Contr.Univ. Mich. Herb. l7:21-3'/.1990.

THE OF JUBELINA ()

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 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 ,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 rarely eglandular,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 betweenthe 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 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. Calyx glandsdistinct (0) to connate(1). Almost all outgroupspecies (and indeed almost all neotropical Malpighiaceae)have distinct calyx glands, so connate glandsare surelya derivedcondition. e. Latcral petals shifting from yellow (0) to pink (l). All speciesof all three outgroupshave yellow petals,so that seemslikely to be the anccstralcolor in Jubelina. f. Inner wing on lateral wing of samarawell devcloped(0) to reducedor absent (1). Intermediatewings are presentrnMezla, and were probablypresent in the common ancestorof the two gcnera; a residualcrest or winglet is oftcn formed in the two speciesof Jubelinathat have experiencedthis loss. g. Transverseoutgrowths of winglets on the lateral wings of thc samaraabsent (0) to present(1). Elaborationslike thesehave evolvedrepeatedly in mascagnioid lines, including Mezia. I consider them an adaptation for dispersalby water, becausethey increasethe surfacearea of the fruit and probably retard wetting. This adaptive significance,added to the complete absenceof even rudiments of such structures in half the speciesof Jubelina, lcads mc tcl consider them a derivedfeature. This decisionis basedprimarily on intragenericconsiderations, not on outgroupcomparison. h. Lateral wings of the samara well dcveloped (0) to partially reduced (1) to rudimentary(2). This step,too, seemslikely to be part of a shift from dispersal by wincl to dispersal by water, so polarity is assignedpartly on the basis of intragenericconsiderations. However, it is alsotrue that almostall speciesin the outgroupshave large,well-dcveloped lateral wings on their samelras.

When the six speciesof Jubelino arescored for theseeight characters.the result is the matrix shown in Thble 1. From this I derivethe diagramshown in Figure 2, which hasten steps,inclucling a parallelloss of the inner wing on the lateralwing of the samara in J. grisebachianaand -/. wilburii. One could construct a tree of the samelength by postulatingthat that inner wing was lost in a stcp betweenuleana and the rcst of the genus.then regained in the common ancestorof rosea,riparia, anclmagnifica. That would not changethe branchingpattern of the tree. In severalways I find the tree in Figure 2 intuitively satisfying.Jubelina nleana close to- and "/. wilburii have many characteristicsin common and surely belong gether in any phylogcnetic analysis.Similarly, J. roseu,J. riparia, and J. magnifica

T,qer-r.1. Species/character matrix for Jubelina. See text for description and discussi

grisebachiana t) 1 I 0 magnifica I I (, I riparia U I 0 2 rosca 0 I (, 0 uleana U 0 U (J t, 0 wilburii (J (l I I 0 (, 1990 W. R. ANDERSON:JUBELINA 25

uleana

wilburii

grisebachiana

rosea

riparia

magnifica

FIG. 2. Phylogenvof Jubelinu.Crossbars indicate shifts from ancestralto derivedcharacter-states; lctters above crossbarsdenote characters.See text for description of charactersand Table 1 for distribu- tion of character-states. form a complex of obviously related species.And J. grisebachiana,while generally more like the roseagroup than the two yellow-floweredspecies, is intermediatein its stem hairs and hooked styles, and probably deservesa somewhatintermediate placeon the tree. On the other hand, I am not so confidentabout the orientation of the tree. Parallelism and reversal have been common in the evolution of the Malpighiaceae,and just a few changesin my analysisof characterswould root the tree differently. For example, the distinct calyx glands of -/. uleana could be a reversal, and the bright yellow petal pigments of J. uleana andJ. wilburii may not be homologouswith the darker pigmentsrn Mezia, rn which casethe ancestralcondi- tion in Jubelina might actually be pink petals, which are common in Mascagnia sectionMascagnia. Therefore, I urge the reader not to regardthe tree in Figure 2 as a definitive phylogeny of Jubelina, but as a first attempt basedon the information now at hand.

