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The Taxonomy of Jubelina (Malpighiaceae)

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The Taxonomy of Jubelina (Malpighiaceae) Contr.Univ. Mich. Herb. l7:21-3'/.1990. THE TAXONOMY OF JUBELINA (MALPIGHIACEAE) William R. Anderson Universityof MichiganHerbarium North University Building Ann Arbor. Michisan48109-1057 The Malpighiaccaeare not as numerousor diversein the Amazonianlowlands as in drier, more open habitatslike the Planalto of central Brazil, but some genera are mostly or entirely Amazonian. One such is Jubelina,four of whose six species occurin the Amazoniandrainage system, the other two beingfound in nearbyareas whosefloras have a strongAmazonian component (Fig. 1).My purposein this little monographis to bring togetherwhat is known about the genus,and to offer some suggestionsabout its evolutionaryhistory. JubelinaAdr. Juss.in Delcssert,Icon. Sel.3: 19,pl.32. 1837[1838]. SprucinaNied., Arbeiten Bot. Inst. Konigl. LyceumsHosianum Braunsberg 3: 18.1908. Diplopteryssubgenus Jubelina (Adr. Juss.)Nied., Arbeiten Bot. Inst. Konigl. LyceumsHosianum Braunsberg 4: 16. 1912. Woody vines. Leavesopposite, the petiole cglandular,the lamina flat or very slightlyrevolute at margin, bearingimpressed glands or rarelyeglandular, the lat- eral veinsprominent below and interconnectedby t parallel"scalariform" tertiary veins;stipulcs small or minute, triangular,borne on baseof pctiole. Inflorescences axillaryand terminal, decompound,thyrsiform. containing much-reduced bractlike leavcsbelow the floriferous bracts. the flowers ultimately borne in umbels of 4 or corymbs of 6; bracts and bracteoleslarge, pubescenton both sides,persistent; peduncleshorter than pedicel.Sepals 5, ncarly distinctnarrowly ovate, obovate. or oblong. spreadingto cxposeoutermost petal in enlargingbud, the anterior sepal cglandular,the lateral4 usuallybearing 1 largegland each.formed by t complete fusion of 2 (except in J. uleana,with 6-8 distinctglands), occasionally all sepals eglandular.Petals pink or yellow,5, at lcast thc antcrior-lateral2 abaxiallyscri- ceous,the latcral4 spreading,the posteriorerect. Receptacle glabrous. Stamens 10, glabrous;filarncnts very stoutopposite posterior-lateral petals. Ovary of 3 carpels,1 antcrior and 2 posterior,all fcrtile, adaxiallyadnate to a common axis; styles3, subterminal.the apex with a large internal stigmaand dorsallytruncate or short- hookcd. Fruit brcaking apart into 3 1-seededsamaras on a high pyramidaltorus; samarawith a narrowly ellipticalor linearventral areole 1-3 mm wide. a semicircu- lar. cntire or rcpand. centraldorsal wing often extendedforward at apex between lateralwings. and 2large lateralwings usually confluent at base,each lateralwing with a complexstructure comprising at leastan outer membranous wing and a stcrile cavitydeveloped in its base,parallel to the fertile locule,during maturationof the fruit, and frequcntly bearing additional wings, winglets.crests, or irregular out- growthsbetwcen outer wing and centraldorsal wing. Embryo with thick flat sub- equalcotyledons. I slightlylonger than the other and bent back over it at thc apex. TypE.Jubelina riparia Adr. Juss. 21 22 CONTR.UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 Jubelina o grisebachiana x magnifica I riparia O rosea Y uleana :.;/-\\'..' '\'r.../. A wilburii rl ) ', \ ,, 1a\ ,1., i ,ll\ e-, I FIG. l. Distribution of all speciesctf Jubelina. Flora Neotropica base map no. 1. copyright the Universitvof Utrecht. NOTEWORTHY GENERIC CHARACTERS Jubelinais one of the vining generathat have the principal wings of the samara lateral (seefurther discussionbelow under Extra- and IntragenericRelationships). It is set off as a genusby the following combination of character-states:scalariform tertiary veins, large decompound terminal inflorescenceswith the flowers in um- bels, large bracts and bracteoles,long narrow sepals.at least the two anterior- lateralpetals abaxially sericeous, and the samaraswith a high narrow ventral areole and complex lateral wings, each of which contains a sterile cavity parallel to the fertile locule and usually bears at least one crest or wing between the outer lateral wing and the central dorsal wing. In addition, five of the six speciesare notable for having the paired calyx glands (if present) connate to form one large gland in the centerof the sepal. The sterile cavities in the fruit are especiallyinteresting, becausethey are unique in this genus.They seem likely to be an adaptationfor dispersalbv water, and suggestthat even in species like Jubelina uleana and J. grisebachiana,which have among the largestsamaras in the family, water may be an important factor in their dispersal. In "/. magnifica there seems to have been a shift toward more dependcnceon water, reflected in reduced fruit wings, and in J. riparia this trend has culminated in samaraswith minimal wings but very