimia amambayensis,G. brasiliensis, andG. platyphylla.

labeledas“Galphimia brasiliensis,”buttheyarereadilydistinguishedfromtrueG. brasiliensisbytheirlargeshowyflowersborneonpedicelssubtendedbypeduncles,whichbeartwobracteolesatorjustbelowtheapex.

Jussieu (�833) was the first to realize that Linnaeus’s Thryallis brasiliensis belongsinGalphimiaandisnotrelatedtospeciesassignedtoThryallisMart.,buthisdescriptionofG. brasiliensiswasbasedonspecimensofG. australis.

16. 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,designatedbyC.Anderson,2005a:G!;isolectotypes:K!P!;photoofPisolectotype:MICH!).

Galphimia brasiliensisvar.pubescensAdr.Juss.inSt.-Hil.,Fl.bras.merid.3:72.�833[“�832”].—type:brAziL.RioGrandedoSul:adripasUruguayjuxtavicumS.-FranciscodeBorja[SãoBorja],Feb,St.-Hilaire C2, 2495bis(holo-type:P!,photo:MICH!).


Galphimia australisf.angustifoliaChodat,Mém.Soc.Phys.Genève3�,part2,no.3:23.�892.Galphimia brasiliensis f.angustifolia(Chodat)Nied.,ArbeitenBot.Inst.Königl.LyceumHosianumBraunsberg5:22.�9�4.—Type:pArA-GuAy:Guairá:páturaged’Itangu,prèsdeVilla-Rica,�7Feb�876,Balansa 2394(holotype:G!).

Fig.20.

Subshrubor suffrutescentherb fromawoodycaudex, to�m; stems sparselypubescentwhenyoung,soonbecomingglabrous,orpubescentandonlytheoldestpartsglabrescent(RioGrandedoSulandUruguay).Vestureofallvegetativepartsofsessile,mostlywavyorcrispedbutsometimesstraight,reddishbrownhairs0.2–0.7mmlong.Laminasofthelargerleaves2.5–7(–8)cmlong,(0.5–)�–2.5(–3.3)cmwide,length/widthratio2.5–7(–9), linear to linear-lanceolate to lanceolate tonarrowlyelliptical,apexapiculate,baseacute,glabrous,secondaryveinsbarelyprominulousornotevident;marginentire;petioles(0.�–)0.2–0.6cmlong,glabrous;leafglandsborneonthemarginnearthebaseofthelaminaorupto0.8cmaboveit,usuallyapair,orsometimeswith�or2additionalglands,orsometimesonly�glandortheglandsabsent,eachgland0.2–0.5mmindiameter,circular,flushwiththemarginorprominentorsometimeswithastalkto0.5mmlong;stipules(�.5–)2–4.2(–5)mmlong,0.5–�mmwide,linear,glabrous.Inflorescenceaterminalraceme,theaxesof-tentomentuloseintheyoungestpart,usuallybecomingglabrateinage,orglabrousthroughout;pedunclesabsentorrudimentaryandupto�.5(–2.5)mmlong,pedicels2–5(–�0)mmlong,tomentuloseorwithsomescatteredhairsorsometimesglabrous;bracts�–5(–7)mmlong,0.4–�mmwide, linear,bracteoles0.5–�.5(–2)mmlong,0.2–0.5(–0.7)mmwide,linear,bractsandbracteolesglabrous.Sepals2.5–3.5(–4)mmlong,�–�.8mmwide,oblong,glabrousbutwithatuftofhairsattheapex(bestseeninbudandearlyanthesis);glands(0–)�–3(–5)percalyx,0.2–0.5mmindiameter,prominent.Petalsdeciduous,subequal,theposteriorpetalsometimesalittlelarger,yellow,becomingredinage,glabrous,marginfinelydenticulate-erose;claw�.3–�.5(–�.7)mmlong,ca.0.5mmwide,limb3–5mmlong,2.5–4mmwide,triangular-ovate,apexobtuse,baseacutetotruncate.Stamensofunequallength;filamentoppositeanteriorsepal2.5–3(–3.4)mmlong,filamentsoppositeanterior-lateralpetals2–2.3(–2.5) mm long, filaments opposite anterior-lateral sepals 2.3–2.5 (–3) mm long,filamentsoppositeposterior-lateralpetals�.8–2(–2.5)mmlong,filamentsoppositeposterior-lateralsepals2–2.3(–3)mmlong,filamentoppositeposteriorpetal(�.8–)2–2.2(–2.5)mmlong;anthers�–�.2mmlong.Ovaryglabrous;styles3.5–4.6(–5.2)mmlong.Coccus3–3.8mmlong,2–2.5mmindiameter,glabrous;areole2.2–3mmlong,�.6–2mmwide;outercotyledon�.8–2.7mmlong,innercotyledon�.5–2mmlong.Chromosomenumberunknown.

Phenology.Collectedinflowerandfruitthroughouttheyear.Distribution(Fig.2�).SouthernBrazil,adjacentBolivia,Paraguay,northeastern

Argentina, western Uruguay; open woodlands and grasslands, cerrado, caatinga,campolimpo,camposujo,matorral,andsecondarysites;90–900m.

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);Depto.SantoTomé,Ea.Timbó,Ayo.Ciriacoyruta40,Schinini et al. 23447(CTES,MICH).—eNtre ríos:Depto.Concordia,ParqueRivadavia,Burkart & Troncoso 27703(NY);Depto.Federación,SantaAna,barrancadelRíoUruguay,Burkart et al. 29359(CTES,GH);ConcepcióndelUruguay,Lorentz 563(BM,F,G,GH,K).—misioNes:9kmalNWdeConcepcióndelaSierra,Krapovickas et al. 15148(C,


FIG.20.Galphimia australis.a.Habit.b.Nodewithstipulesandbasalportionofleaves.c.Flowerbud;notecalyxglands.d.Flower,sideview(adaxialapexoflateralpetalenlarged).e.Distalportionsofstamens,abaxialview(left)andadaxialview(right).f.Gynoecium.g.Apexofstyle.h.Fruit.i.Cocci,abaxialview(left)andadaxialview(right).j.Seed.k.Embryo.Scalebar:a,3cm;b,6mm;c,3mm;d,3.8mm(3mm);e,�.8mm;f,3mm;g,0.3mm;h,i,3.8mm;j,k,2.�mm.(Basedon:a–g,Schinini 14017,MICH;h–k,Krapovickas & Schinini 32151,MICH.)


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,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.CampoGrande,RioAnhandui,Hatschbach 21813(MBM);Mpio.RioBrilhante,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);estradaSantoHipólitoaDia-mantina,Km69,Shepherd et al. 3837(MBM,NY).—pArANá:Ibaréopp.,Morungavapraedium,Dusén 17413(GH,MO,S);Km�27,Mpio.LaranjeirasdoSul,Hatschbach 23123(MBM,MICH).—rio GrANde

FIG.2�.DistributionofGalphimia australis.


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). paraguay.ALto pArANá:Ea.SantaElena,PiraPyta,Schinini & Caballero M. 27152(G,MICH).—AmAmbAy:32kmSEdeBellaVista,Ayo.Negla,Schinini 21495(F,G,MICH).—cAAGuAzú:ColoniaPindo,caminoentreItaquyryyCuruguati,24°35'S,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.Sofía,Rosengurtt B-3278 1/2(GH,MO,SP).

GalphimiaaustralisisavariablespeciesfoundinsouthernBrazil,Bolivia,Para-guay,andeasternArgentinaandadjacentUruguay.TraditionallyandinaccordancewithNiedenzu(�9�4,�928)ithasbeencalledG. brasiliensis,butthatnameappliestoaspeciesrestrictedtoeasternBrazil(seediscussionofG. brasiliensis,no.�5).Gal-phimiaaustralisissympatricwithG. platyphylla,whichhasverybroadlaminasandlargerflowers,andwithG. amabayensis,whichisreadilyseparatedbyitstomentulosecalyx. Ingeneralaspect, it ismost similar to theNorthAmerican G. angustifolia,whichhasapiloseovaryandfruit,andlackscalyxglands,butisalsocharacterizedbyvariationintheshapeofthelamina.MostpopulationsofG. australishavenarrowlaminas,rangingfromlineartolinear-lanceolatetolanceolatetonarrowlyelliptical,whichtapertowardtheapexandare3–6(–9)timesaslongaswide.Occasionally,somelaminasonaspecimenareonly2.5timesaslongaswide,e.g.,Schinini 14017andSchinini et al. 23447 fromArgentina.Chodat (�892)assignedspecimenswithverynarrowleavestohisformaangustifolia,basedonBalansa 2394fromParaguay.Insomepopulationssomeoftheleavesonashoot,orsometimesall,arewiderthanusualandapproachtheleafshapefoundinG. amambayensisandG. platyphylla(e.g.,Hassler 4524,Hatschbach 25071, 49160, Krapovickas 29878, Krapovickas & Cristóbal 25195),butthesespecimensmatchG. australisinallotheraspects.

AvariantfromRioGrandedoSul(Brazil),inwhichthevesturefoundonyoungshootsandinflorescenceaxesisretained,wasnamedbyJussieu(�833)andacceptedbyNiedenzu(�9�4,�928)asG. brasiliensis var.pubescens.Suchexceptionalpopula-tionswithpersistentvestureoccuroccasionallyinotherpartsoftherangeaswell(e.g.,Argentina:Lorentz 563,EntreRíos; Xifreda & Maldonado 381,Misiones)anddonotmeritformalrecognition.

Vegaetal.(2002)reportedthepresenceofarticulatedlacticifersandproductionoflatexinG. australis(as G. brasiliensis).

17. Galphimia amambayensis C. Anderson, Contr. Univ. Michigan Herb. 24: 2.2005.—type:pArAGuAy.Amambay:30kmalNdelcruceBellavistayPedroJuanCaballero,enmatorral,��Feb�982,Fernández Casas & Molero 6190(holotype:NY!;isotype:MO!). Fig.22.

Subshrubto0.6mhigh;stemstomentosewhenyoungbutbecomingglabrousinage.Vestureofsessile,crispedorcurled,reddishbrownhairs0.3–0.7mmlong,butstraightandappressedonthelaminas.Laminasofthelargerleaves4.5–6.5cmlong,


�.5–2.8cmwide,ellipticalornarrowlyso,apexapiculate,baseacute,adaxiallyandabaxiallysparselypubescentwhenyoung,mostlyglabrouswhenmaturebutsome-timeswithsomescatteredhairsremaining,secondaryveinsbarelyprominulousornotevident;marginentire;petioles0.3–0.6cmlong,glabrous;glandsapairborneonthemarginnearthebaseofthelaminaorto0.6mmaboveit,eachgland0.4–0.5mmindiameter,disklikeandflushwiththemargin;stipules4–4.5mmlong,0.7–�mmwide,linear,pubescentonmarginandwithscatteredhairsabaxially.Inflorescenceaterminalraceme,theaxesdenselytomentose,glabrescentinage;pedunclesabsentorrudimentary,pedicels4–6.5mmlong,denselytomentose;bracts2.5–3.5mmlong,0.7–0.8mmwide,linear,bracteoles�.2–�.5mmlong,ca.0.5mmwide,linear,bractsandbracteolesabaxially tomentuloseoronlyalong themargin.Sepals3.5–5mmlong,�.3–�.5mmwide,narrowlyelliptical,abaxiallytomentuloseorpatchilyso inage;glandsabsent.Petalsdeciduous,subequal,yellow,becomingredinage,abaxiallymostlywithscatteredhairsontheclawandonthelimbadjacenttothemidrib,orsometimesglabrous,marginveryfinelydenticulatetosubentire;clawca.2mmlong,ca.0.5mmwide,limb5–5.5mmlong,4–4.5mmwide,broadlytriangular,apexobtuse,basegraduallyacutetosubtruncate.Stamensofunequallength;filamentoppositeanterior sepal3.5–4mmlong,filamentsoppositeanterior-lateralpetals4–4.5mmlong,filamentsoppositeanterior-lateralsepals3.5–4mmlong,filamentsoppositepos-terior-lateralpetals4.5–5mmlong,filamentsoppositeposterior-lateralsepals3.8–4.3mmlong,filamentoppositeposteriorpetal4.8–5.2mmlong;anthersca.�.3mmlong.Ovarypilose,inthedistal�/3onlyonthesuturesandadjacenttothem,hairsca.0.�mmlong;styles5.5–6.5mmlong.Coccusca.4.5mmlong,ca.3mmindiameter,pilosebutglabrescentinage;areoleca.4mmlong,ca.2.5mmwide;outercotyledon3.2–3.4mmlong,innercotyledon2.5–3mmlong.Chromosomenumberunknown.

FIG.22.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,NY.)


Galphimia amambayensis is known only from the type collected in easternParaguay(Fig.�9).It isreadilyseparatedfromtheotherSouthAmericanspeciesbyitsdenselytomentoseinflorescenceaxesandcalyx.LikeG. andustifoliaandG. brasiliensis,ithasapiloseovary,butinthedistal�/3thevestureispresentonlyonthesuturesandadjacenttothem.Thepetalsbearscatteredhairsabaxiallythougharesometimesglabrous,butintheseinstancethehairsmayhavebrokenoff.

18. 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!,photo:MICH!;isotype:G!). Fig.23.