GEOGRAPHICAL DISTRIB UTION

The distribution of all six speciesof Jubelina rs mapped in Figure 1. Several featuresof that distribution merit comment. Only J. grisebachianaand "/.magnifica have been found in the same vicinity, near San Carlos de Rfo Negro, and I do not know that they are actually sympatricthere. Two speciesare extra-Amazonian,but in both casesthere are strong ties between their floras and the Amazonian flora. Jubelinarosea occurs only in the Atlantic drainageof French Guiana and Suriname, where the lowland forest is continuous with that of adjacent Amazonia. Jubelina wilburii is known mostly from Central America. but it has also been collected at Anorf in Colombia. home of disjunctsfrom westernAmazonia like Dicellajulianii (Anderson 1975) and Ectopopteryssoejartoi (Anderson 1980).One related pair, "/. uleana and ,/. wilburii, share the western part of the genus'srange, and another closepair. /. roseaand,/. riparia, sharethe east. 26 CONTR.UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17

KEY TO THE SPECIESOF JUBELINA

l. Leavessparsely sericeous to nearly glabrate,the hairs sessile.+ straight.strongly appressed; lateral 4 petalsyellow. the posterior yellow or pink and white; all -5petals abaxially sericeousl hairson samara0.2-0.1 mm long. 2. Petioleof larger leaves20-40 mm longl floriferousbracts 7.5-10.-5 mm long. .5-6mm wide. with white vesture; sepalstomentose on both sides,the anterior eglandular,thc lateral 4 eachbearing 2 glandsor I or both of the glandsadjacent to the anteriorsepal much rcduced or absent; apex of styleswith an obvious dorsal hook 0.2-0.-5mm long; lateral wing of samarabcanng a well-developedinner wine parallelto centraldorsal wing. 7-1,1mm wide; Amazonian Ecuador and Peru. 7. J. uleana. 2. Petioleof largerleaves 11-20 (-25) mm long; floriferousbracts 2-1(-6) mm long. 1-2 (*3) mm widc. with yellow or brown vesture;sepals abaxially tomentellous. adaxiallv glabrous or sparselypilose near apex, the anterior eglandular,the lateral 4 all eglandularor all bearing I large central gland; apex of stylesdorsally truncate or with a rounded hook up to 0.1 mm long; lateralwing of samarabearing an inner crestor wingletparallel to dorsalwing. up to -5mm wide. this occasionallyabsent; Costa Rica. Panama.and northern Colombia. 2. J. wilburii. 1. Leaves persistently velutinous or tomentose. the hairs stalked. thcir arms varying from subcrcctto partrllelto the lamina and straightto serpentine;lateral 4 pctalspink. the posterior pink or pink and white: lateral ,1petals abaxially sericeous or the posterior-lateral2 glabrous, the posteriorglabrous; longest hairs on samara1-2.7 mm long. 3. Samarawithout wingsor wingletsbetween dorsal and lateralwings, at mostonly a crest1 mm wide parallelto dorsalwing; stemssubsericeous or appressed-tomentose.the limb of hairsat right anglesto the stalk; calyx glandsrevolute at apcx. posterior-lateral2petals eslandular- dcntate or -fimbriate: northwesternBrazil and Amazonian Colombia and Venezuela. 3. J. grisebachiana. 3. Samara with well-developedwings and winglets between central dorsal and outer lateral wings; stems velutinous. the hairs mostly erect; calyx glands attachedat apex; ptrsterior- lateral 2 petalsoften glandular-fimbriate. 4. Lamina of larger leaves 10-21 cm long. -5-1.1cm wide. rounded or abruptly short- acuminateat apex with the acumen up to -5(*10) mm long, bearingbelow on eachside 0-3 glands near base and up to 4 glandsin a singlerow distally:central dorsal wing of samara as widc as to (usually) wider than parallel inner wings borne on lateral wings. -5. Lateral wing of samarawith its outer membranouswing 22-36 mm wide; bractsand bractcolcs1.4-2.5 (-3) mm wide: lamina usuallywithout glandsnear base,occasiL)n- ally 1; Surinameand French Guiana L J. rosea. -5. Lateral wing of samara with its outer membranouswing -5*7mm widel bracts and bractcoles,l--5mm wide; lamina bearing(0-) 1-2 (-3) glandsncar baseon eachside of midrib: northeasternBrazil and French Guiana. 5. J. riparia. 4. Lamina of larger leaves 18-2U cm long, 10-18 cm wide, acuminateat apex with the arcumenoften up to 20 (-25) mm long, bearing(1-) -5-1-5scarttered glands belou'on each side; ccntral dorsal wing of samara narrower than and hidden by parallel inner wings borne on lateral wings; Amazonian Vcnezuela. 6. J. mogniltca.

