REVISION OF GALPHIMIA (MALPIGHIACEAE) Christiane Anderson University of Michigan Herbarium 3600 Varsity Drive Ann Arbor, Michigan 4808-2287 ABSTRACT. Galphimia (Malpighiaceae, Byrsonimoideae, Galphimieae) comprises 26 species of suffrutescent herbs, shrubs, and treelets. Four species (G. amambayensis, G. australis, G. brasiliensis, G. platphylla) occur in South America south of the Amazon basin, and the remainder in Mexico; one species (G. angustifolia) extends into Texas and one (G. speciosa) into Central America. One species, G. gracilis, is cultivated worldwide in warm regions. The genus is characterized by bilaterally symmetrical flowers with yellow to red petals; in eight species the petals are persistent. The sepals lack oil glands, but a few species bear calyx glands that resemble the leaf glands. The fruit is a globose schizocarp breaking apart into three small -seeded cocci.The base chromosome number is x=6. For each species a full synonymy, detailed description, and range map are provided; all but two species are illustrated. Three new species (G. calliantha, G. floribunda, G. speciosa) are proposed. INTRODUCTION The Neotropical genus Galphimia Cav. (Malpighiaceae, tribe Galphimieae) comprises 26 species of large herbs, shrubs, and treelets of mostly drier habitats. Most species occur in Mexico, with one extending to Texas and one ranging to northern Nicaragua, but four are restricted to South America, south of the Amazon basin. Galphimia is characterized by flowers with yellow petals (often suffused with red) borne in racemes, either solitary or arranged in panicles; in eight species the petals are persistent in fruit. The sepals lack the large oil glands typically found in other New World Malpighiaceae, but in a few species the calyx has one or more small glands that resemble the leaf glands. Ten fertile stamens surround a tricarpellate ovary with three subulate styles. The globose fruit is a schizocarp that breaks into three -seeded cocci. The history of Galphimia is one of confusion, caused in part by misinterpretation of the earliest described taxa, competing generic names (Galphimia Cav. vs. Thryal- lis L.), and misapplication of various names to one cultivated species (G. gracilis), but also by the superficial morphological similarity of many of the species. The showy inflorescences catch the eye of collectors, who often label all specimens from Mexico as G. glauca or G. gracilis, and those from South America as G. brasiliensis. This revision builds on preliminary studies of the genus (C. Anderson 2003, 2005a, b) and clarifies the circumscriptions of the species and associated nomenclature. It is intended as a basis for detailed future studies. Many species are still known from only few collections. Questions remain concerning the phylogeny, phytogeographic history, pollination and dispersal strategies, ploidal levels, as well as developmental aspects, such as the nature of the unusual calyx glands, the shift to persistent petals, and the suppression of the peduncle. TAXONOMIC HISTORY The nomenclature of Galphimia and the application of names have been prob- lematical, partly owing to uncertainty about the status of the generic name Thryallis L. and partly to doubtful interpretation of original descriptions without access to type Contr. Univ. Michigan Herb. 25: –82. 2007.
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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.
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.
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.
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).
The margin is entire, but in species bearing tuberculate hairs the margin ofmatureleaves,atleastproximally,bearsthepersistenttubercles.ThelaminasofG. arenicolacommonlybearelongate,multicellular,epidermalprocessesinadditiontosuchtuberclesalongthemargin.Galphimia oaxacanaisdistinctiveinthatthemarginformsathickened,narrow,light-coloredband.
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).
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.
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
2007 C.ANDERSON:GALPHIMIA 7
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).
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.
GalphimiaCav.,Icon.5:6�.�799.Malpighia[subgenus]Galphimia(Cav.)Persoon,Syn. pl. �: 506. �805.—Lectotype, designated by O’Donell and Lourteig,�943:Galphimia glaucaCav.
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
�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
pedicels;sepals3.5–4.2mmlong;laminasellipticaltobroadlyso, obovate, rhombic, ovate, or suborbicular, the base oftendecurrent;Guerrero,Puebla. �2.G. mirandae
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
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.)
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.
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.
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
�8 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25
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.)
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.
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.
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!).
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.)
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).
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.
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.
Shruborsubshrubto3m;stemssparselytomentulosewhenyoung,soongla-brous.Vestureofallvegetativepartscomposedofsessile,straightorwavytocrisped,reddishbrownhairs0.2–0.8mmlong.Laminasof the larger leaves5–�4cmlong,�.5–4.8cmwide,ellipticalornarrowlysotolanceolate,apexacuteorapiculate,base
2007 C.ANDERSON:GALPHIMIA 29
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,
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.