Subshrubto0.5mhigh, fromawoodycaudex;stemsglabrousexcept for theyoungest parts.Vesture of all vegetative parts of sessile, straight, reddish brownhairs0.2–0.7mmlong.Laminasofthelargerleaves3–7.5cmlong,(�.5–)2–4(–5)cmwide,length/widthratio�.2–2.3(–3),ellipticaltolanceolatetoovatetobroadlyovate, occasionally suborbicular, apex obtuse or acute or sometimes apiculate oremarginate,baseacutetotruncate,glabrous,secondaryveinsprominulousorbarelyso;marginentire;petioles0.�–0.6cmlong,glabrous;glandsusuallyapairborneonthemarginnearthebaseofthelaminaorupto0.5cmaboveit,orsometimeswith�or2additionalglands,orsometimesonly�glandortheglandsabsent,eachgland0.2–0.5mm indiameter,flushwith the surfaceor slightlyprominent; stipules2–4mm long, 0.6–�.2 mm wide, triangular to narrowly so, glabrous or with scatteredhairs along the margin. Inflorescence a terminal raceme, the axes with scatteredhairs and glabrescent; peduncles absent, pedicels �.5–8 mm long, 0.7–� mm wide,glabrous;bracts�.5–3mmlong,0.7–�.2mmwide,triangular,bracteoles0.7–�.2mmlong,0.4–0.7mmwide,triangular,bractsandbracteolesglabrous.Sepals4–5.7mmlong,2–2.5mmwide,ellipticalorsometimesoblong,glabrousbutwithatuftofhairsattheapex(bestseeninbudandearlyanthesis);glands(0–)�–5percalyx,glands0.3–0.6mmindiameter.Petalsdeciduous,subequal,theposteriorpetalsometimesalittlelarger,yellow,becomingredinage,glabrous,marginerose-denticulate;claw2–2.5mmlong,ca.0.5mmwide, limb7–8.5mmlong, (4–)5–5.5mmwide,ovate,apexobtuse,baseacuteortruncate.Stamensofunequallength;filamentoppositeanterior sepal 3.3–4.5 mm long, filaments opposite anterior-lateral petals 2.7–3.5mmlong,filamentsoppositeanterior-lateralsepals3–4mmlong,filamentsoppositeposterior-lateralpetals2.5–3.3mmlong,filamentsoppositeposterior-lateralsepals2.8–3.7mmlong,filamentoppositeposteriorpetal2.5–3.5mmlong;anthers�–�.2mmlong.Ovaryglabrous;styles4.2–5.3mmlong.Coccus4–5mmlong,3–3.7mmindiameter,glabrous;areole3–3.5mmlong,2.5–3mmwide;matureseednotseen.Chromosomenumberunknown.

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

Distribution(Fig.�9).EasternParaguay(Amambay,Caaguazú,Canendiyú)andadjacentBrazil(MatoGrossodoSul);campo,cerrado,campolimpio,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);


FIG.23.Galphimia platyphylla.a.Habit.b.Baseoflamina,showingmarginalglands.c.Oldflower(thepetalsfallen)borneonpedicelwithportionofinflorescenceaxis;notecalyxglands.d.Petal.e.Sta-mens,abaxialview,oppositeposterior-lateralsepal(left)andposteriorpetal(right).f.Gynoecium.g.Cocci,adaxialview(left)andlateralview(right).Scalebar:a,4cm;b,8mm;c,d,5mm;e,f,2.7mm;g,4mm.(Basedon:a–c,Hassler 9132,G;d–f,Pedersen 14726,MICH;g,Krapovickas et al. 45943,MICH.)

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,ApepúetTajaira-guay[?],Hassler 4327(BM,F,G,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 platyphylla ischaracterizedbythickandbroadlaminas,whichvaryfrombroadlyellipticalorbroadlylanceolatetobroadlyovateorevensuborbicular;theyareonly�.2–2.3 timesas longaswideandoftenhaveanobtuseapex.Inallotherspecieswithrudimentarypedunclesusuallyonly�–3flowersareopenatonetime,butinG. platyphyllaaracememaybearuptosevenopenflowers(e.g.,Hassler 9132).ThisspecieshassometimesbeenincludedinG. australis,whichhaslineartolanceolateornarrowlyelliptical laminasof thinner texture.Also, theflowersandfruitsofG. platyphyllaarelarger,andtheplantsareapparentlyshorter(to0.5mvs.toca.�mtall)althoughmorerobust,i.e.,theaxestendtobestouter.ShootsofG. platyphyllagenerallyarecomposedofthreetofourinternodesandhaveaterminalinflorescence;sometimeslateralshootsdevelopandthesefollowthesamepattern.Galphimia platyphyllaisalsosympatricwithG.amambayensis,whichiseasilydistin-guishedbyitstomentoseinflorescenceaxesandcalyx.


Group iii (petalspersistent;pedunclespresent)

19. Galphimia glaucaCav., Icon.5:6�,pl.489.May�799.Malpighia glauca (Cav.)Pers.,Syn.pl.�:506.�805[combinationalsoproposedbyPoiretinLamarck,Encycl.Suppl.4:7.�8�6].Thryallis glauca(Cav.)Kuntze,Rev.gen.pl.�:89.�89�.—type:basedonplantscultivatedatthebotanicalgardeninMadridinOctober,�794;provenanceofpropagules:“[mexico:Guanjuato:]AcambaroetSalvatierra,”Née s.n.(holotype:MA-475692,microfiche:MICH!,photo:MO!). Fig.24.

Shrubto2(–3)mtall;stemssericeouswhenyoung,soonglabrous.Vestureofallvegetativepartsofsubsessile,mostlystraightorwavy,reddishbrownhairs0.2–0.6mmlong.Laminasofthelargerleaves(�.5–)2–5.6cmlong,(0.7)�.3–3.5cmwide,ellipticaltolanceolateorovate,rarelysuborbicular,apexapiculate,baseacute,gla-brous,secondaryveinsprominulousabaxially;marginentire;petioles0.4–�.5cmlong,sparselysericeoustosoonglabrous;glandsusuallyapairborneonthemarginwellabovethebaseofthelamina,sometimesnearerthebase,eachgland0.4–0.7mmindiameter,disklikeandsessile,orsometimespeglikeandupto0.5mmlong,occasion-allywithadditionaltinyglandsalongthemarginintheproximalhalf;stipules(�.4–)2–3mmlong,0.5–�mmwide,linearornarrowlytriangular,glabrous,withscatteredhairsonthemargin.Inflorescenceaterminalraceme,theaxesdenselytomentulose;peduncles3–7.5mmlong,pedicels(6–)8–�2mmlong,bothtomentulose,peduncles0.3–0.8(–0.9)timesaslongaspedicels;bracts2–3(–4)mmlong,0.5–0.8mmwide,linear,bracteoles�.5–2.5mmlong,0.4–0.5mmwide,linear,bractsandbracteolesgla-brousorwithscatteredhairsonthemargin;bracteolesborneinthedistal�/5–�/4ofthepeduncle,suboppositeorupto�mmapart.Sepals3–4mmlong,�.3–2mmwide,lineartooblongorelliptical,glabrousorwithafewscatteredhairsatbaseabaxially,marginirregularlydenticulate-erose-ciliate;glandsabsent.Petalspersistent,unequal,yellow,sometimesmarkedwithredinage,glabrous,marginfinelydenticulate;lateralpetals:claw�.5–2mmlong,0.5mmwide,limb7.5–9.5mmlong,5–6.5mmwide,ellip-ticaltolanceolate,apexobtusetosubacute,basegraduallyattenuate;posteriorpetal:claw2.8–4mmlong,�.3–�.7mmwide,limb7.5–9.5mmlong,7–8.5mmwide,broadlytriangular,apexrounded,baseslightlycordate.Stamensofunequallength;filamentoppositeanteriorsepal4–6mmlong,filamentsoppositeanterior-lateralpetals3–5mmlong,filamentsoppositeanterior-lateralsepals4–7mmlong,filamentsoppositeposterior-lateral petals 2.2–4 mm long, filaments opposite posterior-lateral sepals3.8–5.5mmlong,filamentoppositeposteriorpetal2–3.5mmlong;anthers2.4–3mmlong.Ovaryglabrous;styles5.5–6.7mmlong.Coccus2.5mmlong,2mmindiameter,glabrous;areole2mmlong,�.8mmwide;outercotyledon�.8–2mmlong,innercoty-ledon�.3–�.5mmlong.Chromosomenumberunknown.

Phenology.CollectedinflowerfromMaythroughNovember,infruitfromMaythroughJanuary.

Distribution (Fig. 25). Mexico (Aguascalientes, Guanajuato, Hidalgo, Jalisco,NuevoLeón,Querétaro,SanLuisPotosí,Tamaulipas,Zacatecas);indeciduousforest,pineforest,pine-oakforest,matorral,andatroadsides;700–2300m.

represeNtAtive specimeNs. mexico. AGuAscALieNtes: Mpio. Calvillo, Los Lobos, �7 km al S deMalpaso,de la Cerda & García R. 885(ENCB);Mpio.Calvillo,RíoGil,García R. 2705(IEB);withoutlocality, Hartweg 13(BM,G,GH,K,L,NY,W).—GuANAJuAto:DoloresHidalgo,Hernández X. et al. X-2403(MEXU);�3kmalWdeXichú,sobrelacarr.aSanLuisdelaPaz,Rzedowski 41562(ENCB,IEB);Mpio.Atarjea,CasasViejas,8kmalSdeLaJoya,Ventura & López 6516(ENCB,IEB,MEXU,MICH);


FIG.24.Galphimia glauca.a.Floweringbranch,withdetachedleaftoright.b,c.Baseoflaminastoshowmarginalglands.d.Openingflower,borneonpedicelandpedunclewithportionofinflorescenceaxis.e.Lateralpetal.f.Posteriorpetal.g.Androeciumlaidout,abaxialview,thestamenonrightoppositeposteriorpetal.h.Gynoecium.i.Cocci,adaxialview(left)andlateralview(right).j.Seed.k.Embryo.Scalebar:a,4cm;b,d–f,8mm;c,g,h,4mm;i–k,2.7mm.(Basedon:a,d–h,Ventura & López 6919,MICH;b,c,González 970,IEB;i–k,Ventura & López 6516,MICH.)


Lagunilla,�5kmalNdeVictoria,Ventura & López 6919(CHAPA,GUADA,IEB,MICH);LaPresa,8kmalNdeDoctorMora,Ventura & López 7048(GUADA,IEB,MICH);Mpio.Victoria,RancholaLuz,Ventura & López 8481(IEB,MICH,XAL).—hidALGo: 2.3miNEofZimapán,Mex.Hwy85,Anderson & Laskowski 4027(DUKE,ENCB,GH,MICH,NY,US);Mpio.Jacala,Chase 7281(F,GH,MICH,MO);SofPachuca,Clark 7361(MO,NY);Mpio.Zimapán,3kmalNEdeZimapán,carr.aJacala,García P. 1039(CAS,CHAPA,ENCB,GUADA,IBUG,LL,MEXU,MICH,TEX,XAL);40miNNWofIxmiquil-pan,Hitchcock & Stanford 7004(DS,GH,NY,UC,US);Mpio.Ixmiquilpan,barrancadeTolantongo,40kmalNEdeIxmiquilpan,Tenorio 513(ENCB,UAMIZ,XAL).—JALisco:descentfromMesadeCristoto Huejuquilla elAlto, �8 mi by rd SW ofValparaíso, Anderson & Laskowski 3617 (DUKE, ENCB,GH,MICH,US);Salcillo,Jones 20(US).—Nuevo LeóN:Mpio.Zaragoza,Potreritos,Hinton et al. 24313 (MEXU,NY,TEX);nearElCaracol,3miNEofDulcesNombres,24°N,99°30'–�00°30'W,Meyer & Rog-ers 2532(BM,BR,G,GH,MO).—QuerétAro: alongHwy�20betweenCadereytaandVizarrón,3.5miSofVizarrón,20°50'N,99°45'W,Daniel 378(CAS,CHAPA,MICH);6.5miWofColón,Denton 1996(ENCB,MICH,UC);Vizarrón,Zamudio R. 2318(GUADA,IEB,MICH,MO,TEX);Mpio.Jalpan,4–5kmalSdeLaParada,Servín 268(IEB,MICH);Mpio.Landa,PuertodelMalpaís,3.5kmalNWdeElMadroño,González 970(IEB);Mpio.Peñamiller,MagüeyVerde�7kmalSEdePinaldeAmoles,carr.PinaldeAmoles–Peñamiller,Tenorio L. 2337(MEXU,MO);Mpio.PinaldeAmoles,�kmalNEdeElLlano,Carranza 679(ENCB,IBUG,IEB,MEXU);Mpio.Tolimán,RanchoMagüeyManso,Zamudio R. 2203(IEB).—sAN Luis potosí:Km53,nearLasRusias,ca.4miEofSanFrancisco,22°04'N,�00°33'W,Moran 6339 (MEXU, SD); Km 54 carr. San Luis Potosí–Ríoverde, Rzedowski 8057 (ENCB, MEXU,MICH,TEX).—tAmAuLipAs: arroyoalongHwy�0�, 0.2miNofLaPresita, 23°�5'N,99°40'W,Daniel 286(CAS,ENCB,MICH);Mpio.Jaumave,AlvarezyUrbina,29kmalNdeMagdalenoAguilar(San-tiaguillo),González M. 9731(MEXU,MICH);Mpio.Bustamente,caminodeFelipeAngelesalOjodeAguaSanJosé,Hernández 1608(ENCB,MO,TEX);Mpio.Hidalgo,LosCaballos–Potreritos,Hinton et al. 24875(TEX);Mpio.Tula,LaPresita,Km66carr.Tula–Victoria,Martínez 1224(ENCB,MEXU,MO);Mpio.VilladeCasas,rdfromRancho“LasYucas”toSantaMaríadelosNogales,Martínez M. & Borja

FIG.25.DistributionofGalphimia elegans, G. glauca, G. mexiae,andG. radialis.


L. F-1972(TEX);Mpio.GómezFarías,caminodeJulioaJoyadeSalas,Puig 7173(ENCB);6kmWofMiquihuana,23°42'N,99°45'W,Stanford et al. 721(DS,GH,MO,NY);3kmNofHuisachal,30kmfromVictoria,Stanford et al. 2031(DS,G,UC,US).—zAcAtecAs:ridge2miNofPotrerodelasYeguas,�3miSofValparaíso,McVaugh 17773(ENCB,MICH).