The terminology used here follows that of my 1981treatment of the Mal- pighiaceaeof thc GuayanaHighland. Seepp. 24-26 of that paperfor definitionsof vesturetypes. bract, peduncle, bracteole, and pedicel.and for an explanationof the floral symmetry in the family. Here, as there. I have describedall wings of the samarasuch that width is measuredat right anglesto the ventralareole and heightis measuredparallel to it. l. Jubelinauleana (Nied.) Cuatr., Webbia 13:445. 1958. Diplopterys uleano Nied., Arbeiten Bot. Inst. Konigl. Lyceums Hosianum Braunsbcrs4: 18. 1972.

Stemssericeous to glabrate,the hairswhite or pale yellowish,sessilc. straight and mostly stronglyappressed, very short. Lamina of largcr leaves14-24 cm long, 1990 W. R. ANDERSON:JUBELINA 27

8-17 cm wide, broadly ellipticalor somewhatovate or obovate,obtuse or rounded at base,usually abruptly acuminateat apex with the acumen 5-20 mm long. bcaring 2--5 glands below on each side in a row set in from margin. sparselysericeous to nearlyglabrate on both sides,the hairssessile, straight. strongly appressed.0.2-0.4 mm long; petiole 20-40 mm long, sericeousto glabrate. Inflorcscenceproximally sericeous,distally tomentose or velutinous;bracts and bracteolesreddish, densely white-tomentellousor velutinous,7.5-10.5 mm long, 5-6 mm wide, broadlyovate, obtuse,the bracteolessomewhat falcate; peduncle2.5-4.5 mm long; pedicel6-9.5 mm long, short-velutinous.Sepals reddish,4.5-7 mm long.1.5-2 mm wide, nar- rowly oblong, acute or obtuse at apex, reflexed, apparently not inflated with aerenchyma,tomentose on both sides,the anterior eglandular,the lateral 4 each bearing2 ellipticalor ovateglands 1.6-2 mm long and 0.8-1.5 mm wide, or 1or both of the glands adjacent to anterior sepal much reduced or absent. Lateral 4 petals yellow. orange in age, abaxially sericeousexcept glabrousnear margin. the claw 2-3.5 mm long, the limb 3.5-6.3 mm long, 3.5-6 mm wide, rotund, concave, subentireor erose, the anterior-lateral2 notably larger than the posterior-lateral2 posteriorpetal white with red veins. abaxiallysericeous only in center.the claw 2.8-4.3 mm long, the limb 2.6-3.5 mm long and wide, flat, ovateor nearlysquare, bearing small sessileor short-stalkedglands all around margin. Filamcnt opposite posteriorpetal notably shorter than others,1.5-2 mm long,the rest2.3-4 mm long, longestopposite anterior sepal, connate up to 1 mm; anthers0.8-1.3 mm long, the connectivesflat or somewhat enlarged opposite sepals.Ovary velutinous; styles glabrousor piliferous,3-3.7 mm long, subequalor the anteriorshorter, the anterior nearly straightor somewhatsigmoid or arcuate,the posterior2 arcuate-or sigmoid- ascending,the apex of all 3 with a protuberant internal stigma and an obvious dorsal hook 0.2-0.5 mm long. Samaraelliptical, 80-100 x 30-40 mm, minutely velutinousor (on the wings)subsericeous, the hairs0.1-0.4 mm long, V-shapedor + straightand subappressed;fertile loculeca 5 mm in diameter;ventral areole 11- 16 mm high; central dorsal wing 9-15 mm wide. 20-28 mm high; lateral wings continuousat base,free at apex, each40-50 mm widc. 30-40 mm high, flat, entire or repand,containing an open or chamberedsterile cavity in the base2-4.5 x 6-7 mm acrossand bcaring an inner wing parallel to and almost as large as the central dorsalwing, 1-14 mm wide, 76-22 mm high, repandor coarselydcntate. TvpE. Ppnu. San Martfn: Near Tarapoto, 1855-6, fr. Spruce4950 (lectotype, here designated:C, photo at MICH! F neg. 23018). Habitat and Phenology.Tropical wet forest, at elevationsof 180-4-50m; col- lectedin flower and fruit from Septemberto January. Distribution.Amazonian Ecuador and Peru:see Fis. 1.