large inflated bladders. Another significanttrend in the genusis for the reducedleaves. bracts, bracteoles, and sepalsin the large inflorescenceto becomecolored. Thesecharacteristics com- bine to produce a picture of plants well adaptedto life in wet lowland forests.They are strong vines that can reachthe tops of largc trees.Their large inflorescencesfull of colored parts presumably attract pollinators from some distance.an important adaptation since individual plants are often far apart in such forests. And their fruits are well adapted for primary dispersalby wind followed by secondarydis- persalby water. ^a 1990 W. R. ANDERSON:JUBELINA L-) EXTRA- AND INTRAGENERIC RELATIONSHIPS Most neotropicalgenera of Malpighiaceaecan be placedin one of two subfami- lies. The one that has most of the lessspecialized character-states is the Byrsonim- oideae,in which the pollen is tricolporateand the plantsare treesor shrubsand bear unwinged fruits (Anderson 1978). The other large subfamily must be called the Malpighioideae,because it includesthe genusMalpighia, which is atypicalin bearing fleshyfruits but is descendedfrom the wing-fruitedgenus Mascagnia. Most speciesin the Malpighioideae are vines and bear winged fruits, and the pollen is of several derivedtypes. Within the MalpighioideaeMascagnia is centralto a group of generain which the principal wings of the samara are lateral, not dorsal. When one looks beyond the samaras,one finds the mascagnioidgenera to be rather diverse.Hiraea andMascagnia were separatedlong ago, and only recentlyJohnson (1986) has segre- gated Callaeum from Mascagnia, which still remains a probably unnatural assem- blage.Jubelina is mascagnioid,but it is not very similar in most charactersto other mascagnioidgenera. Niedenzu (1928)included Jubelinawith Mezia asa subgenusof his genus Diplopterys. Gates (1982) has shown that Diplopterys sensu stricto is a segregatefrom Banisteriopsls,not at all closely related to Jubelina and Mezia. The latter two certainly deserveto be recognizedas separategenera, as was done by Cuatrecasasin 1958, but I agree with Niedenzu that Mezia is closely related to Jubelina. It has a similar inflorescence,long narrow sepals,and samaraswhose lateral wings are basallyconfluent and often bear intermediateelaborations. How- ever, Mezia is advanced in several characters(Anderson 1981) and can only be regarded as a sister genus to Jubelina,'neither could reasonablybe considereddi- rectly ancestralto the other. Moving further afield,one naturallylooks to Mascagnia and its segregatesfor other outgroupswith which to compareJubelina. The best of theseis probably Callaeurn(Johnson 1986), which often hasumbellate inflorescences and sericeouspetals, and one speciesof which (C. nicaraguense)bears a complex structure between the lateral and dorsal wings of the samara. Within Mascagnia itseff, the best group to which I can compare Jubelina is Niedenzu's section Pleuropterys(1928), which hashairy petalsand sometimesbears intermediate elabo- rationson the lateralwings of the samara.However, the lateralwingsare usually free at the baseand the inflorescenceis racemose,so the similarityis lessthan compelling. No help is to be obtained in this case from consideration of chromosome numbers or pollen. The only chromosomenumber known for Jubelinais my count of n : l0 in J. magnifica.Most wing-fruited MalpighiaceaeI have counted have ten pairs of chromosomesor a multiple of that number, so that count simply confirms that Jubelinais correctly placed in the Malpighioideae.The pollen, as describedby Lowrie (1982), is of a moderatelyspecialized rugose type common among mas- cagnioidMalpighiaceae. A believablephylogenetic analysis of Jubelinais made difficult by the lack of an obvictusoutgroup to guide the assessmentof the polarity of characters.Neverthe- less,I am includinga simple analysis,based on rather few characters,in order to summarizesome clear relationshipsbetween the speciesand presentmy best esti- mate of phyletic trends. The principal outgroup employed was Mezia, with secon- dary consideration given to Callaeum and Mascagnia section Pleuropterys. The charactersand character-statesuscd in the analysisare given below, with my rea- sonsfor assigningpolarity as I did: a. Leaf hairsshifting from sessile(0) to stalked(1). Sessilehairs are commonin all three outgroupsand stalkedhairs are rare, so it seemsbest to considerstalked hairs derived within Jubelina. 24 CONTR.UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 b. Leaf glandsshifting from uniseriate(0) to scattered(1). Having the leaf glands scatteredis an unusualcondition, apparentlynot found in any of the outgroups, so it is surely derived within Jubelina. c. Sepalsuninflated (0) to distallyinflated with aerenchyma(1). This conditionis unknown in any of the outgroups, so it is probably derived within .lubelina. d.
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