Galphimia glaucaisashruborsuffrutescentherbofnortheasternandcentralMexico.Theflowersarearrangedinracemesterminatingbranchesbearingrathersmallleaveswithellipticaltoovatelaminas.Theleafglandsareborneonthemarginofthelaminawellabovethebase(Fig.24a–c);theglandsvaryfromflushwiththemargintoprominentorsometimespeglike.Theposteriorpetalhasabroadlytriangu-larlimbthatcontrastswiththeellipticaltolanceolatelimbofthelateralpetals.Thecocciarethesmallestinthegenus,onlyca.2.5mmlongandca.2mmindiameter.

ThenameGalphimia glaucahasbeenwidelyandindiscriminatelyappliedtoallMexicanspeciesofGalphimia withpersistentpetals,aswellastosomespecieswithdeciduouspetals,inpartbecausetheearlyintroductionsofGalphimiatohorticulturewereallsolabeledandthenwidelydistributed;forexample,Maund’s(�837)accountandillustrationof“G. glauca”appliestoG. gracilis,andBartling’s(�840)treatmentof“G. glauca”appliestoG. multicaulis.MostnotablythecommonlycultivatedG. graci-lis,whichhasdeciduouspetals,continuestobeidentifiederroneouslyasG. glaucainthenurserytradeandinbotanicalgardens.Imagesof“G. glauca”inhorticulturalpublicationsandonwebsitesarealmostalwaysofG. gracilis.ThatspeciesisnativetoeasternMexicoandispartlysympatricwithG. glauca,fromwhichitdiffersinaddi-tiontothepetalcharactersinitslargecocci(4.5–5mmlong),bracteolesplacedatorjustbelowtheapexofthepeduncle,andhairsoftenborneontubercles.

The Cavanilles Herbarium, housed at MA, has two sheets of G. glauca, MA47569�andMA475692.AsnotedbyGarilleti(�993),thesheetannotatedbyCava-nilles (MA 475692) is the holotype; the other sheet (MA 47569�) should not beconsideredtypematerial.

20. Galphimia elegansBaillon,Hist.pl.5:43�,figs.429–435.�874.—type:specimentakenfromplantscultivatedattheJardindesPlantesinParis(holotype:P!,photo:MICH!).

Galphimiahirsutaf.virescensK.Koch,BerlinerAllg.Gartenzeitung�857(50):394,t.9.�857.—type:basedonalivingplantcultivatedatthebotanicalgar-deninBerlin;lectotype,heredesignated:theillustrationintheprotologue.

Fig.26.

Shrubto3(–5)m;stemsstrigosewhenyoung,soonbecomingglabrous.Vestureofallvegetativepartsofsubsessile,crisped,reddishbrownhairs(0.�–)0.2–0.8mmlong.Laminasofthelargerleaves3.5–6cmlong,�.5–3.5cmwide,narrowlyellipti-caltolanceolatetoovatetorhombic,apexacuteorapiculate,baseacute,adaxiallysparselytomentulosetoglabrescentinage,abaxiallytomentulose,becomingpatchilytosparselytomentulose,eventuallyglabrescent,secondaryveinsprominentabaxially;marginentire;petioles0.8–2cmlong,tomentulosetoglabrescent;glandsusuallyapairplacedadjacentorupto2mmapartatthemiddleofpetioleorsometimesneartheapexofpetiole(especiallyinsmall/youngleaves),sometimeswithonly�glandortheglandsabsent,eachgland0.4–�.2mmindiameter,disklikeandveryprominent,orpeglikeandupto�.5mmlong;stipules�.5–2mmlong,0.7–�.2mmwide,linearornarrowlytriangular,glabrous.Inflorescenceaterminalracemeorsometimesasmallpanicle,theaxestomentosebutpatchilysoinage;peduncles3.5–6(–8)mmlong,pedicels7–�0.5mmlong,bothtomentulose,peduncles0.3–0.7(–�)timesaslongas


pedicels;bracts�.5–3mmlong,0.4–0.8mmwide,linear,bracteoles�–�.7mmlong,0.3–0.5 mm wide, linear, bracts and bracteoles tomentulose abaxially; bracteolesborneatmiddleofthepeduncle,subopposite.Sepals3–3.5mmlong,�.5–2mmwide,oblongorelliptical,withscatteredhairs,marginirregularlydenticulateanddistallyciliate;glandsabsent.Petalspersistent,unequal,yellow,glabrous,margindenticulate;lateralpetals:claw�.8–3mmlong,0.5mmwide,limb7–9mmlong,4.5–5.2mmwide,lanceolate,apexobtuse,baseacute;posteriorpetal:claw2–3.5mmlong,ca.�mmwide,limb7–9mmlong,5–6mmwide,broadlyovate,apexobtusetorounded,basetruncateorslightlycordate.Stamensofunequallength;filamentoppositeanteriorsepal5–5.8mmlong,filamentsoppositeanterior-lateralpetals4–4.3mmlong,fila-mentsoppositeanterior-lateralsepals4.5–4.7mmlong,filamentsoppositeposterior-lateralpetals3–3.5mmlong,filamentsoppositeposterior-lateralsepals5–5.5mmlong,filamentoppositeposteriorpetal3–3.2mmlong;anthers�.8–2mmlong.Ovaryglabrous;styles6.2–7mmlong.Coccusca.3mmlong,ca.2.5mmindiameter,smooth,glabrous;areoleca.2.7mmlong,ca.2.2mmwide;matureseednotseen.Chromo-somenumberunknown.

Phenology.CollectedinflowerandfruitfromJulythroughNovember.Distribution (Fig. 25).Mexico (Oaxaca);matorral,pine-oak forest,deciduous

forest;�740–2200m.

AdditioNAL specimeNs exAmiNed.mexico.oAxAcA:�.5kmNofIxtlándeJuárez, junctiononrdtoValleNacional,Breedlove 60033(CAS);cercaníasdeOaxaca,Conzatti 1373(US);CerroSanFelipe,Dtto.Centro,Conzatti 2283 (F,GH,MEXU);without locality,�859,Cuming s.n. (G,P);alongroadto

FIG.26.Galphimia elegans.a.Floweringbranch.b.Largeleaf.c.Detailofpetioletoshowglands.d.Openingflower,borneonpedicelandpedunclewithportionofinflorescenceaxis.e.Petals,lateral(right)andposterior(left).f.Partialandroeciumlaidout,abaxialview,thestamenonrightoppositeposteriorpetal.g.Gynoecium.h.Cocci,adaxialview(left)andlateralview(right).Scalebar:a,b,4cm;c,f,g,4mm;d,e,8mm;h,2.7mm.(Basedon:a,c–g,Lorence 4734,MEXU;b,Conzatti 2283,MEXU;h,Saynes V. 29,ENCB.)


MicroondasNuevasPuntas,7.7miSEjct.rdtoMitlaalongHwy�90,Daniel 1271(MICH);withoutlocal-ity,�842,Franco s.n.(G);Yavezia,Sep�844,Galeotti 4335(BR,G,K,MEXU,P,W);Yodonocuito,5kmalEdeTeposcolula,Dtto.TeposcolulaMixtecaAlta,García M. 1044(ENCB,MEXU,MO);withoutlocality,Karwinski s.n.(G);Mpio.Ixtlán,SierradeJuárez,Ruta�75a0.5kmalNdeIxtlán,Lorence 4734(MEXU,MICH);PlayadeRíoCulebra,Macuiltianguis,Lucero L-92(ENCB,MEXU,XAL);MonteAlbán,nearOaxaca,Pringle 4873(A,BM,BR,G,GH,GOET,MEL,MEXU,MO,NY,S,UC,US,W);Mpio.Oaxaca,CerrodeSanFelipedelAgua,Saynes V. 29(CHAPA,ENCB,GUADA,IEB,MEXU,MO);Mpio.Oaxaca,CerrodeDonaji,Saynes V. 852(IEB,NY,XAL);Jayacatlán,Smith 83(GH);SierradeSanFelipe,Smith 647(NY);MonteAlbán,nearOaxaca,Smith 648(MO,NY,UC,US).

Galphimia elegansistheonlyspecieswithpersistentpetalsthathaspubescentleaves.Youngleavesandthoseneartheinflorescenceareabaxiallysparselytoabun-dantlytomentulose;thevestureiscomposedofreddishhairs,whichcommonlyaresloughedoffeventually inage.In larger leavestheglandsareborneataboutthemiddleof thepetioleandareveryprominentorpeglike(Fig.26c);onveryshortpetiolestheglandsareclosetotheapex.Galphimia elegansisalsodistinctiveinitsshortanthers(�.8–2mmlong)anddistallyciliatesepals.ThespeciesisendemictoOaxacaandsympatricwiththevariableG. speciosaandG. multicaulis,whichhavealways glabrous leaves with marginal glands as well as larger anthers (2.2–3 mmlong).Inaddition,theleavesofG. multicaulishaverudimentarytoveryshortpetioles[0.3–0.8(–�)mmlong].

AlthoughBaillondescribedG. elegansin�874,theidentityofthisspeciesanditsdistinctnesshavegonemostlyunrecognized.ThenameG. eleganswasoverlookedandisnotlistedinIndexKewensisandtheGrayIndex;yet,Baillon’sbriefdescrip-tion and illustrations fulfill the requirement for valid publication, and there is aholotypeatP.Thespecieswasintroducedtobotanicalgardensinthe�9thcenturywith thename“G. glauca,” andcollectionsare still commonly somisdetermined.Earlierbotanistswhonoted thepresenceof leafvesturemisapplied thenameG.hirsutatogardenplantsandherbariumspecimensofG. elegans(e.g.,Bartling�840;Neumann�847).Jussieu(�843)citedGaleotti 4335,acollectionofG. eleganswithwidelydistributedduplicates,asanexampleofG. hirsuta,whichhelpedtoperpetuatetheerror(e.g.,Koch�857;Rose�909;Niedenzu�9�4,�928).OccasionallyspecimensofG. elegansinbotanicalgardensandinthehorticulturaltradewerealsolabeled“Galphimiamollis”;thenameGalphimiamollisH.B.K.isasynonymofTetrapterys cotoneasterAdr.Juss.

21. Galphimia speciosaC.Anderson,sp.nov.—type: mexico.Chiapas:2.5kmNWofComitánonrdtoSanCristóbaldelasCasas,brushycut-overroadside,�820m,27Oct�988,Anderson 13555(holotype:MICH!;isotypes:CHAPA!DUKE!IEB!MBM!MO!NY!). Fig.27.

Frutexvelarborparva.Laminaefoliorummajorum2.5–7cmlongae,�.5–4cmlatae,ellipticaevellanceolataevelovatae,glabrae,margineintegra,glandulis0.5–�.5mm diametro, sessilibus vel pedicellatis; petioli 0.8–2.5 cm longi. Inflorescentiaracemosa, axibus tomentulosis vel sparse tomentulosis. Petala persistentia; petalalateraliaungue(�.6–)2–2.5mmlongo,limbo7–9(–�0)mmlongo,5–6(–7)mmlato,oblongovelovato;petalumposticumungue2.5–3.5mmlongo,limbo7–8(–9.5)mmlongo,6–7.8mmlato,triangularivelsuborbiculari.Antherae2.3–2.5(–3)mmlongae.Ovariumglabrum;styli5.5–6mmlongi.Coccus3.5–4mmlongus,glaber.

Shruborsometimesatreeletto3(–6)mtall;stemsstrigosetotomentulosewhenyoung,soonglabrescenttoglabrous.Vestureofallvegetativepartsofsubsessile,mostlystraightorwavy,reddishbrownhairs0.2–0.6(–0.8)mmlong.Laminasofthelarger


FIG.27.Galphimia speciosa. a.Floweringbranch.b.Detachedlargeleaf.c.Variationinpositionofglandsonpetioleandbaseoflamina.d.Flowersubtendedbypedicelandpedunclewithportionofinflo-rescenceaxis.e.Pedicel-pedunclejointwithbracteolesonpeduncle.f.Lateralpetals.g.Posteriorpetal.h.Partialandroeciumlaidout,abaxialview,thestamenonleftoppositeposteriorpetal.i.Gynoecium.j.Cocci,lateralview(left)andadaxialview(right).k.Embryo.Scalebar:a,b,4cm;c,h–j,4mm;d,�cm;e,2mm;f,g,8mm;k,2.7mm.(Basedon:a–i,Anderson 13555,MICH;j,k,Prigge 3237,MICH.)

leaves2.5–7cmlong,�.5–4cmwide,ellipticaltolanceolateorovate,rarelysubor-bicular,apexacutetoapiculate,baseacute,glabrous,secondaryveinsprominulousabaxially;marginentire;petioles0.8–2.5cmlong,withscatteredhairsorglabrous;glandsusuallyapairborneatthebaseofthelaminaorattheapexofthepetioleorondecurrentlaminartissueinthedistal�/3–�/2ofthepetiole,sometimeswithonly�glandortheglandsabsent,eachgland0.3–0.8mmindiameter,disklikeandprominent,orsometimespeglikeandupto0.5mmlong;stipules�.5–2.5(–3)mmlong,0.5–�mmwide,linearornarrowlytriangular,withscatteredhairsalongthemargin.