Aootltott,tt. SpncrlrpNsEx,rnrNEo. Ecunoon. Napo: Auca Oil Field. 60 km S of Coca, Besseet al. 055(MICH,SEL);ReservaBiokigicaJatunSacha,RioNapo,likmfromPuertoMisahualli. Cer(tn2l58 (MICH, MO)l ParqucNacional Yasuni. C'er6n & Coellct3220, Pulac'ios 2352,2113 (all MICH & MO).- Pr.ttu.Amazonas: Rfo Santiago.Caterpiza. Huushikut I199 (MICH. MO)l QuebradaHuampami. monte al laclode Huampami [ca 4'30'5.7tl'1-5'W.fidc A. Gentry. pers.comm.]. Ka-,-ap2-5S (MO). San Martin: Thrapoto,Spruce 1607 (svntypc. BM, F. GH. K. NY); Pongode Clainarachi.Ille6-147 (synr.vpc. photo at MICH of sheet formerly at B); trapoto. Ll. Williams 6603 (F). Loreto: Washintsaand vicinitv, Rio Huasaga,3'2(l'S. 76'20'W. Lew'iset ttl. llB27 (MO)l Previsto.ravine of Yurac River, Woytkowski7551 (F.MICH).

I have selectcdas lectotype the sheet of Spruce4950 at C becauseNiedenzu presumblysaw and annotatedit. 28 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17

2. Jubelinawilburii W. R. Anderson, Brittonia 28: 470.1976. Fig. 3.

Stems sericeousor eventually glabrescent,the hairs golden fading to white, sessile,straight and strongly appressed,very short. Lamina of larger leaves1I-27 cm long,6-11.5 (-14) cm wide, elliptical,ovate, or rotund, obtuseor rounded at base, acuminate or obtuse at apex, bearing (0-) 1-3 (-5) glands below on each side between midrib and margin, sparselysericeous to eventuallyglabrate on both sides,the hairs sessile,-* straight,strongly appressed,0.2-0.4 mm long; petiole 11-20 (-25) mm long, sericeous. Inflorescence proximally sericeous, distally short-tomentose;floriferous bracts 2-4 (-6) mm long, I-2 (-3) mm widc, ovate or elliptical, rounded at apex, yellow- or brown-tomentellous;peduncle 1-3.5 mm long; bracteoleslike bracts but somewhat smaller; pedicel 2-5 mm long, tomen- tellous.Sepals 2.5-3.5 mm long, 1-1.8 mm wide, narrowly ovate,acute or obtuse and revolute at apex, not inflated with aerenchyma,abaxially tomentellous, adaxiallyglabrous or sparselypilose near apex, the anterior eglandular,the lateral 4 all eglandularor all bearing 1 largegland ca 1.5 mm long and wide, ovate,acute and revolute at apex. Petals greenish yellow, abaxially sericeousexcept glabrous toward margin; lateral4 petalswith the claw 1.5-2.5 mm long, the limb 3-4.5 mm long, 2.5-4 mm wide, rotund or very broadly obovate, erose at margin or the posterior-lateral2 bearing a few glands; posterior petal with the claw 2-2.4 mm long, the limb 2.5-2.8 mm long,I.5-2.5 mm wide, flat, ellipticalor obovate, bearing glands all around margin. Filaments 1.8-3 mm long, longest opposite anterior sepal, connate up to 1 mm; anthers 0.8-1 mm long, the connectives glandular-swollen.Ovary tomentellous;styles 2-2.5 mm long, glabrousor pili- ferous, the anterior straight and inclined toward posterior petal, the postenor 2 arcuate-ascending,the apex of all 3 dorsally truncate or with a rounded hook up to 0.1 mm long. Samaraelliptical,43-80 x 20-35 mm, thinly sericeous,the hairs 0.2*0.4 mm long, straightand appressedor slightlyraised; fertile locule3-4.5 mm in diameter;ventral areole 9-14 mm high; dorsalwing 6-13 mm wide, 74-27 mm high; lateral wings continuous at base, free at apex, each22-40 mm wide, 20-35 mm high, flat, entire or repand, containinga small sterilecavity t-2 x 1-3 mm acrossin the base, this empty or partly filled with aerenchyma,and bearing an inner crest or winglet parallel to dorsal wing, up to 5 mm wide, this occasionally absent. TvpE. PnNa,ua.Veraguas: Road beyond Escuela Agrfcola Alta Piedra. above Santa F6, Pacificwatershed, 800-1000 m, 1 Jan 1975,fllfr. Luteyn & Wilbtrr 4574 (holotype:DUKE!). Habitat and Phenology.Wet forest at elevationsof 100-1000m; collected in flower from December to March, and in July and September,and with fruits from Januaryto April. Distribution. Northern Colombia to Costa Rica; to be expectedalso in the Atlantic lowlandsof Nicaragua;see Fie. 1.