2007 C.ANDERSON:GALPHIMIA 3�
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.
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.
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).
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.
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.)
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.
2007 C.ANDERSON:GALPHIMIA 37
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.)
38 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25
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. Collected in flower and fruit from April through November(–December),inSonoraalsoinJanuaryandFebruary,inBajaCaliforniaSurfromAugustthroughMay.
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.
42 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25
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.
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
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
Jussieu (�833) was the first to realize that Linnaeus’s Thryallis brasiliensis belongsinGalphimiaandisnotrelatedtospeciesassignedtoThryallisMart.,buthisdescriptionofG. brasiliensiswasbasedonspecimensofG. australis.
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.
Galphimia amambayensis is known only from the type collected in easternParaguay(Fig.�9).It isreadilyseparatedfromtheotherSouthAmericanspeciesbyitsdenselytomentoseinflorescenceaxesandcalyx.LikeG. andustifoliaandG. brasiliensis,ithasapiloseovary,butinthedistal�/3thevestureispresentonlyonthesuturesandadjacenttothem.Thepetalsbearscatteredhairsabaxiallythougharesometimesglabrous,butintheseinstancethehairsmayhavebrokenoff.
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. 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);
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.
58 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25
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).
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.
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.
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.)
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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.
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”).
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.
2007 C.ANDERSON:GALPHIMIA 63
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
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G. speciosaareG. elegans,withdistalleavesabaxiallypubescent,leafglandsbornenearthemiddleofthepetiole,andsmalleranthers(�.8–2mmlong),andG. multi-caulis,withrudimentarytoveryshortpetioles(eveninthelargestleaveslessthan�cmlong).
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
2007 C.ANDERSON:GALPHIMIA 65
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.)
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
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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.)
2007 C.ANDERSON:GALPHIMIA 69
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�.
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
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.
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.
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,
2007 C.ANDERSON:GALPHIMIA 75
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.
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:
76 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25
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).
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”).
Distribution(Fig.30).Mexico(Sonora,Chihuahua,Sinaloa,Nayarit,Durango,Jalisco); on dry slopes in deciduous forest, oak forest, and pine-oak forest;500–�300m.
34�–354.———.200�.Malpighiaceae.InFlora of the Venezuelan Guayana,ed.P.E.Berry,K.Yatskievych,andB.
K.Holst,6:82–�85.St.Louis:MissouriBotanicalGardenPress.Baillon,H.�874.Galphimia elegans.Histoire des plantes,5:43�,figs.429–435.�874.Bartling,F.T.�840.DeGalphimiismexicanisannotationes.Linnaea�3:549–558.Blake,S.F.�9�7.Descriptionsofnewspermatophytes,chieflyfromthecollectionsofProf.M.E.Peckin
Galphimia brasiliensis(Malpighiaceae).Amer.J.Bot.88:�935–�944.Cavanilles,A.J.�799.Galphimia. Icones et descriptiones plantarum,5:6�–62,t.489.———.�80�.Galphimia glandulosa.Icones et descriptiones plantarum,6:43,t.563.Chodat,R.�890.MalpighiacéesduParaguay.Arch.Sci.Phys.Nat.,sér.3,24:495–50�.———.�892.ContributionsalafloreduParaguay.V.Malpighiacées.Mém.Soc.Phys.Genève3�,part2,
393–395.Kunth,C.S.�822[“�82�”].Galphimia.InNova genera et species plantarumbyF.W.H.A.Humboldt,A.J.
A.Bonpland,andC.S.Kunth,5:�72–�74.Paris.Kuntze,O.�89�.Thryallis.InRevisio genera plantarum,�:88.Kyhos,D.W.�966.Galphimia glauca. In“Documentedchromosomenumbersofplants.”Madroño�8:
———.2000.Botanical results of the Sessé & Mociño expedition (1787–1803). VII. A guide to relevant scientific names of plants.Pittsburgh:HuntInstituteforBotanicalDocumentation,CarnegieMellonUniversity.
67:95–96.Rose,J.N.�897.StudiesofMexicanandCentralAmericanplants.Contr.U.S.Natl.Herb.5:�09–�44.———. �909. Studies of Mexican and Central American plants—No. 6. Contr. U.S. Natl. Herb. �2:
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
82 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25
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