Excerpt'from'. Floweing Plantsof the Neotroplcs,ed. N. Smith,S A. Mori.A. Henderson,D. W. Stevenson,and S. V. Heald.2004. (Malpighia Family)

Wrllrau R. ANontsoN Figures1 15, 116; Plate28 @*ffi*. o ,, vines, and a few perennralherbs

o indument of unicellular hairs, usually 2-branched

o leaves usually opposite, simple, many with large multicellular glands on pettole, abaxial surface, or l.- margtns 4 or all 5 a\ . flowers mostly bilaterally symmetrrcal; =.\-.) sepals usually bearing 2 large abaxral glands petals " -l 5, mostly clawed;starnens mostly 10; ovary \)i superior, mostly tncarpellate with the sVes dtstrncf, ovules 1 Per locule 1/ . mostly schizocarpic and winged in vines. fruits \', mostly unwinged and dry or fleshy in shrubs and trees s,##.w" t';j I

l::l l3mh Numbers of genera and . Currentlyapproximately 65 generaand 1,260species of Malpighiaceaeare recognized worldwide.Fifiy generaand approximately1,1 l0 speciesoc- cur only in the Westem Hemisphere,except fbr two speeies found also in western Africa (.Heteropterysleona and Stig- WWA li,I14 E ll' maphyllctnbannisterioides). The largest genera,all of them mostlyor entirelyNeotropical, are Banisteriop,lls (94 species), Bunchosia(68), Byrsonima(.121), Heteropre:s (136),SrlS- Figure115. MALPIGHIACEAE.Byrsonima aeruga. A. Stemwith Nodeshowing petiole bases and con- maphyllon(92), and T'etrapterys(70). Sixteen generahave leavesand inflorescences, B. nateintraoetiolar stipules. C. Lateralview of flowerbud with sess e onlyone species. pedicelsubtended by two shortbracteoles and one long, reflexed bract;note oil glands on calyx.D. Laieral(left) and apical (right) views Distribution and habitat. The Malpighiaceaehave little of fiowers.E. Adaxial (left) and lateral (right) views of stamens.F. Lat- tolerancefor extreme cold and, therefore,show a typically eraiview of gynoecium.G. Partof infructescencewith one fruit re- tropicaldistribution. In the WesternHemisphere, a few spe- moved(center) and transverse section of fruit(left), showing three cies reach southernFlorida, Texas, New Mexico, and Ari- seedsin a commonstony endocarp surrounded by fleshymesocarp; zona,and in the south the family is rnoderatelywell repre- detailof sericeousapex of fruit(above right). Reprinted with permis' sentedto 35oS, the latitude of Buenos Aires; only a f'ew sionfrom Mori et al.(2002). Arlist: Bobbi Angell. speciesoccur farther south,to about 39'S. A similar pattern existsin the paleotropics,but it is much less dramatic there becausethere are only about 150 speciesin the Eastern Byrsonimahas many speciesin southernVenezuela and the Hemisphere. Guianasand rather f'ew in Ecuador,while the reverseis true The great center o1 diversity of Malpighiaceaeis South for Stigntaphr-llon.Thefamily is well representedin the trop- Americanorth of the tropic of Capricorn.Except tbr Chile, ical West Indies,but not in the extratropicalBahamas. which has only two species,all the countries in tropical The Malpighiaceaehave adaptedto diversehabitats in the SouthAmerica have substantialnumbers of species,Brazil neotropics,including wet, mesic, and seasonallydry forests, having far more than any other. In the cerrados of Minas shrubbysavannas, and grasslands.Few grow high in the An- Gerais,Brazil, fbr example,there is an astonishingdiversity des becausefew can toleratethe temperaturesat higher ele- of Malpighiaceae.Some specieshave extensivedistributions vations.and few havesucceeded in extremedeserts. Although (e'9.,Baniste.riopsis muricata, from Mexico to Argentina); they are not rare in wet fbrestslike thoseof Amazonia,Mal- Dutmany more speciesare restrictedto much smaller re- pighiaceaeare more numerousin well-drainedsavannas like gions,and narrow local endemismis fairly common.Patterns those of the Brazilian Pianalto,and in the shrubby associa- of speciesrichness differ fiom genusto ;for example, tions lbund in upland habitatsof the VenezuelanGuayana.