AontrtoNnt- SpEcrrrlrNsExnmrr.rE'o: ColoMsrn. Antioquia: Mpio San Luis. autopista Medellfn- Sogotir. vereda La Josefina,Canon de la quebrada La Salada.Ho1'os & Herndndez757 (MO); Anori. betwecn Providenciaand Alhibe , Soejartoet al.4521 (MICH).-PaNa,nrn. Col6n: Santa Rita Ridge. Duke I5-100(MO). Dtt'y'er& Gentry 9544 (MICH. MO). Lewis et ol. 5281(MICH. MO). San Blas: El Llano-Cartf road. Km 27, llevers & Charnle.v--5089 (MICH).-Cosrn Rrcq. San Josi: vicinity of El Gcneral. Skutc'h40-lB (A. MO. NY). Heredia: Finca La Sclva, Rfo Pucrto Viejo just E of its junction with the Rio Sarapiqui.Chac1n G. I04B (MICH), D. Sntith239 (DUKE). 1990 W. R. ANDERSON:JUBELINA 29

tA T JVI I f-P' "L4 1;,c f.g {. tv "d

F-5cm cml -s- ---,

FIG.3. Jubelirutwilhurii; a. flowcring branch; b. flowcr; c. gvnoL-cium.with anterior style in centertd, ovary, crossscction; e. samura.abaxial view; f. samara.cross scction. u-d drawn from Sftutclr 1038, e-f trom Lutet,n & Wilbur 4574. 30 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17

3. Jubelinagrisebachiana W. R. Anderson, sp. nov. Jubelinabracteosa sensu auctt., non Mascagniabracteosa Grisebach in Martius, Fl. Bras. l2(1): 97. 1tl58.