Inflorescence a terminal raceme, in vigorous plants 3 racemes sometimes aggre-gatedinaleafyternatepanicle,theaxestomentulose,sparselysoinage;peduncles(2.5–)3–6(–7.5)mmlong,pedicels(6.5–)8–�3mmlong,bothtomentulose,peduncles0.2–0.5timesaslongaspedicels;bracts�.5–2.8mmlong,0.4–0.8mmwide,lineartonarrowlytriangular,bracteoles�–2(–2.5)mmlong,0.3–0.5mmwide,linear,bractsandbracteolesglabrousorwithscatteredhairsalongthemargin;bracteolesborneattheapexofthepeduncleorinthedistal�/4–�/2,suboppositeorupto0.5mmapart.Sepals2.5–3.8mmlong,�.5–2mmwide,oblong,glabrous,marginirregularlydenticulate-eroseandoftendistallyciliate;glandsabsent.Petalspersistent,unequal,yellowandoftenmarkedwithredespeciallyontheclawandmidrib,glabrous,marginfinelydenticulatetosubentire;lateralpetals:claw(�.6–)2–2.5mmlong,0.5mmwide,limb7–9(–�0)mmlong,5–6(–7)mmwide,oblongorovate,apexbroadlyrounded,basetruncateorverybrieflyacute;posteriorpetal:claw2.5–3.5mmlong,�–�.2mmwide,limb7–8(–9.5)mmlong,6–7.8mmwide,broadlytriangulartosuborbicular,apexbroadlyrounded,basetruncate.Stamensofunequallength;filamentoppositeanterior sepal4–5.5mmlong,filamentsoppositeanterior-lateralpetals3–4.5mmlong,filamentsoppositeanterior-lateralsepals3.8–5.3mmlong,filamentsoppositeposterior-lateralpetals2.2–3(–4)mmlong,filamentsoppositeposterior-lateralsepals3.5–5mmlong,filamentoppositeposteriorpetal2.2–3(–4)mmlong;anthers2.3–2.5(–3)mmlong.Ovaryglabrous;styles5.5–6mmlong.Coccus3.5–4mmlong,2.7–3mmindiameter,glabrous;areole2.5–3mmlong,2.3–2.8mmwide;outercotyledon3.2–3.5mmlong,innercotyledon�.5–�mmlong.Chromosomenumber:n=6(Kyhos�966;W.R.Anderson�993;bothas“G. glauca”).

Phenology.CollectedinflowerandfruitthroughouttheyearbutmostcommonlyfromJunethroughDecember.

Distribution(Fig.28).SouthernMexico (southernPueblaandadjacentVera-cruz, Oaxaca, Chiapas) and CentralAmerica (Guatemala, Honduras, Nicaragua);insubtropicaldeciduousforest,pine-oakforest,matorral,andatroadsides;(��0–)800–2400m.

FIG.28.DistributionofGalphimia speciosa.


represeNtAtive specimeNs.mexico.chiApAs: 8.7–9.0miNofTuxtlaGutiérrezonrdtoSumidero,Anderson 4229(DUKE,ENCB,GH,MICH,MO,NY,US);5kmNofSoyaló,rdtoBochil,Breedlove 21324(CHAPA,DS,F,LL,MEXU,MICH,MO,NY);Mpio.Totolapa,6–8kmWofTeopiscaonsideofCerroChenek'ultik,Breedlove 27081(CHAPA,DS,MEXU,MICH,MO);Mpio.Berriozábal,�.3–2.6kmNofBerriozábaltowardLaCabaña,�6°48'N,93°�7'W,Daniel 8369(MICH);Mpio.Escuintla,Mt.Ovando,Matuda 16240(MEXU,MICH,MO,US);Mpio.Escuintla,Buenavista,Matuda 1878(DS,F,GH,MEXU,MICH,NY,US);Miramar,Matuda 395(LL,MEXU,MICH,US).—oAxAcA: 97kmSofTeotitlánonrdtoOaxaca,Anderson 12991(CAS,DUKE,IEB,MBM,MEXU,MICH,MO,NY);20kmSEofMitla,Con-rad 3044(GH,MO,SC,XAL);�kmalSdeMagdalenaJicotlán,Dtto.Coixtlahuaca,García M. 1923(F,MEXU,MICH,MO,UAMIZ,XAL);2kmWofTutlaonrdtoTamazulapanalongPan-AmericanHwy,Km382,Iltis 1333(F,K,MEXU,MICH,SP,US,WIS);nearTamazulapan,Nelson 1957(GH,US);CerroSolo,ca.7kmNEofTepelmemedeMorelos,Prigge 3237(ENCB,MEXU,MICH);Mpio.Albarradas,4kmalNEdeSanBártoloAlbarradas,caminoaZacatepec,Rzedowski 37038(ENCB,IEB,MEXU);5kmalNEdeTeotongo,porlaterraceríaaTepelmeme,�7°46'N,97°30'W,Salinas T. F-3507(CHAPA,F,MEXU,MO);2kmalNdeCuyotepeji,carreteraaTehuacán,�7°57'N,97°4�'W,Salinas T. F-3720(IEB,MEXU,WIS);Mpio.ConcepciónBuenavista,CerroLaCulebra,alSWdeElEnebro,Tenorio L. 7143(MEXU,MO,TEX).—puebLA: SantiagoMiahuatlán,�8°50'N,Cházaro B. 652(MEXU,XAL);ca.�2kmNEdeTehuacán,porlacarr.aEsperanza,Chiang F-270(CHAPA,IEB,MEXU,MO);vicinityofSanLuisTul-titlanapa,SierradeMixteca,Purpus 2733(BM,F,G,GH,MO,NY,UC,US);Mpio.Caltepec,LaLaguna,faldasdelCerroElGavilán,2kmalEdeCaltepec,�8°�0'N,93°38'E,Salinas F-3452(F,IEB,MEXU,MICH,WIS); 2.5–3.5 mi NW of Coalcoapan, �8°34'N, 97°40'W, Webster 17239 (DUKE, GH, MEXU,MICH,MO).—verAcruz:Acultzingo,Güiluapa,Ventura A. 15419(ENCB,IEB,MEXU,MO,XAL).

Guatemala.ALtA verApAz: Rd to San Juan Chamelco, Molina R. 12285 (F, LL, NY, US).—chi-mALteNANGo:rdfromChimaltenangotoSanMartínJilotepeque,Standley 57864(A,ECON,F,MICH,NY,US).—huehueteNANGo:entreNaranjalesyBoquerón,carreterahaciaLaMesilla,Castillo 1373(MICH,NY);SanMiguelAcatán,Skutch 999(A,BM,F,G,NY,US);CanyonofRíoSeligua,40kmNWofHue-huetenango,Williams 41147(BM,F,NY,US).—GuAtemALA: Amatitlán,Ruano 834(US).—Quiché:SanMiguelUspantán,Heyde 3272 (US).—sAcAtepéQuez:VolcánAcatenango,Smith 2548 (GH,US);nearPastores,Standley 59913(F);alongRíoGuacalate,rdbetweenAntiguaandChimaltenango,Standley 80988(F,US).—sANtA rosA:Barberena,Heyde 3704(G,GH,MO,NY,US).—soLoLá:abovePanajachel,LakeAtitlán,Hunnewell 14708(GH,MICH);slopesaboveLakeAtitlán,3–5kmWofPanajachel,Williams 25260(F,GH,NY,US,WIS). honduras. comAyAGuA:Km�28Tegucigalpa–Siguatepéquehwy,Edwards P-511(A,F,S,UC,US);entreLasLajasyYure,90kmNWdeComayagua,Nelson 6054(MEXU,MO,TEFH).—eL pArAíso:Hac.SanIsidrobetw.QuebradaElPescaderoandElParaíso,Molina R. 18786(F,NY);Danlí,Valerio R. 1989(F).—frANcisco morAzáN:rdtoAgalteca,Molina R. 18715(F,GH,NY);ValledeAmarateca,32kmNdeTegucigalpa,Navarro O. 58 (TEFH,MO).—oLANcho:RíoJuticalpa,6kmdeJuticalpa,Molina R. 13225(F,LL,NY,US);MontañaUval,carreteraaSanFranciscoLaPaz,Molina R. 13327(F,LL,NY).—yoro:Yoro,Edwards P-738(A,F,MO,NY,US);Victoria,orilladelRíoSulaco,Nelson 7013(MICH,MO,TEFH). Nicaragua. esteLí:EstelíRiver,5kmfromEstelí,Molina R. 23031(NY);SWdeSantaCruz,�kmdeCarreteraPanamericana,�3°00'N,86°�8'W,Moreno 5564(MICH,MO);SanJosédeLaguna,�3°29'N,86°2�'W,Moreno 8089(MICH,MO);alongnewrdfromHwy�(atca.Km�35.5andca.�0.6kmWofbridgeatLaTrinidad)toSanNicolás,ca.8.0kmfromHwy�,Stevens 10243(CAS,MEXU,MICH,MO);LlanoAlmaciguera,8.4kmSofHwy�,justSofEstelí,rdthroughEstanzuela2.8kmSofRíoEstanzuelabridge,�3°00'N,86°2�'W,Stevens 10753(DUKE,ENCB,MICH,MO,TEX);mountainsnearEstelí,Williams 10959(A,F,MICH,UC,US);RíoEstanzuela,8kmSWofEstelí,Williams 20185(F,NY,US).—JiNoteGA:caminoviejoMatagalpa-Jinotega,ca.6kmalNOdeciudadMatagalpa,Grijalva 4027(MO);ca.20.3kmNEofHwy�atEstelíonrdtoYalí,�3°�3'N,86°�5'W,Stevens 15775(MICH,MO).—LeóN:caminoaSanNicolás,LaGuayaba,a��kmdelaCarreteraPanamericana,�2°57'N,86°2�'W,Moreno 17790(MICH).

Galphimiaspeciosaiswidespreadandhighlyvariable;futurestudiesmayrevealevidence to support recognition of varieties.The shrubs and treelets occur fromsouthernMexicotonorthernNicaragua,butG. speciosahasnotbeenrecordedfromElSalvador.Theleavesusuallybeartheglandsonthemarginofthelaminanearthebaseorattheapexofthepetiole,butthepositionisvariableevenonthesameshoot.Thelimboftheallpetalsisusuallytriangular,broadlysointheposteriorpetal,andhasaroundedapex,butsometimesthelimbofthelateralpetalsisbroadlyellipticaltoovate.Theonlyotherspecieswithpersistentpetalsoccuringwithintherangeof


G. speciosaareG. elegans,withdistalleavesabaxiallypubescent,leafglandsbornenearthemiddleofthepetiole,andsmalleranthers(�.8–2mmlong),andG. multi-caulis,withrudimentarytoveryshortpetioles(eveninthelargestleaveslessthan�cmlong).

22. Galphimia multicaulisAdr.Juss.,Ann.Sci.Nat.Bot.,sér.2,�3:327.�840.Thryal-lis multicaulis(Adr.Juss.)Kuntze,Rev.gen.pl.�:89.�89�.—type: mexico.Oaxaca:“juxtaHuauapan,”Andrieux 496(holotype:K!;isotypes:G!K!M,P!P-JU,microfiche!;photoofPandP-JUisotype:MICH!;photosofMisotype:F!GH!MICH!MO!). Fig.29.

Smallmulti-branchedshrub to�.5m tall; stemspubescentwhenyoung, soonglabrous.Vestureofallvegetativepartsofsubsessile,crisped,reddishbrownhairs0.2–0.6mmlong.Laminasofthelargerleaves4–8.5cmlong,2–5.2cmwide,ellip-tical to lanceolate or ovate, apex apiculate, base acute, glabrous, secondary veinscommonlyprominentabaxially;marginentire;petioles0.3–0.8(–�)cm long,withscatteredhairsabaxiallytoglabrous;glandsborneonthemarginnearthebaseofthe lamina(toca.4mmabovethebase)orsometimesat thebase,usuallyapairbutsometimeswithanadditionalgland,orsometimesonly�glandor theglandsabsent,eachgland0.3–�mmindiameter,disklikeandprominent,orpeglikeandupto�mmlong,sometimesthelaminartissueextendedintoatooth;stipules2–5mmlong,0.7–�.8mmwide,triangular,glabrous.Inflorescenceadenseterminalraceme,inlargerplantstheracemesternateorgroupedinasmallpanicle,theaxestomentu-lose,glabrescentinage;peduncles�.5–6.5mmlong,pedicels3.5–�3mmlong,bothtomentulose,peduncles0.�–�.8 timesas longaspedicels;bracts2.2–4.5mm long,0.5–�mmwide,linear,bracteoles�.5–3mmlong,0.3–0.7mmwide,linear,bractsandbracteolesglabrousorwithscatteredhairsalongthemargin;bracteolesborneintheproximal�/4–2/3ofthepeduncle,suboppositeorupto�mmapart.Sepals2.5–3.8mmlong,�.4–�.8mmwide,oblongtoelliptical,glabrous,marginentireorfinelyerose;glandsabsent.Petalspersistent,unequal,yellow,glabrous,marginentireorfinelyandirregularlydenticulate;lateralpetals:claw�.5–2mmlong,0.5mmwide,limb7.5–�0mmlong,4.5–5.2mmwide,narrowlyovatetonarrowlyelliptical,apexsubacute,basesubtruncateorsubacute;posteriorpetal:claw2–3.5mmlong,�.2–�.3mmwide,limb8–�0.5mmlong,6–7mmwide,triangularorsometimesovate,apexobtuse,basetrun-cateorshallowlycordate.Stamensofunequallength;filamentoppositeanteriorsepal4–5.5mmlong,filamentsoppositeanterior-lateralpetals(3–)4–5mmlong,filamentsopposite anterior-lateral sepals (3.7–) 4–5 mm long, filaments opposite posterior-lateralpetals(2.5–)3.2–4mmlong,filamentsoppositeposterior-lateralsepals(3.7–)4.3–5mmlong,filamentoppositeposteriorpetal2.5–3.5mmlong;anthers2.2–3mmlong.Ovaryglabrous;styles5–6.5mmlong.Coccusca.3mmlong,2–2.6mmindiam-eter,glabrous;areole3mmlong,2mmwide;outercotyledonca.2.5mmlong,innercotyledonca.2.3mmlong.Chromosomenumberunknown.