MALPIGHIACEAE.229 Family classification, Cronquist lbllowed earlier workers ing most of the wing-fruited genera.6) Winged fruits mny and put the Malpighiaceaein his order Polygalales,on the have evolved several times or once; but if only once. the basisof morphologicalsimilarities. That placementis not sup- wings were * completely lost in severaidescendant clades. ported by molecular data, and recent authorshave recognized 7) There is a strongcorrelation between the vining habit and the morphologically diverse order . comprising winged fruits versus the shrubby or arborescenthabit and Malpighiaceae,Euphorbiaceae, Passifloraceae, Violaceae, and unwingedfruits, but the phylogeneticpath to that correlation other families. Both morphological and molecular data agree is probably not simple, suggestingrepeated evoiutionary that the family is monophyletic, but so isolated that one still convergenceon an adaptiveassociation of habit and method cannotsay which familiesare closestto it. of dispersal. The infiafamilial classificationof the Malpighiaceaeis in As morphologicaldata are combinedwith more and better a stateof flux. Earlier authorsdivided the family on the basis molecular data, the resolutionof the phylogeny of the Mal- of fiuits: two subfamilieswith the fruits unwinged versus pighiaceaeshould improve enoughto aliow the proposalof winged or bristly, with each subfamily then divided into a new infrafamilial classificationthat will be both usetuland tribes defined principally by details of the fiuits. Some of phylogeneticallyaccurate. these groupings have been supportedby later studies that incorporatedother aspectsof the morphology and molecular data, but in many casesthe family is proving to be much more complicated.William R. Andersonasserted many years ago that the three genera with similar fleshy fruits (Br.in- chosia, Byrsonima, and Malpighia) were quite unreiatedto each other, and chloroplastDNA supportsthat conclusion. Bristly fruits like those found in Echinopterys,HenLeophy- tum, Lasiocarpus,Ptilochaeta, and Tricomaria, rather than marking a naturaland convenienttribe, clearly evolvedinde- pendently at least three times in the family. While it does seem likely that the relative developmentof the lateral and dorsal wings on the samaraswill continue to be useful in /: grouping many generaof wing-fruited vines, those "rules" have been violated a number of times. For example, Hetero- pterys,with a large dorsalwing, belongsin a cladeof lateral- winged genera(a molecularresult supportedby the nonfruit moryhology); on the other hand, Diploptery-sand Cordobia, both with the lateral wing dominant, are sistersto genera with a dominantdorsal wing, and this result is supportedby nonfiuit morphology.A number of generathat are morpho- logically isolatedare still not satisfactorilyplaced by mole- cules or moryhology, and the large wing-fiuited clade that comprisesthe vast majority of the family's generaand spe- cies is still poorly resolved.Nevertheless, certain things can now be statedwith some confidence:1) The Malpighiaceae probably originatedin the Westem'Hemisphereand the Mal- pighiaceaeof the EasternHemisphcre are probably derived fiom at leasteight independentdispersal events. 2) The origi- nal Malpighiaceaewere very likely shrubsor trees,with the 't1 habit of woody vines evolving betweenone and four times. ,._.1 3) The earliestMalpighiaccae probably producedfruits that lacked flesh,wings, or bristles;dispersal by water may have Figure 116. MALPIGHIACEAE.Jubelina rosea. A. Stem with leaves and beenancestral. 4) The basechromosome number in the fam- and inflorescence,B. Detailof abaxialleaf surface showing glanC stalked,bifurcate hair, C. Obliqueapical view of flower shor,u,ng ily was probablyx = 6; however,all but the lowest branches bud spatulatebracteoles and sepals,each Jateral sepal bearing one arge of the family seem to have numbers based on r = 10, abaxialgland, D. Oblique-apicalview of flower,the posteriorpetal and that shift (perhapsby aneuploidyfrom an ancestorwith uppermost.E. Adaxialview of parl of the androecium;the centralsta n= 12) probably occurredat about the time of the origin of men would be oppositeone of the postenor-lateralpetals, F. Lateral winged fruits. -5)The pollen was originally radially symmet- view of gynoeciumwith antenorstyle in center,G. Abaxialvrer'v rical, probably tricolporate,and the shift to globally symmet- (above)and medialsection of fruit(below), Beprinted with permssion rical pollen happenedonce, at the baseof the clade contain- -orn Mo.i er al. (2002).Artist: Bobbr Angerl.

