Leandra 7: 11. 1978 ["1977"]
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I,EANDRA VI-VII DEZI]MBRO DE 1977 Ne7 I}YRSONI},{OIDTAE, A NI]W SL]BF'AT{ILY OF THI] I{ALPIGHIACEAI \flilliam R. Anderson,r 'I'he Malpigliiaceaecan easiiybe dividcd into In'o groui)sby their fruits, which are usually either winged or unwinged. Specieswith winged fruits are generally but not always vines, while speciesri'ith unwinged fruits are usually shrubs or trees. On tl.rebasis of thesecharacteristics, plus the shapeof the reccptacle,the family N'as divided two subfamilies by Franz Niedenzu, rvho published his rnonographof the family in 1928. Niedenzu used thc nanrc Pyramidotoraefor the group with rvinged fruits and Planitoraefor the groulr rvrtir unwinged fruits. Morton(1968) pointed out that the Code of Nomenclaturerequires that tl.renamcs of subfamiiiesbe basedon namesof included genera. lior Nieclenzu'sPyramido- toraehe proposedthe nane Gaudichauclioideae;for tire Planitoractirc naruervould hnve to be Malpighioideaebecausc the subfamrlyas circumscribcdby NiedenzLr irrcludedthc genus Malpighia, thc type o1' the family. The subfamily Malpighioideaesensu Morton is rnostly a n:rturai assemblage of genera linked together by nutnerous ciraracters. Horvcver, ccrt,rin generr included by Niedenzu do nclt scem to me to belong n'itlr thc othcrs, ancl I would excludethen.r. Unfortunatcly, orre of the excludedgcrlera is h{alpighia,and wherr it is removed the nanreof the subfamily goes rviih it. Therefore it is necessar)' to describe a ne\\r subfarnily i:or the rcmaining genera. In this paper I shall describeand circumscribethat subfamilt,lncl tlrcn discussnly reasonsfor excluding It{alpigliia ancl scveralotlrcr genera. ' Tliq ljnies.sity of Michigon Herbarium, Ann Arbor, Michigon 48109 USA. Reseorcb supported in port by Ncrlionol Science Foundation Grcrnt GB-373i4. Byrsonimoideaew R Anderson, Ir{alpighiacearutnsubfaru no\r. Arbores ct f ruiices, rafo suffruticcs. Stipulae bene evolutae, plerurnque intrapetiolares,liberac vel connltac. Inflorescentiathyrsus ex cincinnis2-1O-floris constans,vel pseudoracemusvel spica vcl fasciculus(re vera ex cincinnisunifloris -+- constans).Gemrna floris saepecircinata. Antherae inter se similares.Pollen plerumcluetricolporatrrnt, raro tetracolporatum (in Galphimia Sect. Galphimia parasyncolporatum),10-30 p in diametro. Styii plerumque3, subulati,stigmate minuto (in Spachea2-3, crassi,stigmate truncato vel subpeltato). Fructus caroosus vel siccgs,nunquam alatus nec setifer, ex 3 carpellis liberis vcI z'3 lnericarpiis constansvel compositusloculis 1-1 -j (-hronrosotnatttm ntllncrtls: t) 6 vel tl' Typus. Byrsonima1.. c. Richarclex Flun'rboldt,Bonpland .\ Kttnth, Nov Gen. Sp. (.1red) . ):I47 1s21(1822)' plants of subfamilyByrsonirnoideae are all nativesof the tropics anclsubtropics of the New .J(orld. There are no vines among thetn, irnd I believethat the vining habit 6as never evolvedin this group, This is in rnarkedcoutrast to many other genera(e. g. Banisteriopsis,HeteroPterys, Tetrapterys), in'n'hich the vining habit is probably ancestraland extant shrubs are probably derived. The best single characterfor recognizingthis subfamily is the presenceof tlrrce subulate styles sith minute stigmas; the only excePtionis Spachea,which cannot be excluded b:causeit seemsto be closelyrelatecl to Lophanthera. Smali tricolporatepollen is anoiherimportant charactcrthart ur.rites tiris subfamily,and anotheris a chrotnosome nurnberof n - 6 or n : 12. Almost all chromosomenuntbers knorvn for other New World lr{alpighiaceaearc n : 10 or a multiplecf 10,or eneuploidderivatives Of such rnultiples.However, very few chromosomecounts have been pubiishedfor more lire lvfalpighiaceae,ancl this gencmlizationnray or miy not be supportedrvllen data are available. It would be especiallyinteresting to have a chrotttosomecount for the troublesomegenus Spachea. any No member of this subfamilyhas the fruit rvingedor setose,nor is there evidenceof reductionfrom a *,inged fruit comparableto the u'inglessspecies found rn Banisteriopsis,Heteropterys, Stigmaphyllon, I\{ascagnia' etc. In tnost generathe fruit is probably dispers.,iby rvater or blown about with other detritus-likeseeds and fruits. Byrsonimahas acirieveda rnaior distinction by evolving a fleshy, bird- dispersedfruit, and it is probably no coincidencethat Byrsonimais the only geuus in ihe subfamilyn'ith more than 1i species.\flith the adventof small birds thete has probably been strong selectionin various lines for fleshy fruits. and it is my opinion that suchfruits have evolvedseparately and repeatedlyin the Malpighiaceae. It happenedonly once in this subfamily,in the ancestorof Byrsonima,which pro- bably had a friut like that of Blepharandra. Somewhatsimilar fruits also evolved in the ancestorsof Malpighia and Bunchosia,but other characteristicsof those generado not suPpoft a probable origin in this subfamily,and they are therefore r:xcluded. KtsY TO THI] TRIBES OT BYRSONIIVIOIDEAI] I (-;rrpelsconnate in flower, bearing apical styles. 