Phenology.Collected inflower fromAugust throughDecember, in fruit fromSeptemberthroughNovember,inOaxacaalsoinflowerinJuneandinbudinMay.

Distribution (Fig. 30). Mexico (Guanajuato, Jalisco, Michoacán and adjacentMéxico,disjuncttoOaxaca);pine-oakforestandmatorral;�450–2350m.

represeNtAtive specimeNs.mexico.GuANAJuAto:Mpio.Victoria,Malinto,Badillo 3(MICH);MontesdeObrajuelo,�2Oct�9�3,Salazar s.n.(MEXU,US).—JALisco:Mpio.ZapotlándelRey,CerrodeSantaFé,Cházaro B. 5033(CHAPA,ENCB,IBUG,IEB,XAL);Mpio.CiudadGuzmán,20kmalNWdeCd.Guzmán,brechaaYerbalito,Fuentes O. 827(CHAPA,IEB,MICH);Mpio.Jocotepec,CerroViejo,veredaa


FIG.29.Galphimia multicaulis.a.Floweringbranch.b–d.Leafbases,abaxialview,toshowvariationinglands.e.Flowerborneonpedicelandpeduncle,withportionof inflorescenceaxis. f.Lateralpetal.g.Posteriorpetal.h.Stamens,adaxialview,stamenatrightoppositeposteriorpetal,stamenatleftoppo-siteposterior-lateralsepal.i.Gynoecium.j.Cocci,adaxialview(left)andlateralview(right).k.Embryo.Scalebar:a,4cm;b–e,8mm;f,g,5mm;h,i,4mm;j,k,2.7mm.(Basedon:a,Jones 40975,MO;b,Díaz & Barriga 4760,MICH;c,Machuca N. 4412,XAL;d,e,j,k,King & Soderstrom 4671,MICH;f–i,Cházaro B. 5033,CHAPA.)


BarrancadelAgua,enfrentedeZapotitándeHidalgo,Machuca N. 3551(MICH,XAL);Mpio.Jocotepec,CerroViejo,laderadeexposiciónS,enfrentedeSanJuanCozalá,20°�5–30'N,�03°20–40'W,Machuca N. 4412(XAL);SierradelTigre,3miSofMazamitla,McVaugh 12988(BM,G,MEXU,MICH,US);baseoftrailleadingtohilltoweringbehindIxtlahuacándelosMembrillos,offRt35toChapala,Norman 661(MICH);5kmSofQuitupan,Villarreal 8215 (ENCB,MICH).—méxico:ValledeBravo,Lyonnet s.n.(MEXU);5kmNofValledeBravo,Ripley & Barneby 14869 (CAS,NY).—michoAcáN:Cerrode lasNalgas,vic.ofMorelia,Arsène 2654 (MEXU,US);vicinityofMorelia,NWofPungarato,Arsène 5164(A,BM,MO,NY,US);Morelia,Pungarato,Arsène 8402(GH,MICH,NY,UC);Mexhwy�5atSanJosédePuruard,Barr & Barr 64-554(DS,UC);Mpio.Morelia,entreelCerroPrietoyElPunhuato,Díaz B. 4760(IEB,MICH);Mpio.Morelia,SanJoséCoapa,Escobedo 1850(GUADA,IEB,MICH,TEX);slopesofCerroSantaMaría,8–�0kmSWofJiquilpan,5kmNEofQuitupan,Feddema 93(DS,DUKE,ENCB,MICH,TEX);8miSofZitácuaro,Jones 40975(MEXU,MO);CerroPotrerillos,5miNofCotija,22miSofJiquilpan,King & Soderstrom 4671(MICH,NY,TEX,UC,US);Mpio.Chilchota,300malNEdelpuebloHuancito,Romero B. 13(IBUG);Mpio.Huaniqueo,0.8kmalSWdeTendeparacua,centroNEdepedregalpequeño,Silva-Sáenz 139(MICH).—oAxAcA: Huauclilla,Nochixtlán,Conzatti & González 1199(GH);CuestadeHuauclilla,Dtto.Nochixtlán,Conzatti 1881(F,GH,MEXU);Mpio.Nochixtlán,Huauclilla,Conzatti 3965(MEXU);LaCarbonera,Dtto.Etla,Conzatti 4210(MEXU,US);Nochistlán,Jun�844,Galeotti s.n.(BR);withoutlocality,Jurgensen 288(DS,G,K);cuestadeSanJuandelEstado,Pringle 4828(A,BM,BR,F,G,GH,GOET,MEL,MEXU,MO,NY,S,UC,US,W);propeOaxaca,Seler& Seler 98(GH);mountainsnearSanJuandelEstado,Smith 82(GH).

Galphimiamulticaulis isdistinguishedby its leavesandthecrowded inflores-cences.Thepetiolesareveryshort(usually3–8mmlong),andtheleafglands,borneonthemarginandabovethebaseoftheellipticaltoovatelaminas,varyfrompromi-nenttopeglike(Fig.29b–d).Galphimia multicaulisoccursincentralMexicoandisalsodisjunctinnorthernOaxaca,wherethisshowyanddistinctivespecieshasbeen

FIG.30.DistributionofGalphimia floribunda, G. hirsuta,andG. multicaulis.


collectedonlytentimes,mostrecentlyin�92�(Conzatti 4210).TheOaxacancollec-tionsdifferonlyinthattheyhaveshorterpeduncles(�.5–3mmlong;0.�–0.4timesaslongaspedicels)thanspecimensfromelsewhere(3–6.5mmlong;0.3–�.8timesaslongaspedicels).AllbutoneotherspecieswithwhichG. multicaulisissympatrichavewell-developedpetioles.InOaxacathedistributionofG. multicaulisoverlapswiththerangeofG. sessilifolia,ararelycollectedspecieswithdeciduouspetalsandsessiletosubsessileleaves.

Jussieunoted“v.s.herb.J.Gay”(nowatK)intheprotologueforG. multicaulis.Therefore,IconsiderthesheetatKfromtheGayherbarium,whichbearsJussieu’sannotation,theholotype;theduplicateatP-JUisanisotype.

GalphimiamulticauliswastreatedbyBartling(�840)underthenameG. glaucaandbySmall(�9�0)underThryallis multicaulis.

23. Galphimia paniculataBartl.,Linnaea�3:556.�840[“�839”].Thryallis paniculata (Bartl.)Kuntze,Rev.gen.pl.�:89.�89�.—type: mexico.“InregnoMexi-cano,”Hegewisch s.n.(holotype:GOET!;isotype:NY!).

Galphimia glandulosaCav. [var.]αovalifoliaDC.,Prodr.�:582.�824.—type: Sessé&Mociñodrawingno.0269,TornerCollectionofSesséandMociñoBiologicalIllustrationsattheHuntInstituteforBotanicalDocumentation(holotype,digitalimage!;photosofCandollecopyofdrawingatG:F!GH!GOET!MICH!MO!).

Galphimia glandulosaCav.[var.]βoblongifoliaDC.,Prodr.�:582.�824.—type: Sessé&Mociñodrawingno.0�48,TornerCollectionofSesséandMociñoBiologicalIllustrationsattheHuntInstituteforBotanicalDocumentation(holotype,digitalimage!;photosofCandollecopyofdrawingatG:F!GH!MICH!MO!).

Galphimia latifolia Bartl., Linnaea �3: 553. �840 [“�839”]. Thryallis latifolia(Bartl.)Kuntze,Rev.gen.pl.�:89.�89�,nonThryallis latifoliaMart.,�829.—type: BasedonplantsraisedfromseedsentbyHunnemannfromMexicoin�837(holotype:unknown).—SpecimentakenfromthebotanicalgardenatGöttingeninJune,�840,andpreservedintheBartlingHerbarium(neotype,heredesignated).

Galphimia humboldtianaBartl.,Linnaea�3:555.�840[“�839”].Thryallis hum-boldtiana(Bartl.)Kuntze,Rev.gen.pl.�:89.�89�.Galphimia paniculatavar.humboldtiana(Bartl.)Nied.,ArbeitenBot.Inst.Königl.LyceumHosianumBraunsberg5:26.�9�4.—type: mexico.[Guerrero:]“interAltodelosCax-onesetAcaguisotla,”Humboldt & Bonpland s.n.(holotype:P-HBK,micro-fiche:MICH!,photo:MICH!;isotype:P!).

Fig.3�.

Largeherbtoshrubortreeletto5m;stemspubescentwhenyoung,soongla-brous.Vestureofallvegetativepartsofsubsessile,straighttocrisped,reddishbrownhairs 0.2–0.6 mm long. Laminas of the larger leaves 5–�0 cm long, 2–6 cm wide,elliptical toovate,apexroundedorapiculate,baseattenuate,glabrous, secondaryveinsprominentabaxially;marginentire;petioles�.5–3.5cmlong,glabrous;glandsusuallyapairborneataboutthemiddleofthepetiole,suboppositeorupto�mmapart,sometimeswith�(2)smallergland(s)distally,eachgland�.5–2mmindiam-eter,disklikeandprominent;stipules�–2(–3)mmlong,0.5–�.5mmwide,narrowlytriangular,glabrousorwithscatteredhairsonthemargin.Inflorescenceusuallyalargepaniclecomposedofracemes(inveryyoungspecimensaterminalraceme),the


FIG. 3�. Galphimia paniculata. a. Flowering branch. b. Petiole, adaxial view; note glands. c. Par-tialandroeciumlaidout,abaxialview,thestamenatleftoppositeposteriorpetal.d.Fruitfromabove,withpetalspersistent,posteriorpetaluppermost.e.Cocci,abaxialview(left)andadaxialview(right).f.Seed.Scalebar:a,4cm;b,�.3cm;c,e,4mm;d,�cm;f,2.7mm.(Basedon:a–c,Machuca N. 5023,MICH;d,McVaugh 14237,MICH;e,f,González T. 9,MICH.)


axespubescent;peduncles3–7.5mmlong,pedicels(7.5–)9–�3(–�6.5)mmlong,bothsparselypubescenttoglabrous,peduncles0.3–0.4(–0.5)timesas longaspedicels;bracts�.4–3(–5)mmlong,0.4–0.8(–�)mmwide,linear,bracteoles�–2(–2.8)mmlong,0.3–0.6mmwide,linear,bractsandbracteolesglabrous;bracteolesborneinthedistal�/4–�/2ofthepeduncle,suboppositeorupto2mmapart.Sepals3.3–4.5mmlong,�.4–2.5mmwide,oblongornarrowlyellipticaltolinear-lanceolate,glabrousorwithafewscatteredhairsatbase,marginirregularlydenticulate-erose-ciliate;glandsabsent.Petalspersistent,unequal,yellow,theclawandmidriboftensuffusedwithred,glabrous,marginirregularyfinelydenticulate;lateralpetals:claw�.3–2mmlong,ca.0.5mmwide,limb8–��mmlong,4.5–6.5mmwide,ellipticalorlanceolate,apexsub-acutetoobtuse,basegraduallyattenuate;posteriorpetal:claw2–3mmlong,�.�–�.3mmwide,limb9–��mmlong,6–8mmwide,ovateorbroadlyelliptical,apexobtuse,basesubacute.Stamensofunequallength;filamentoppositeanteriorsepal3.5–4.4mmlong,filamentsoppositeanterior-lateralpetals(2–)2.5–3.5mmlong,filamentsoppositeanterior-lateralsepals3.3–4.4mmlong,filamentsoppositeposterior-lateralpetals�.5–2.5(–3)mmlong,filamentsoppositeposterior-lateralsepals3.3–4.2mmlong, filament opposite posterior petal 2–2.8 mm long; anthers 2.3–2.8 mm long.Ovaryglabrous;styles(4–)5.2–6.7mmlong.Coccus3–3.8mmlong,2.4–2.7mmindiameter;glabrous;areole2.2–2.8mmlong,�.9–2.5mmwide;outercotyledonca.3mmlong,innercotyledon2–2.5mmlong.Chromosomenumberunknown.Fig.3�.

Phenology.CollectedinflowerandfruitfromAugustthroughApril,sometimesalsoinMayandJune.