230 . N,,1ALP GHTACEAE Features of the family. Habit; Trees,shrubs, erect or trail- cidal, or at least not sufficiently so to allow the seed to ing subshrubsor herbs with perennial undergroundstems, escape);endosperm absent. andwoody to herbaceous,perennial vines that climb by twin- The family is easierto recognizein flower than in fruit ing stems. Latex rarely present (Galphimia, Lophanthera, becausethe flowers are so uniform, but once one has placed Spnchea, and Ve rrucularia). lndument: hairs always unicel- a in the Malpighiaceae,it is much easierto identify it lular,usually 2-branchcdand medifixedor submedifixed,ba- to genuswith fruits than with flowersbecause the generaare sifixedor stellatein a few genera.Stipules usually present definedprimarily by charactersof the fruits. besideor on the petiole or axillary to it, distinct or variously connate,minute to more than l4 cm long, absentin some Natural history. As noted above, most Neotropical Mal- generaor species.Leaves usually opposite and decussate, pighiaceaeare recognizcdeasily by the large paired glands iometimeswhorled, suboppositeor alternatein a few genera on the abaxial surfaceof the sepalsand by the clawedpetals. or species,often bearing large multicellular glands on the StefanVogel has shown that thesestructures are adaptations petioieor blade (usually the abaxial surface or margin) or for pollination by oil-collecting bees, which land on the both;blade simple, mostly entire, rarely lobed. the margins flower, reach between the petals (hence the importanceof nevertruly toothed but sometimespseudodentate or ciliate the spacelefl by the claws), and collect the oil producedby at the location of marginal glands or stout bristlelike hairs. the glands. They mix this oil with pollen and pack it into lnflorescencesterminal or axillary, very diverse,most often brood cells with one egg each,and the mixture eventuallyis racemoseor paniculatebut with the flowers often ultimately consumedby the growing larvae. The flower producesno bornein umbelsor corymbsof 4 or more. Flowers subtly to sugarynectar so, exceptfor the oil, pollen is the only reward stronglybilaterally symmetricalin most Neotropicalspecies for pollinators.Malpighiaceae that have lost the calyx glands with the plane of symmetrypassing through the anterior(of- must rely on pollen to attractpollinators, and in some such reneglandular) sepal and the posterior(often erect and dif- groups (e.g., Galphimia) the anthersare enlarged.Almost ferentiated)"flag" petal,a few radially symmetricalor nearly nothing has beenpublished on the pollination of theseeglan- so,mostly bisexual, a few generadioecious or functionallydi- dular species.William R. Anderson has postulatedthat the oecious,hypogynous except perigynoustn Barnebya, small Malpighiaceaenow in the EasternHcmisphere descended (about6 mm in diameter)to fairly large (about4 cm diam.); fiom severalspecies that emigratedfrom Sor.rthAmerica to sepals5, mostly imbricatein bud, the greatmajority of Neo- Africa after the separationof the continents.The oil-bees tropicalspecies bearing 2 (rarely only l) large, multicellular, that pollinate Neotropical Malpighiaceaedid not reach the abaxialglands on all 5 sepalsor on lateral4 (most Paleotrop- EasternHemisphere, so it is not surprising that the calyx ical specieswith calyx glandsmuch reducedin number and glandsof EasternHcmisphere species are mostly reducedor sizeor absent);petals 5, distinct, mostly clawed, alternating absent. with sepals,imbricate, the posteriorinnermost and I of ante- The majority of Malpighiaceaein the Wcstem Hemisphere rior-lateralpair outermost,most often yellow, pink, or