2 . F'ruit quite indehiscent,neither opening nor breakingapart into mericarps I . Byrsonimeae 2. F-ruit breaking apart into dry 1-seededrnericarps, the nrericarpsoften dorsally loculicidal z. Galphimieae I . Carpels free from each other, borne on a flat or slightly pyramidal torus, bearing ventriflxed sub apical to almost basal styles f. i.,,,"nrt "..u. i ByrsonimeaeW. R. Anderson,trib. nov. Gemma floris saepecircinata. Glandescalycis l0 vel nullae. Staminat0 (vel rn Diacidia spp. 9-6), aniherisglabris vel piliferis, inter se -+ similaribus. Carpella 3 omnino (praeter stylos) connata,ovario 3- vel abortu 2- vel 1-loculato,stylis i apicalibussubulatisque, stigmate nrinuto apicali vel parum interno. Fructusinde, hiscens,drupaceus vel siccus,1-3-spermus. Typus. ByrsonimaL. C. Richardcx Humboldt,Bonpland, & Kunth, Nov, Gen. Sp (4r ed.) 1:1471821 (1822). KEY TO THE GENERA OIi BYRSONIMEAE Leaves,bracts, and bracteoleseglandular (except for gland-tipped marginal teeth or cilia on the bractsand bracteolesof some species). 2. Fruit drupaceous;hairs on the anthers,if. any,medifixed or sub-medifixed, with 2 branches;New World tropics and subtropics . BytsonimaRich. ex H. B. K 2. l"ruit dry; hairson thc anthersbasifixed Petalsall yellow; anthersbcaring 2(-4) stout, apical,ax'n-like hairs, these strongly differentiated from any other hairs on the stamen; AmazonianBrazil, southernVenezuela, and easternColombia Diacidia Grisebach Petals white an<l/or pink, or 4 white ancl the fifth pale yellow: anthersbearing many stiff but fine hairs, thoseof thc apex hardly or not at all different from other hairs on the stamen;Amazonian Brazil, southern Venezucla,and rvesternGuyana BlepharandraGrisebach Leavesand altcrnatebracteoles bearing large dorsal glands. 4. Stipulesconnate intrapetiolarly, persistent; bud spherical;connective of the anther enlarged, greatly exceeding the apically rounded locules; AmazonianBrazil, Colombia, and Venezuela,and Guyana BurdachiaAdr. Juss. 4. Stipulesconnate interpetiolarly, caducous; bud pyramidal;connective of the anther exceeded by extensions of the apically tapered locules; AmazonianBrttzil, Venezuela,and probably Colombia GlandoniaGrisebach All of thesegenera are well representedin the GuayanaHighland and I shall describeand discussthem further in my fotthcoming treatmentof the Malpighiaceae frorn that region. I am also preparing a monographof the whole genusByrsonima, ). GalphimieaeNiedenzu, Ber. Deutsch. Bot. Ges. 8:192 1890 Floriferous peduncie well developed,cven rvhen cinciuuus is reduced to 1 flower. Calyxglands 10 or fewer through fusionand loss,to none. Petalssubequal or the posteriorpetal moderatelydifferentiatcd. Stnmcns10, tlrc filamentsopposite thc sepalsusually longer than those opposite the petals, the anthers +- alike, glabrous. Carpels 2-3, connate in flower for their whole length (except styles), both or all 3 loculesuniovulatc; styles 2-3, aprcal. Fruit breaking apart into 2-3 (or fewer by abortion) 1-seededmericarps, the mericarpsoften dorsally loculicidal but not opening sufficiently to rcleasethe seed; baseof the mericarp often with an ernpty or aerenchyma-filledcavity. Type. GalphimiaCavanilles, Icon. et descr. pl . 5:61. 1799 *- 8 ,,- KEY TO THE GENERA OF GALPHIMIEAE Stylesslender and subulate,tapered distally to a minute stigma. 2. Anthers unwinged, at most apically verruculate;bracteoles egiandr,lar. 3 . Anthers without apicalornamentation; calyx glands 5 or fewer, often none; leavesusually bearingglands on baseof lamina, sometimeson petiole;from Texasto Argentina Galphimia Cav, ) . Anthers ornamented at apex rvith 2 wart-like outgrowths (<<ver- ruculate>>);calyx glands 10, with some neighboring pairs partially fused; leaveseglandular; Bahia and Amazonas,Brazil . Verucularia Adr. Juss. 2. Anthers longitudinally winged; alternatebracteoles bearing rarge glands, Amazonian Brazil, Venezuela,and probabiy Colonbia Lophanthera Adr. Juss, I . Styies stout, truncatc or subpeltatc at tl.reapex; northern Soutir America, Panam6,and the W'est Indics SpacheaAdr. Juss. As circumscribedhe rc this tribe is drastically changed from the group recognizedby Niedcnzu in 1928, Sevcralof his genera have been excludedfrom the subfamilf, and others have been assignedto other tribcs. The four genera left form two natural pairs (Gatphimia and Verrucularia vs. Lophanthera antl Spachea)that have in common similar fruits and a tendencytoward reducliol ol' tire calycineglands. tr'Iostof thesegenera were recentlystudiecl by a4acBryde(1970). I-Iegave little considerationto thc relationshipsbetween g€nera, but he did describe tltcm