FIG.32.DistributionofGalphimia arenicolaandG. paniculata


Distribution(Fig.32).Mexico(Guerrero,Jalisco,México,Michoacán,Morelos,Nayarit,westernOaxaca,Puebla);tropicaldeciduousandsubdeciduousforest,oakandpine-oakforest,pedregal,matorral,roadside,pasture;700–2220m.

represeNtAtive specimeNs.mexico.Guerrero: �0kmalNEdeIguala,LaCumbre,desv.alaTorredeMicro-ondas,Díaz M. 241(DS,ENCB,MICH,WIS);ca.8km(airline)SWofXochipala,Feddema 2773(DUKE,ENCB,MICH,MO);Manchón,Hinton 11292(BM,DS,F,GH,MICH,NY,UC,US);Mpio.Mina,CampoMorado,Hinton et al. 14846(GH,LL,MICH,NY,UC,US);Mpio.Chilpancingo,SaltodeValadeza6kmalNdeMazatlán,López H. 1030(ENCB,MEXU);Mpio.Mina,SierraMadredelSur,HuentaVieja,Mexia 9010a(GH,UC,US);Mpio.Alcozauca,Alcozauca,LasMesitas,Viveros S. & Casas 331(ENCB,MEXU,MO).—JALisco:7miNEofTecolotlánonroadtoGuadalajara,Anderson 13564(MICH);Mpio.Autlán,entreTecopatlányManantlán,�9°37'�0"N,�04°�2'�8"W,Cárdenas T. et al. 28(WIS);Mpio.Tequila,2–3kmadelantedeTequilarumboalcerrodeTequila,Cházaro 4386(IBUG,XAL,WIS);alongHwy80,ca.6miSWofCasimiroCastilloandca.4�miNEofCihuatlán,Clarke et al. 1987-1(MICH);Mpio.Ixtla-huacándelRío,LosPitayitos,González T. 9(DS,ENCB,IBUG,MICH,TEX,WIS);Mpio.VillaGuerrero,4kmalNEdeElRefugioBolañosy26kmalSWdeVillaGuerrero,Lott 2120(CHAPA,F,IEB,MEXU,MICH,XAL);Mpio.AcatlándeJuárez,“ElCuarenta”alNEdeAcatlán,Machuca N. 5023(MICH);aboveTalpadeAllende,nearrdtoMascota,McVaugh 14237(BM,MEXU,MICH,US);Mpio.Zapopan,cercadelasTimbres,brechaSta.Lucia–PaloGordo,Navarro G 195(IBUG);RíoBlanco,Palmer 684in�886(GH,MICH,NY,US);slopesofbarrancanearGuadalajara,Pringle 1728(A,BM,BR,F,G,GH,MO,NY,S,UC,US,W);barrancaofGuadalajara,Pringle 11391(CAS,F,GH,L,MICH,MO,S,UC,US);Mpio.Tolimán,PuertodeToxín,6kmalSdeSanPedroToxín,Ramírez S. et al. 208(IBUG);rdbetweenBolañosandGua-dalajara,Rose 3021(NY,US).—méxico: Bejucos,Temascaltepec,Hinton et al. 2546(A,G,NY,US);CerroLaCorona,Zacualpan,Matuda 30349(MEXU);Tejupilco,Matuda 27477(MEXU);Mpio.ValledeBravo,CeritodePeña,Matuda 28046(MEXU);Mpio.SanTomás,2kmSofSanNicolásTolentino,8kmNEofZuluapan,�9°09'N,�00°�5'W,Solheim & Benz 1064(MEXU,XAL,WIS).—michoAcáN:Mpio.Villamar,CerrodeCotijarán,García 1272(IEB);brechaaDosAguas,González & Martínez 720(ENCB,IBUG,MEXU);Mpio.Coalcomán,Coalcomán,Hinton 12694(G,LL,MICH,NY,S,UC,US);Mpio.Coalcomán,Aquila,Hinton 16245(ENCB,G,LL,MICH,NY,P,UC,S,W);MontedeSta.Helena,Langlassé 64(G,GH,K,P,US);steepdrymountainsidesca.8kmNWofAguililla,rdtoAserraderoDosAguas,McVaugh 22665(ENCB,MICH);Susupuato,Ramos et al. 588(MEXU,XAL);a2kmalSWdeAriodeRosales,Soto N. 580(BM,ENCB,MEXU);26kmalSWdeZitácuaro,entrandoporlapresadelBosque,Torres & Ramírez 13640(ENCB,IEB,MICH).—moreLos: Mpio.Xochicalco,zonaarqueológica,3kmalOdelacarr.Fed.México–Acapulco,Anaya R. 8(CHAPA,F,XAL);Km�6delacarreteraCuautla–Tepoztlán,Arreguín 327(CAS,CHAPA,ENCB,MEXU,NY);Mpio.Tepoztlán,8kmdespuésdeTepoztlán,rumboaSto.DomingoOcotitlán.,Espejo 3426(CHAPA,ENCB,MEXU,UAMIZ);Mpio.Yautepec,�kmalSEdelpobladodeLaJoya,carr.Cuernavaca–Yautepec,�8°52'�5"N,99°07'05"W,Flores F. 366(IEB,MEXU);entreCuernavavaetlevillagedeSochil,Ghiesbreght 303(K,P);Mpio.Tlayacapan,nearTlayacapan,rdtoOaxtepec,Lott 247(CHAPA);Xochitepec,Lyonnet 1089(MEXU);Km�6carr.Cuautla–Tepoztlán,Or-dorica 160(CHAPA,ENCB,MEXU,XAL);Cuernavaca,Pringle 7215(GH,MICH,US).—NAyArit:hillsbackof[townof]Jalisco,Ferris 5943(A,DS,US);mountains20miSEofAhuacatlánonrdtoBarrancadelOroandAmatlán,McVaugh & Koelz 847(ENCB,G,MICH,NY,US);trailYxtlántoBarrancadelOro,Mexia 784(A,BM,CAS,DS,F,G,GH,MICH,MO,NY,UC);Tepic,Palmer 2030in�892(BM,C,F,GH,MICH,NY,S,US);Mpio.Huajícori,RíodelosTalladeros,Solís 604(ENCB,MEXU,MICH,NY,TEX);Mpio.LaYesca,6.2kmalSEdePuentedeCamotlán,caminoaHuajimic,2�°36'N,�04°04'W,Tenorio L. 16675(IEB,MEXU).—oAxAcA:ArroyodelaPastor,S.Vicente,Putla,MacDougall H204(NY);Mpio.SanJuanMixtepec,SWdelRíoTimbre,Sánchez B. 41(CAS,CHAPA,ENCB,IEB,MEXU,MO,NY);Mpio.SanJuanMixtepec,NWdeLomadeLobo,Dtto.Juxtlahuaca,Sánchez B. 66(ENCB,IEB,MEXU);25.5kmalNdePutla,carr.Putla–HuajuapandeLeón,Distr.Putla,Torres C. 1956(CAS,F,MEXU,MO);Mpio.Silacayoapan,Dtto.Silacayoapan,�0kmalSEdeSanMartíndelEstado,haciaSilacayoapan,�7°33'N,98°07'W,Torres C. 14143(MEXU,MO,TEX).—puebLA:Mpio.Jolalpan,EjidodeXochitepec,Arreola et al. Xo-634(CAS);Mpio.Jolalpan,2kmalNWdeSanPedrodelasPalmas,Razo & García IIc-33(IEB,MEXU).—zAcAtecAs:4miNofElLimón,5.2miNofborderofJalisco,Croat 45091(MEXU,MO,NY);8miSofMoyahua,Webster 1081(ENCB,MICH).

Galphimia paniculataisacoarse,showyspeciescommonlycollectedinwesternMexicofromNayarittowesternmostOaxaca.Itisdistinguishedbyitslargecom-pound inflorescencesand leaves thathave the glandsborneat the middle of thepetiole;thelaminasvaryfromellipticaltoovate.Collectionsofthisspeciesareoften


misdeterminedasG. glauca(ofeasternandcentralMexico),whichhassmallerleaveswiththeglandsborneonthemarginofthelaminaandalsosmallerflowersandfruits;thelimboftheposteriorpetalisbroadlytriangular.Galphimia paniculataisatleastpartlysympatricwithfourspecieswithpersistentpetals,butnonehavetheleafglandsatthemiddleofthepetiole.GalphimiamulticaulisandG. callianthabearleafglandsonthemarginofthelaminawellabovethebase;inaddition,G. multicaulisdiffersinitsveryshortpetioles(lessthan�cm)andG. callianthainitsfringedsepals.Galphi-mia floribundaandG. grandiflorahavetheleafglandsatthebaseofthelaminaortheapexofthepetiole;especiallyinlargerlaminas,thebasemaybedecurrentwiththeglandssuperficiallyappearingtobeborneonthepetiole,orthelaminartissuemaybedrawnoutintoatoothterminatinginagland.Galphimiafloribundaalsodif-fersfromG. paniculatainitsalmostorcompletelyglabrousinflorescences;G. gran-difloraisnamedforitslargepetals(thelimbofthelateralpetals��–�5.5mmlong).

Bartling(�840)basedhisG. paniculataonaherbariumcollection,buthisG. lati-foliaonlivingplantsofthesamespecieswithunusuallybroadlaminas,whichweregrownatthebotanicalgardeninGöttingen.ThespecimenherechosenasneotypeofG. latifolia consistsonlyofaninflorescencebranch;itistheonlyspecimenIhaveseenthatmayhavebeencollectedfromtheplantsstudiedbyBartling,butitisnotpossibletoascertainwhetheritispartofthetypematerial.BartlingalsoprovidedanadditionalsynonymforG. paniculata.HerealizedthatKunth(�822)hadmisap-pliedthenameG. glandulosaCav.andprovidedthenameG. humboldtiana,whichis validated by Bartling’s reference to Kunth’s description and the Humboldt & Bonplandspecimen;Bartlingalsogaveabriefdescriptionbutdidnotciteanyotherspecimens.

Candolle’svarietiesof“Galphimi glandulosa”werebasedondrawingspreparedfortheSesséandMociñoexpedition.Thatof“β oblongifolia”isclearlyG. paniculata.Thedrawingof“α ovalifolia”isquitestylizedandthereisnospecimenintheSesséandMociñoherbarium(housedatMA);however,atGisaspecimenofG. paniculatalabeled,likethedrawing“Tlaxacoxochitl”andsaidtobefrom“Hb.Pavon”(i.e.,exMA),whichmaybeauthenticmaterial.

Small(�9�0)includedG. paniculata, G. latifolia,and G. humboldtianawithinhisconceptofThryallis glauca.

24. Galphimia callianthaC.Anderson,sp.nov.—type: mexico.Guerrero:23–24kmEofPetatlánonrdtoTlapa[“Teapa”onlabel],Km�23–�24,2000m,�2–�3Feb�970,Anderson & Anderson 5758(holotype:MICH!;isotypes:DUKE!ENCB!SD!). Fig.33.

Frutexvelarborparva.Laminaefoliorummajorum3–6.5cmlongae,�.6–3.5cmlatae,oblongaevelellipticaevelovataevelangusteovatae,glabrae,margineintegra,glandulis ca. 0.5 mm diametro; petioli 0.8–�.8 cm longi. Inflorescentia racemosa,axibustomentosis.Sepalamargineciliata.Petalapersistentia;petalalateraliaungueca.�.5mmlongo,limbo�0–�2mmlongo,5.5–9.5mmlato,ovatovelangusteovato;petalumposticumungueca.2mmlongo,limbo�0.3–�2.5mmlongo,6.5–��mmlato,ovato-triangulari.Antherae2.5–2.7mmlongae.Ovariumglabrum;styli4.5–5.2mmlongi.Coccusca.4.5mmlongus,glaber.

Shrubortreeletto3m;stemsdenselytomentosewhenyoung,soonglabrous.Vestureofallvegetativepartsofsubsessile,crispedtocurled,reddishbrownhairs0.5–�.5mmlong.Laminasofthelargerleaves3–6.5cmlong,�.6–3.5cmwide,oblongtoellipticalornarrowlyovate,apexobtuseorapiculate,baseacute,glabrous,second-ary veins barely prominulous; margin entire; petioles 0.8–�.8 cm long, tomentose


FIG.33.Galphimia calliantha.a.Floweringbranch.b.Leaf,abaxialview.c.Baseofleaf,adaxialview;noteglandsonmargin.d.Flower,sideview.e.Sepal,abaxialview.f.Lateralpetal,adaxialview.g.Posteriorpetal,adaxialview.h.Partialandroeciumlaidout,abaxialview,stamenatleftoppositeposteriorpetal.i.Gynoecium.j.Cocci,adaxialview(left)andlateralview(right).k.Embryo.Scalebar:a,b,4cm;c,f,g,8mm;d,�.3cm;e,h–j,4mm;k,2mm.(Basedon:a,d–k,Rzedowski 18095,MICH;b,c,Hinton 9848,MO.)

whenyoung,becomingglabrous;glandsapairborneonthemarginofthelamina,commonly0.5–�.5cmabovethebase,eachglandca.0.5mmindiameter,prominent,oftenwithadditionalglandsalongthemargin,especiallyinthesmallerleavesneartheinflorescence;stipules2.7–4mmlong,0.8–�mmwide,narrowlytriangular,gla-brous.Inflorescenceadenseterminalracemeoraternatepanicle,theaxesdenselyreddishtomentose;peduncles2.5–7(–9)mmlong,pedicels5–��(–�3)mmlong,bothtomentose,peduncles0.3–0.7(–0.9)timesaslongaspedicels;bracts3–5.5mmlong,0.5–0.8mmwide,linear,bracteoles�.6–3mmlong,0.3–0.5mmwide,linear,bractsandbracteolesglabrousandwithscatteredhairsalongthemargin;bracteolesborneinthedistal�/4–�/2ofthepeduncle,subopposite.Sepals4–4.5mmlong,�.6–2.2mmwide,oblongtonarrowlyelliptical,glabrousorwithscatteredhairsintheproximal


�/3,marginfringedwithreddishhairsto0.2(–0.3)mmlong;glandsabsent.Petalspersistent,unequal,yellow,theclawoftensuffusedwithred,glabrous,marginentireorfinelyerose;lateralpetals:clawca.�.5mmlong,ca.0.5mmwide,limb�0–�2mmlong,5.5–9.5mmwide,ovateornarrowlyso,apexobtuse,basesubacute;posteriorpetal:clawca.2mmlong,�–�.2mmwide,limb�0.3–�2.5mmlong,6.5–��mmwide,triangular-ovate,apexobtuse,baseslightlycordate.Stamensofunequallength;fila-mentoppositeanteriorsepal4.5–5mmlong,filamentsoppositeanterior-lateralpetals3.7–4.2mmlong,filamentsoppositeanterior-lateralsepals4–4.8mmlong,filamentsopposite posterior-lateral petals 3–3.5 mm long, filaments opposite posterior-lat-eralsepals4–4.7mmlong,filamentoppositeposteriorpetal3–3.2mmlong;anthers2.5–2.7mmlong.Ovaryglabrous;styles4.5–5.2mmlong.Coccusca.4.5mmlong,ca.3.5mmindiameter,glabrous;areoleca.3mmlong,ca.2.2mmwide;outercotyledonca.2.7mmlong,innercotyledonca.2mmlong.Chromosomenumberunknown.

Phenology.Collected inflower fromSeptember throughMarch, in fruit fromJanuarythroughMay.

Distribution(Fig.35).Mexico(GuerreroandwesternmostOaxaca);inpine-oakforestonlimestone;�600–2200m.