Liana ramis subsericeisvel appresso-tomentosis;folia utrinque velutina vel tomentosa;4 sepalalateralia omnia glandulamagna centrali apice revoluta munital petalarosea, 4 lateraliaabaxialiter sericea, 2 postico-latcraliaeglandulosa, posticum glabrum;samarae alae laterales 25-30 mm latae,25-43 mm altae,sine ala vel alulis intermediis,vel tantum crista ca 1 mm lata ala centrali dorsaliparallela munita. Stemssubsericeous or appressed-tomentose,the hairs with a very short stalk and the limb at right anglesto it and straight to somewhatserpentine. Lamina of largerleaves 13-18 cm long,7-12 cm wide, broadly elliptical,cuneate or rounded at base.roundcd or more often abruptly short-acuminateat apexwith the acumen 2-13 (-20) mm long, bearingbelow on eachside 1-2 (-3) glandsat baseand 1-4 in a singlc row distally, persistently velutinous to tomentose on both sides, more densely so below. sometimes glabrescentabove, the hairs with arms longer than stalk and suberectto parallel to lamina and becomingserpentine; petiole 74-23 mm long, subsericeous.Inflorescence tomentose or subsericeous;bracts and bracteoles pink, 4-6.5 mm long, 1.5-2 (-2.5) mm wide, narrowlyobovate or spatulate,obtuse or rounded at apex, appressed-tomentoseor subsericeous;peduncle 0-1.5 mm long, pcdicel 5-7 mm long, subvelutinous,thickened in fruit. Sepals4.5-6.5 rnm long, 1-1.5 mm wide, narrowly obovate,acute or obtuseat apex,reflexed, distally somewhatinflated with aerenchyma,appressed-tomentose on both sides,the ante- rior eglandular,the lateral 4 all bearing l large gland 1.5-2 mm long and wide, broadly ovate, acute or emarginateand revolute at apex, often emarginateat base. Petalspink; lateral 4 petals sparselyto denselysericeous abaxially, eglandular, the claw 2.5-3.5 mm long, the limb 4.5-6.5 mm long, 5-7.5 mm wide, rotund, the antcrior-latcral 2 deeply concaveand entire to erose, the posterior-lateral2 shal- lowly concave and dentate to fimbriate; posterior petal glabrous, the claw 2.5-4 mm long, the limb 3-5 mm long, 2.5-4.5 mm wide, + flat and rectangular, glandular-fimbriateall around margin. Filaments2-3.3 mm long, longestopposite anterior sepal, up to Vt connate; anthers 0.9-1.4 mm long, those opposite sepals with enlargedglobose connectives. Ovary hispid;styles glabrous, the anterior 1.9- 2.5 mm long. straight,the postenor22-2.8 mm long, arcuate-ascending,the apex of all 3 with a rounded dorsal hook 0.7-0.2 mm long. Samaraelliptical. 60-80 x 25-43 mm, sericeousto glabrescent,the longesthairs (on the nut) 1-1.5 mm long, straight,appressed; fertile locule ca 4 mm in diameter,ventral areole 10-12 mm high; dorsal wing 70-12 mm wide, 24-30 mm high; lateral wings continuous at base. free at apex, each 25-30 mm wide,25-43 mm high, flat, entire or repand, containinga chamberedsterile cavity in the base2-3 x 2-5 mm acrossand bearing no intermediatewing or winglets,or at most only a crestca I mm wide, parallelto dorsalwing. Tvpe. VpNEzuEr-n.Terr. Fed. Amazonas: IVIC study site 4 km NE of San Carlos de Rfo Negro, I20 m, 1o56'N, 67'03'W, 4 Apr 1979, fl, Liesner 6114 (holotypc:MICH! ; isotypes:MO! VEN). Habitat and Phenology.Open areasand secondaryforest. at elevationsof 100- 200 m; collected in flower in April and May and from October to December, and with fruits in May and November. Distribution. Rio Uaup6s/Vaup6sand upper Rio Negro, Amazonian Brazil, Venezuela.and Colombia: secFig. 1. r990 W R. ANDERSON:JUBELINA 31

Aootltora,r. Spr-c'rnnNsExarrrrNeo. Bnazll. Amaz6nas: Pari Cachoeira. Rio Tiqui6 [0'15'N, 69'4-5'Wl,Coelho & F'rancisco274 (INPA): Rio Papury.Vaup6s. Rio Negro, Froh 21173(lAN. NY); Panurd, Rio Uaup6s. Spruc'e2853, syntype of Masc'agniabracteosa Griseb. (BM. G. K, NY).- Vr-Nr-.zur.r.e.Terr. Fed. Amazonas: Vicinity of San Carlos de Rfo Negro. Altmartl et al. 3538(MICH). Croar59637, Liesner -1696,696-1,7523 (all MICH & MO). 7555(MO); CarreteraSan Carlos-Solano.2- 6 km SW of Solano, Morillo et al. 3980(MfCH)l San Sim6n de Cocuy, l-2 km NW of the Piedra de Cocuy. Morillo et al. 1l4l (VEN).-Cor.oMBtA. Vaup6s: Mit:'r, Cuatrecasas7255 (US); Piracuara. Romero Castanedu-175-l (COL).