white, have winged or bristly iruits adaptedfor dispersalby wind. sometimesother coiorsbut very rarely blue; androeciumusu- Three Neotropical genera(Buncho.sia, Byrsonima, and Mal- ally of l0 stamensin a single whorl, the stamenssometimes pighia) produce edible, fleshy fruits that presumably are fewer.up to 15 in Lasiocarpu.s,borne on receptaclcbetween bird-dispersed.Several genera, mostly treesthat grow along perianthand gynoecium,the filamentsalways present,short Amazonian rivers, have smooth aerenchymatousfruits that to long,alike or heteromorphic,distinct or partially connate, surelyare dispersedby water,and scatteredspecies in genera theanthers alike or heteromorphic,4-locular, mostly iongitu- with winged fruits have more or less completely lost their dinallydehiscent along inner edgeof eachlocule, with apical wings and become adaptedsecondarily for water dispersal. or subapicalpores or very short slits in a t'ew genera;gynoe- Finally, a number of genera,most relatively near the baseof ciumsuperior, usually comprising3 distinct to connatecar- the phylogenetictree, produce small, smooth,dry fruits with- pels,mostly I anterioron plane of symmetryand 2 posterior out any obvious adaptationfor dispersal.These f'ruits, gener- on each side of plane of symmetry, the carpeis only 2 in ally 1 to severalmillimeters in diameter,presumably are dis- severalgenera and very rarely 4, mostly all fcrtile, eachfer- persedby wind or rainwater.It is difficult to understandhow tile loculecontaining I pendentanatropous ovule, the slyles some oI those groups (e.g., Pterandrn) have achievedtheir mostlyas many as carpels and distinct, but connateor re- extensivepresent-day distributions. ducedin numberin a few genera.Fruits fleshy or dry; fleshy truttsmostly an indehiscentdmpe or beny, yellow, red, blue, Economic uses. The Malpighiaceaeare of modesteconomic or black;dry fruits indehiscentin a few genera.but schizo- importance.One of the specieswrth fleshy fruits, Malpighia catpicin most, splitting apart into mericarps(typically up to emarginata (often erroneouslycalled M. ptrnicffblla), has J);dry fruits or mericarpsof somegenera nutlets with smooth long been cultivated for the red cherrylike fruits, which are walls.those of some generaor specrescontaining aeren- rich in vitamin C. It bearsmany common names,of which cnyma.but mostbearing wings or bristles.Seeds I per loc- the onesmost frequentlyencountered are acerola and Barba- uleor mericarp,never released (i.e., dehiscencenever loculi- dos cherry. In recentyears, manufacturers of vitamins have

IvALPIGHIACEAE. 231 ANnnnsox, C. 199'7.Monograph ol Stignro- taken to adding vitamin C from M. entorginaldto their con- References. (Malpighiaceae).S,r;s/. Bot' Monogr. 51:1-313.AN- coctions in order to make them more "natural" (and core- phytton Revision of Pterandra'(Malpighiaceae). spondinglymore expensive),and commercialplantations fbr DERSoN,C. 1997. Michigan Herb. 2l I-27. ANlensoN, W' R. the cultivation of the speciesexist in Mexico and probably Contr. Univ. Byrsonimoideae,a new subtamily of the elsewhere.The fruits of Byrsonimo crassifolia, which look 1978 1"191'7"1. 7:5-18. ANletsoN, W. R. 1979. like yellow cherries,are consumedcommoniy in Mexico, Malpighiaceae.Leanclra in neotropicalMalpighiaceae' Biotropicct Centrai America, and northern South America. In Spanish- Fioral conservatism W. R. 1981.Malpighiaceae' ln The speaking areas, the common namc is usually nance or ll:219-23. ANosnsoN, Highland-Part XI. Mem. New'Yctrk nanche,whereas in Brazil it rs murici or murucf' Bunchosia botany of the Guayana ANoansoN,W. R. 1990' The origin glandutiJbrahas a pleasant-tastingfruit the size of a small Bot. Garrl. 