AdditioNAL specimeNs exAmiNed. mexico. Guerrero: Mpio. Tixtla de Guerrero, Atliaca, entreAcatempa yAtliaca, �7°38'02"N, 99°2�'03"W, Hall 778 (MICH); Distr. Mina, Chiriagua, Hinton 9848(BM,G,MO);SierraMadre,Langlassé 780(F,G,GH,K,NY,P,US);Mpio.Chichihualco,2kmadelantedeCruzdeOcote,Lozano V.171(ENCB);Mpio.ChilpancingodelosBravos,a2kmalEdeOmiltemi,caminoaChilpancingo,Martínez S. & Téllez 285(MBM,MEXU);Mpio.ChilapadeAlvarez,Km72–73de la carretera Chilapa–Tlapa, �7°35'53.�"N, 99°06'02.2"W, Panero 3912 (IEB,TEX); �� km alW deAmojileca,caminoChilpancingo–Amojileca,Rodríguez B. 65(ENCB,MEXU,MO);Mpio.Chichihualco,CruzdeOcote,25kmalWSWdeCamotla,Rzedowski 18095(DS,ENCB,MICH);Mpio.Chilpancingo,LaSimaroa,�0kmalEdeOmiltemi, Rzedowski 30261(ENCB,SD);Mpio.Mochitlán,ElVoladerito,cercadeCoxtlahuacán,Rzedowski 30290(ENCB);Mpio.Chichihualco,CruzdeOcote,sobreelcaminodeXochipalaalaserraderoAguaFría,Rzedowski & McVaugh 284(ENCB,MICH);WofChilpancingo,Sharp 441453(GH);Hwy�34betweenZihuaquioandIxtapa,crestofmtsbetweenPacificandRíoBalsaswatersheds,�8°00'N,�0�°�3'W,Soulé 3228B (MO,TEX).—oAxAcA:Mpio.SantiagoJuxtlahuaca,Dtto.Juxtlahuaca,ElManzanal,�7°�3'�3.20"N,98°03'38.30"W,Calzada 21372(CICY,MO).

Galphimiacallianthaisahandsomeshrubortreeletofthepine-oakforestsofGuerreroandwesternmostOaxaca.Itslargeflowersareaggregatedindenseracemeswithreddishtomentoseaxes.Itistheonlyspeciesinwhichthesepalsarefringedwithreddishhairsalongthemargin(lessobviousinolderflowers).Thelaminasareoftenapicallyobtuseandbeartheleafglandsonthemargin;oftenseveralsmallerglandsarepresentinadditiontothelargerpairnearthebase.TheonlysympatricspecieswithpersistentpetalsisG. paniculata,whichhastheleafglandsatthemiddleofthepetiole.

25. Galphimia grandifloraBartl.,Linnaea�3:554.�840[“�839”].Thryallis grandiflora(Bartl.)Kuntze,Rev.gen.pl.�:89.�89�.Galphimia gracilisvar.grandiflora(Bartl.)Nied.,ArbeitenBot.Inst.Königl.LyceumHosianumBraunsberg5:24.�9�4.—type: mexico.“InregnoMexicano,”Hegewisch s.n. (holotype:GOET!;isotype:NY!)

Galphimia schiedeana Nied., Arbeiten Bot. Inst. Königl. Lyceum HosianumBraunsberg5: 28. �9�4.—type: mexico. [Michoacán:]pr.Ario,Dec�829,Schiede s.n. (lectotype,heredesignated:NY!,fragmentsfromspecimenfor-merlyatB,nowdestroyed;photosofSchiede s.n.syntype:F!GH!MICH!MO!NY!).

Fig.34.


FIG.34.Galphimia grandiflora.a.Floweringbranch.b.Baseoflamina,abaxialview,showingpairofmarginalglands.c.Flowerbud,borneonpedicelandpedunclewithportionofinflorescenceaxis.d.Lateralpetal.e.Posteriorpetal.f.Partialandroeciumlaidout,abaxialview,thestamenatleftoppositeposteriorpetal.g.Gynoecium.Scalebar:a,4cm;b–e,8mm;f,g,4mm.(Basedon:a,c–g,McVaugh 26100,MICH;b,Anderson & Anderson 5881,DUKE.)

Large herb, shrub, or treelet to 5 m tall; stems strigose to tomentulose whenyoung,soonglabrous.Vestureofallvegetativepartsofsubsessile,straighttocrisped,reddishbrownhairs0.2–0.8mmlong.Laminasofthelargerleaves5–�3.5cmlong,2–7.3cmwide,narrowly tobroadlyelliptic,ovate,oblanceolate,or rhombic,apexapiculateorrarelyacute,baseattenuateandoftenbrieflydecurrent,glabrous,second-aryveinsprominulousorbarelysoabaxially;marginentire;petioles�.5–5cmlong,sparselystrigosetoglabrescent;glandsapairborneatthebaseofthelamina(ifbasedecurrent,glandsuperficiallyappearingasborneonthepetiole),oftenattheapexofatoothoflaminartissue,sometimesonthemarginjustabovethelaminarbase,


orsometimesoneglandatthelaminarbaseandtheotherattheapexofthepetiole,sometimeswith�–2additionalsmallerglandsonthemarginnearthebase,rarelywithonly�gland,eachgland0.6–�.3mmindiameter,disklikeandprominent,orpeglikeandupto�.5mmlong;stipules2–3.5(–4.5)mmlong,0.5–�(–�.5)mmlong,linearornarrowlytriangular,glabrous,themargincommonlyciliate.Inflorescenceadenseterminal racemeorapanicle, theaxes tomentulose, sparsely so inage;peduncles6–�� mm long, pedicels 6–�2 mm long, both tomentulose, peduncles (0.5–) 0.6–�(–�.6)timesaslongaspedicels;bracts(3–)4–6.7mmlong,0.5–0.8mmwide,linear,bracteoles 2–4 mm long, 0.3–0.5 mm wide, linear, bracts and bracteoles glabrous;bracteolesborneinthedistal�/3–�/2ofthepeduncle,mostlysuboppositeorsome-timesupto�mmapart.Sepals4.2–5.5mmlong,2–2.5mmwide,ellipticalornarrowlyso or sometimes oblong, glabrous, margin entire; glands absent. Petals persistent,yellow,theclawoftensuffusedwithred,sometimesalsothelimbinage,thecentralveinoftengreen,glabrous,margindenticulate;lateralpetals:claw�.8–2.5mmlong,0.5–0.7mmwide,limb��–�5.5mmlong,6–9mmwide,ellipticaltolanceolate,apexsubacute,baseattenuate;posteriorpetal:claw2–3(–3.5)mmlong,�–�.5mmwide,limb��.5–�5.5mmlong,7–9mmwide,ovateorbroadlyelliptical,apexsubacute,baseattenuate.Stamensofunequallength;filamentoppositeanteriorsepal4.3–5.5mmlong,filamentsoppositeanterior-lateralpetals3.5–4.3mmlong,filamentsoppositeanterior-lateralsepals3.8–5mmlong,filamentsoppositeposterior-lateralpetals3–4mmlong,filamentsoppositeposterior-lateralsepals4–5.2mmlong,filamentoppositeposteriorpetal3–3.5mmlong;anthers(2–)2.5–3.5mmlong.Ovaryglabrous;styles5.2–6 mm long. Coccus 3.�–3.6 mm long, 2.4–3 mm in diameter, glabrous; areole2.5–2.7mmlong,2–2.3mmwide;outercotyledon2.3–2.5mmlong,innercotyledon�.8–2mmlong.Chromosomenumberunknown.

Phenology.CollectedinflowerfromNovemberthroughApril,infruitfromJanu-arythroughApril.

Distribution(Fig.35).Mexico(Colima,Jalisco,México,Michoacán,Morelos);mostlyinwetorhumidplacesinforestsdominatedbypines,oaks,andfirs,alsoatroadside;�360–2800m.

represeNtAtive specimeNs.mexico.coLimA:Mpio.Comala,fraccionamiento5kmantesdellegaraSanAntoniocarr.Queseria–SanAntonio,Acevedo R. & López 1153(IEB,MICH,TEX,WIS,XAL);Vol-cándeColima,Zepeda R. s.n.(IBUG).—JALisco: WslopesofSierradeSanSebastián,�5–30km(strtline)NofMascota,rdtoSanSebastián,Anderson & Anderson 5962(ENCB,MICH,SD);SierradeMinatlánaboveHaceradero[aserradero?],Boutin & Kimnach 2960(CAS,MEXU);Km33sobrelabrechaaTalpadeAllende,entrandoporlacarr.PuertoVallarta–ElTuito,Campos V. 4531(MEXU,TEX);Mpio.Atengo,VeredadeTacotaaRanchoViejo,Cházaro B. et al. 6874(IEB,MICH,TEX,XAL);Mpio.SanSebastián,23.6kmfromSanSebastiánonrdtoTalpa,Cowan 4760(NY,TEX);EstacióndeMicro-ondas,VolcándeFuego,Díaz L. 3125(CAS,ENCB,GUADA);Mpio.Autlán,�6–�7kmalSSEdeAutlán,4–5kmalSdeAhuacapan,Zacahuautla,�9°37'43"N�04°�9'00"W,Guzmán & López 629(MEXU,WIS,ZEA);Mpio.VenustianoCarranza,CerroViejosubidaporApango,Huerta et al. 216(MICH,TEX,XAL);�.5–3kmalEdelaEst.Biol.“LasJoyas,”caminoalCerroElAlmeal,�9°34–35'N,�04°�6'W,Judziewicz & Cochrane 4845(IBUG,MEXU,WIS,ZEA);�2–�5km(strtline)SWofBolaños,nearsummitsalong“CaminoViejo”toBerbería,McVaugh 25894(MICH);foothilsofSierradeManantlán,alongrdtosummit-pass,�6–22kmSofElChante,McVaugh 26100(MICH);fromSanSebastiántoRealAlto,Mexia 1637-a(A,CAS,GH,MICH,MO,UC,US);Km36delacarreteraaLasMinasdeCuale,lacualcomienzaaproximadamentea7–9kmalWdeElTuito,Panero 2866(TEX);Mpio.Gúzman,faldasdelNevadodeColima,Pérez A. 30(IBUG);Mpio.Tolimán,7–9kmalWdeLosSauces,caminoaTerreros,ElTerrero,Vázquez & Guzmán 4154 (WIS, ZEA).—méxico: Mpio.Valle de Bravo, 3 mi NE ofTemascaltepec, Dunn & Dunn 18965(ENCB,NY);RanchoSanLorenzo,neartownofValledeBravo,Gilly 2(MICH,NY);Nanchititla,Temas-caltepec,Hinton 7364(BM,F,MEXU,NY,S,US);CañadadeNanchititla,Matuda 38268(LL,MEXU);6kmEofNanchititla,30kmSWofTejupilco,�8°50'N,�00°45'W,Neill 5365(MEXU,MICH,MO);Mpio.Tejupilco,Ocotepec,Rzedowski 25296(DS,ENCB,LL,MICH);ca.�5.5miNofTejupilco,ca.3.5miSofTemascaltepeconHwy�34toToluca,�9°02'N,�00°03'W,Woodruff 271(F,MO,TEX).—michoAcáN:


mountainsWofAguililla,rdtoAserraderoDosAguas,Anderson & Anderson 5881(DUKE,ENCB,MBM,MICH,SD);LosCantiles,carr.Morelia,Cedillo T. 974(ENCB,LL,MEXU,MO);Mpio.Coalcomán,20.9kmalWdeCoalcománhaciaCoahuayana,Cowan 4910(MICH,TEX);Mpio.Uruapan,PresaCupa-titzio,Km89delacarr.Uruapan–PlayaAzul,desviaciónaMatarguarán,Fernández N. 4939(ENCB,IEB,MICH);Mpio.Jiquilpan,García R. 825 (WIS);Mpio.Zitácuaro,Zitácuaro–Lomas,Hinton et al. 13435(BM,DS,F,GH,MEXU,MICH,NY,UC,US);Mpio.Uruapan,Tancítaro,Hinton et al. 15684(GH,LL,NY,TEX,US);S-facingslopesofmtsbetweenRíodelSaltoandLaPolvilla,�8miEofMorelia,King & Soderstrom 5098(MEXU,MICH,NY,TEX,UC,US);3–4kmalNdeTacámbaro,Kishler 158(MEXU,MO);Mpio.Zacapu,5kmalSdeLasCanoas,Martínez L. 623(ENCB,IBUG,IEB,MEXU);Mpio.Ziracuaretiro,2kmalNEdeSanAndrésCorú,Martínez L. 967(ENCB,IEB,XAL);Mpio.Zinapécuaro,cercadeSantaCruz,Santos M. 1205(CHAPA,ENCB,IBUG,IEB,MEXU);3kmdelaentradaaTin-gambato,carr.aHuetamo,Torres & Ramírez 13762(ENCB,IEB,MEXU,MICH).—moreLos:Tepoztlán,Paray 1291(ENCB,MEXU,MICH);aboveCuernavaca,Pringle 8036(A,BM,BR,C,F,G,GH,GOET,L,MEXU,MO,NY,S,UC,US,W).

Galphimiagrandiflora isnamedfor itsunusually largepetals;atmaturity, thelimbsareupto�5.5mmlongand9mmwide.Thelaminasbearlarge,prominenttopeglikeglandsatornearthebase.Thisshowyandattractivespeciesisacommonlargeherborshrub,oreventreelet,ofwesternMexico,andoftenhasbeenconfusedwithotherpartlysympatricspecieswithpersistentpetals.Its leavesaresimilartothoseG. floribunda,whichiseasilydistinguishedbyitsnearlytoentirelyglabrousinflorescences.Galphimiamulticaulisdiffersinitsveryshortpetioles(mostly3–8mmlong)andG. paniculatainhavingtheleafglandsborneatthemiddleofthepetiole.

Small(�9�0)includedG. grandiflora inhisbroadcircumscriptionofThryallis glauca.

FIG.35.DistributionofGalphimia callianthaandG. grandiflora.


26. Galphimia floribundaC.Anderson,sp.nov.—type: mexico.Sonora:SierraLaChuna,RíoMayo,27Feb�935,Gentry 1372(holotype:MICH!;isotypes:A!ECON!K!MEXU!MO!S!UC!). Fig.36

Frutex.Laminaefoliorummajorum6–�4.5cmlongae,2.7–9cmlatae,ellipticaevelovataevelrhombicae,glabrae,margineintegra,glandulis0.5–�.5mmdiametro,sessilibusvelpedicellatis;petioli2.4–6.5cmlongi.Inflorescentiapaniculata,axibusglabris.Petalapersistentia;petalalateraliaungue(�.3–)�.5–2.2mmlongo,limbo7.5–�0mmlongo,4–6mmlato,ellipticovellanceolato;petalumposticumungue2.2–3.3mmlongo,limbo8.5–�2mmlongo,6–8mmlato,lateellipticovelovato.Antherae2.4–3.5(–3.8)mmlongae.Ovariumglabrum;styli4.5–5.7mmlongi.Coccusca.3.8mmlongus,glaber.

Shrub to 4 m; plants entirely glabrous or with sparsely scattered hairs in theinflorescence;hairs0.�–0.5mmlong,subsessile,straightorwavytocrisped,reddish.Laminasof the larger leaves6–�4.5cm long,2.7–9cmwide,elliptical toovateorrhombic,thelargerleavesverybroadlyso,apexapiculateoracute,baseattenuate,glabrous, secondaryveinsprominentabaxially;petioles2.4–6.5cm long,glabrous;glandsapairborneatbaseofbladeorapexofpetiole,eachgland0.5–�.5mmindiameter,disklikeandprominent,orpeglikeandupto�mmlong;stipules�.5–3.3(–4)mmlong,0.8–�.5mmwide,glabrous.Inflorescencea largepaniclecomposedof racemes, the axes glabrous or with a few scattered hairs; peduncles (4–) 5–��mmlong,pedicels4–8(–�2.5)mmlong,peduncles0.6–�.8timesaslongaspedicels,bothglabrousorwitha fewscatteredhairs;bracts (�.7–)2.5–4.5 (–6.2)mm long,0.5–�mmwide, linear tonarrowly triangular,bracteoles�.5–3.3mmlong,0.5–0.7mmwide,linearorsometimesnarrowlytriangular,bractsandbracteolesglabrousorsometimeswithscatteredhairsalongthemargin;bracteolesborneinthedistal�/4–�/2ofthepeduncle,suboppositeorupto�.5mmapart,themoredistalinserted0.7–4.5mmbelowtheapexofthepeduncle.Sepals3.5–5mmlong,�.5–2.2mmwide,ellipticaltooblong,glabrous,marginentire;glandsabsent.Petalspersistent,unequal,yellow,glabrous,marginfinelydenticulate;lateralpetals:claw(�.3–)�.5–2.2mmlong,0.5–0.8mmwide,limb7.5–�0mmlong,4–6mmwide,ellipticaltolanceolate,apexsubacute,baseattenuate;posteriorpetal:claw2.2–3.3mmlong,0.8–�mmwide,limb8.5–�2mmlong,6–8mmwide,broadlyellipticaltoovate,apexobtuse,baseattenuate.Stamensofunequallength;filamentoppositeanteriorsepal(3.5–)4.�–5.5mmlong,filamentsoppositeanterior-lateralpetals3.5–4.3(–4.7)mmlong,filamentsoppositeanterior-lateral sepals3.5–4.5 (–5.2)mmlong,filamentsoppositeposterior-lateralpetals2.7–4.2mmlong,filamentsoppositeposterior-lateralsepals3–4.2(–5.3)mmlong,filamentoppositeposteriorpetal(�.7–)2.2–2.7(–3.5)mmlong;anthers2.4–3.5(–3.8)mmlong.Ovaryglabrous;styles4.5–5.7mmlong.Coccusca.3.8mmlong,2.2mmindiameter,glabrous;areoleca.3mmlong,ca.2.�mmwide;outercotyledonca.2.5mmlong,innercotyledonca.�.7mmlong.Chromosomenumber:n=6(Seavey�975,as“G. glauca”).

Phenology.CollectedinflowerfromlateDecemberthroughJune(onecollectionfromSeptember,Urbina s.n.MEXU-�6562),infruitfromJanuarythroughJune.

Distribution(Fig.30).Mexico(Sonora,Chihuahua,Sinaloa,Nayarit,Durango,Jalisco); on dry slopes in deciduous forest, oak forest, and pine-oak forest;500–�300m.

represeNtAtive specimeNs. mexico.chihuAhuA: Mpio.Batopilas,barrancadeBatopilas,vic.ofArroyoSanFernando,WofLaBufa,Bye 3580 (TEX);CumbresdeLoreto,26°50'N�07°35'W,Hewitt 268 (GH);RíoBatopilasbridge,�.5miEofLaBufa,Kimnach 904 (UC,MEXU);RíoUriqueCanyon,


FIG.36.Galphimia floribunda.a.Floweringbranch.b.Detachedlargeleaf.c.Baseoflamina.d.Open-ingflower,borneonpedicelandpeduncle,withportionofinflorescenceaxis.e.Lateralpetal.f.Posteriorpetal.g.Androeciumlaidout,abaxialview,thestamenat leftoppositeposteriorpetal.h.Gynoecium.i.Cocci, adaxial view (above)and lateral view (below). j.Embryo.Scalebar: a,b, 4 cm; c, e, f, 8mm;d,5.7mm;g,i,4mm;h,j,2.7mm.(Basedon:a,i,j,Palmer 284 in�890,US;b,c,Rose 13095,US;d–h,Reina G. 97-338,MICH.)


Pennington 111 (TEX).—durANGo:Mexicohwy40,6.8miEofElPalmito,ca.53miEofConcordia,Breedlove 4254 (DS,DUKE,MICH);nearChacala,Goldman 326 (GH,NY,US);Mpio.Sianori,González Ortega 5379 (DS,K,MEXU,US);SanRamón,Palmer 101in�906 (CM,F,GH,K,MO,NY,UC,US).—JALisco:rdfromTequilatomicro-ondasonVolcánTequila(MontanaAzul),Daniel 4791 (CAS,DUKE,MICH,NY);Mpio.IxtlahuacándelRío,30kmalNdeIxtlahuacándelRío,ejidoPitallitos,Flores M. 549 (IBUG,MEXU);Mpio.Mascota,CerrodelaBarranca,González T. 36 (ENCB,MICH).—NAyArit:enlabasedelCerroSangangüey,a20kmalSEdeTepic,2�°28'N,�04°43'W,Téllez V. 10398(MEXU,MO).—siNALoA: Mpio.Rosario,4.9miEofSantaLucía,ca.23miEofConcordiaonMexico40,Breedlove 1663 (DS,DUKE,MICH);Mpio.SanIgnacio,Balboa,SanJavier,González Ortega 5094 (MEXU,US);Mpio.Cosalá,LaCopetona,arribadeGuadelupedeLosReyes,Tenorio L. 8358 (MEXU,MICH);Mpio.Concordia,carr.Mazatlán–DurangoentreKm240al256,entreOjodeAguaylaGuayamera,Vega 3617 (GUADA,MEXU).—soNorA:5.�miEofSantaRosaonoldroadtoYecora,Goldberg 76-155(ENCB);SierradelosAlamos,Palmer 284in�890(GH,MICH,NY,US);Mpio.Yecora,ArroyodelPilladito,nearTepoca,28°26'20"N,�09°�5'30"W,Reina 97-338(MICH);vicinityofAlamos,SierradeAlamos,Rose 12794, 13095(NY,US).

Galphimiafloribundaisahandsomeshrubnamedforitsunusuallylarge,distallyoftennoddinginflorescences,whichareglabrousorhaveveryfewscatteredhairsevenintheyoungestparts.Inmostspecies(exceptG. glandulosaandG. sessilifolia)theinflorescenceaxesarepubescent,theimmatureportionsoftendenselyso;theabsenceofsuchvesturereadilyseparatesG.floribundafromG. grandifloraand G. paniculata, twootherspecieswithpersistentpetalswithwhichitissympatricinthesouthernpartofitsrange.InG.floribunda theleafglandsareborneatthebaseofthelamina,oftenattheapexofatooth(Fig.36c)inlargerleaves.Galphimia grandifloramayhavesimilarleafglands,butthoseofG. paniculataareplacedatthemiddleofthepetiole.

excLuded NAmes

Galphimiasect.CosmogalphimiaNied.,ArbeitenBot.Inst.Königl.LyceumHosia-numBraunsberg5:23.�9�4.—Notvalidlypublished(ICBNArt.22.2).

Galphimia pubescens(Adr.Juss.)Herter,Fl.illustr.Uruguay�4(��):49�.�956.—Notvalidlypublished(ICBNArt.33.3).

Galphimia caucaSpreng.,Syst.veg.2:385.�825.=Caucanthus edulisForssk.

Galphimia chrysophylla (H. B. K.) Spreng., Syst. veg. 2: 385. �825. = Byrsonima chrysophyllaH.B.K.

Galphimia longifolia H. B. K., Nov. gen. sp. 5: �73. �822 [“�82�”]. = Lophanthera longifolia(H.B.K.)Griseb.

Galphimia mollisH.B.K.,Nov.gen.sp.5:�73.�822[“�82�”].=Tetrapterys cotoneasterAdr.Juss.

Thryallis hirsutaSessé&Mociño,Fl.mex.��9.�894;ed.2,�09.�894.—TheSessé&Mociñoherbarium(atMA)doesnotincludeacollectionlabeledwiththisname.Mostlikely,thedescriptionwasbasedonaspecimenofEchinopterys eglandulosa(Adr.Juss.)Small(W.R.Anderson,pers.comm.).


ACKNOWLEDGMENTS

IamindebtedtoWilliamR.Andersonforhisgenerousadviceandassistance,andtoJohnH.BeamanandAlanPratherfortheirhelpwithbibliographicmatters.KarinDouthitdrewthebeautifulillustrations.IthankthecuratorsofthefollowingherbariaformakingtheirmaterialofGalphimiaavailabletomeandfortheirassistanceandcourtesiesduringvisits:A,BM,BR,C,CAS,CEPEC,CHAPA,CTES,DS,DUKE,ECON,ENCB,F,G,GH,GOET,GUADA,HB,IBUG,IEB,K,L,LL,MBM,MEL,MEXU,MICH,MO,NY,P,RB,S,SD,SP,TEX,UAMIZ,UB,UC,US,W,WIS,XAL,ZEA.ThisstudywassupportedinpartbyagrantfromtheNationalScienceFoundationtotheUniversityofMichigan(DEB-0543909).

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APPENDIX

species of GALphimiA occurriNG iN mexico, Listed by stAte

Aguascalientes: G. glaucaBajaCaliforniaSur: G. angustifoliaChiapas: G. speciosaChihuahua:G. floribunda, G. vestita.Coahuila: G. angustifoliaColima: G. grandiflora, G. langlassei, G. montana, G. tuberculataDurango: G. angustifolia, G. floribunda, G. montanaGuanajuato: G. glauca, G. multicaulisGuerrero: G. calliantha, G. glandulosa, G. hirsuta, G. langlassei, G. mirandae, G. montana, G. paniculata,

G. radialis, G. tuberculataHidalgo: G. glauca, G. gracilisJalisco: G. floribunda, G. glauca, G. grandiflora, G. langlassei, G. mexiae, G. montana, G. multicaulis, G.

paniculata, G. tuberculataMéxico: G. glandulosa, G. grandiflora, G. multicaulis, G. paniculataMichoacán: G. glandulosa, G. grandiflora, G. langlassei, G. montana, G. multicaulis, G. paniculataMorelos: G. grandiflora, G. paniculataNayarit: G. floribunda, G. mexiae, G. montana, G. paniculata, G. tuberculata, G. vestita


NuevoLéon: G. angustifolia, G. glaucaOaxaca: G. arenicola, G. calliantha,G. elegans, G. multicaulis, G. oaxacana, G. paniculata, G. sessilifolia,

G. speciosaPuebla: G. gracilis, G. mirandae, G. paniculata, G. speciosaQuerétaro: G. glauca, G. gracilisSanLuisPotosí: G. angustifolia, G. glauca, G. gracilisSinaloa: G. angustifolia, G. floribunda, G. glandulosa, G. montana, G. tuberculata, G. vestitaSonora: G. angustifolia, G. floribunda, G. vestitaTamaulipas: G. angustifolia, G. glauca, G. gracilisVeracruz: G. gracilis, G. speciosa, G. glandulosa(?)Zacatecas: G. glauca, G. montana, G. paniculata

INdExTOSPECIES

G. amambayensisC.Anderson51G. angustifoliaBenth.38G. arenicolaC.Anderson23G. australisChodat47G. brasiliensis(L.)Adr.Juss.44G. callianthaC.Anderson71G. elegansBaillon58G. floribunda C.Anderson77G. glandulosaCav.25G. glaucaCav.55G. gracilisBartl.12G. grandifloraBartl.73G. hirsutaCav.19

G. langlassei(S.F.Blake)C.Anderson28G. mexiaeC.Anderson31G. mirandaeC.Anderson36G. montana(Rose)Nied.16G. multicaulisAdr.Juss.64G. oaxacanaC.Anderson17G. paniculataBartl.67G. platyphyllaChodat53G. radialisC.Anderson33G. sessilifoliaRose34G. speciosaC.Anderson60G. tuberculata(Rose)Nied.21G. vestitaS.Watson42

revision of galphimia (malpighiaceae)

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