32:21-305. evidence from morphology. plum and surely has been cultivated by indigenouspeoples of the Malpighiaceae-The Bot. Gard. 64:210-24. ANopnsoN, W. R. in South America for a very long time; it is not known fiom Mem. New York numbersof neotropicalMalpighiaceae. unequivocallywild populations.Fruits of various other spe- 1993. Chromosome Herb. 19:341-54.ANpEnsox' W. R. cies of Bunchosia,Byrsonima, andMalpighia are consumed Contr. [Jniv. Mic'hiean In Flora of the VenezuelanGuatttnc4 from Mexico to Brazil, and the (a11shrubs or small 2001.Malpighiaceae. K. Yatskievych,and B. K. Holst,6:82-185. trees)often are grown in dooryards,both as ornamentalsand ecls.P. E. Berry, Press,St' Louis.BlnN'q'1. H. Y., fbr the fiuits. MissouriBotanical Garden 1994. Malpighiaceae'In Especies A numberof other Malpighiaceaeare piantedas ornamen- AND J. E. Connsn.{ Q. de los paises del Convenio Antlrds tals, and some are availablein warm regionsfrom nurseries' vegetalespromisorias de Bogotd,D. E., Colombia:Secre- The commonestof these is Gctlphimiagracilis (often mis- Bello,Tomo X. Santaf6 Andr6sBello. C.quenoN.K. M., identif,edas G. glauca), a with bright yellow flowers taria Ejecutivadel Convenio W. R. ANlrRSoN,AND H. G' HIr-ls. 2001. that give it common nameslike Lluviade ttro, ramittt de oro, M. W. Cuese, of Malpighiaceae: Evidence tiont and spray of gold. Other Neotropical species popular in Molecular systematics sequences'Amer. -I. .Bot.88:1811 -62- warm gardens ancl greenhousesate Malpighia toc'cigera plastid rbcL andmatK Phylogeneticsof seedplants: (Singaporeholly), Stigmaphyllon ciliatum (Brazilian gold Cn.qsE,M. W., Er AL. 1993. analysis of nucleotide sequencesfrom the plastid gene vine), and S. floribundurrr(orchid vine). LophcLntheraluctes- an Bot. Gard. 80:528-80. Dlvts, C. C.. cens. a handsometree with long inflorescencesof yellow rbcl. Arut. Missouri ANDM. J. DoNocHus. 2001.Phylogeny flowers. is known in the wild from only one small area in W. R. ANocRSON. Evidence from chloroplast ndhF' and easternAmazonia, but it thrives when planted along busy, of Malpighiaceae: Amer' J. Bor. 88:1830 1846' polluted streets and is now popular in towns and cities trnL-F nucleotidesequences' Banisteriopsis,Diplopterys (Malpighia- throughoutBrazil. Two speciesnative to the EastemHemi- G.qrr.s, B. 1982. 30l-237. Julo, W' S ' C S' sphereare cultivatedoccasionally in the neotropics,Hiptage ceae).F/. Neotrop.Monogr. ANDP. F. Srnvr'Ns.1999 Plant benghale nsi s and Trist ellate ia (tustalasiae (ba gnit)' Many C.qNtpeEI-r,E. A. KsI-rocc, Approach' l-'16:l Suth- other membersof the family are very showy and, hence,are Systematics:A Phylogenetic Ii]-xvi' Associates,Inc. NtnosNzu. F' 1928' alsocandidates for cultivation' erland. MA: Sinauer PJlanzenreich'Series IV' 141(Hefte At leastone memberof the family , Banisteriopsiscaapi, is Malpighiaceae.In Das gl, Wilhelm Engelmann'Vec'r' A S" a potent hallucinogenand has becomefamous among those 93,94):1-870. Leipzig: R. ANosnsoN.2002. Occunence interestedin drug plants under its scientific name and such M. A. C,\srno, AND W. of latex in the Malpighiaceae vernacularnames as ayahuosca,cuapi, andyag6. It is culti- and phylogeneticsignificance Vocnr, S. 1974.Olblumen vated widely in AmazonianSouth America, where it is used Amer. J. Bot. 89(11):1125_29. Tropische und subtrt'fisclrc by native populationsas one ingredientin the preparationof und 6lsammelnde Bienen. Mainz: Akademieder Wissenschaf- a beveragethat is said to produce spectacularmulticolored Pflanzenweh7:l1l-261 . visions after somehours of preliminary vomiting. ten und der Literatur.