When Grisebachpublished Mascagniabracteosd he cited two syntypes,Spruce U0931from near Manaus and Spruce l2\53l from near Panurd [:Ipanor6]. These represent different species.tn 1912 Niedenzu selectedSpruce 2853 as lectotype when he transferredthe epithetbracteo,sato Diplopterys;that choicewas followed by Cuatrecasasin 1958,when he transferredthe speciestoJubelina,and by me in 1981.Unfortunately I now find, after careful study of the original description,that Niedenzu'schoice of a lectotype was inappropriate and cannot stand. A point-by- point comparison of Grisebach's description with Spruce 1093 and 2853 shows almost perfect agreement with 1093 and very extensivedisagreement with 2853. This pattern holds true for size and vesture of the lamina, length of the petiole, length of the peduncle and pedicel, vestureof the bracteoles,shape and vestureof the sepals,number of calyx glands,and shapeof the style apex. The only evidence that Grisebachconsulted 2853at all in drawing up the descriptictncomes from the bracteoles,for which he gave a length closer to that of 2853 than 1093. I cannot avoid the conclusionthat Grisebachbased his descriptionalmost entirely on 1093, which shouldbe the lectotypeaccording to Article 8.1 and Recommendation7tl-.3 of the International Code of Botanical Nomenclature.Therefore, I here designate the sheet of Spruce 1093 at M the lectotype of Mascagniabracteosa Grisebach, which becomesthe correct name for the speciesI named Mascagniaheterocarpa in 1981.That leaves"Jubelina bracteosa" without a name. so I am namine it here in honorof A. H. R. Grisebach.

4. Jubelinarosea (Miq.) Nied. in Pulle, Enum. Pl. Surinam472. 1906. Hiroearoseo Miq., Stirp.Surin. Sel. in Natuurk.Verh. Holl. Maatsch.Wetensch. Haarlemser. 2. l:84: 1tt5l. Jubelina riparia var. hiraeoidesSagot, Ann. Sci. Nat. Bclt. 6' 56r. 12: 187. 1881.-TypE. FneNcHGutnNn. Mdlinon [9a] in 1862(lectotype, here desig- nated:P!; isolectotype:P!). Diplopterysrose a (Miq.) Nied., Arbeiten Bot. Inst. Konigl. LyceumsHosianum Braunsbers1: 20. 1912.

Stemsvelutinous to eventuallyglabratc. the hairs up to 0.6 mm long, mostly crect and -t Y-shaped. Lamina of larger leaves 10-20 cm long. 5-13 cm wide, obovateor broadlyelliptical to almostrotund. cuneateor truncateat base.rounded or abruptly short-acuminatcat apex with the acumen up to 5 (-10) mm long, bcaringbelow on each side 0 (-1) gland near baseand 2-4 glandsin a singlerow distally.persistently vclutinous on both sides,more denselyso below, the hairs aboveY-shaped or simpledue to suppressionof 1 branch.Y-shaped below; petiole 7-21 mm long, vclutinous.Inflorescence velutinous; bracts and bracteolespink. 6- 9 mm long, I.1-2.5 (-3) mm wide. oblanceolate,acutc. appressed-tomentose; peduncle1-3 mm lclng;pedicel 6-12 mm long. tomentose.somewhat thickened in fruit. Sepalspink, -5-8.5mm long. 1.2-1.7mm widc, narrowlyoblong or obovate, -)L CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 obtuse or acute at apex, reflexed, distally inflated with aerenchyma, loosely subsericeouson both sides,the anterior eglandular,the lateral 4 all bearing 1 large pink gland 2-2.5 mm long, 7.5-Z mm wide. rounded or emarginateand non- revolute at its apex. Petals pink; anterior-lateral2 petals with the claw 2-3 mm long, thc limb 4-6 mm long, 4.5-6 mm wide, deeplyconcave, erose and eglandular at the margin, abaxiallysericeous; posterior-lateral 2 petalswith the claw2-2.5 mm long. the limb 3.5-5 mm long and wide. moderatelyconcave, short-fimbriate all around margin with the fimbriae mostly glandular to mostly eglandular,abaxially glabrousto sericeousin center; posterior petal glabrous,the claw 3-3.5 mm long, the limb 3-4 mm long, 2.5-4 mm wide, rotund or quadrate,nearly flat, glandular- fimbriate all around margin. Filaments1.5-2.5 mm long. longer oppositesepals than petals,up to % connate;anthers 0.7-l .2mm long, largestopposite posterior- lateral petals, the connectivesmoderately glandular-swollen.Ovary hispid; styles glabrousor hispid at base, 1.5-2 mm long, equal or the anterior slightlyshorter than posterior 2, the anterior straight, the posterror 2 arcuate at base and then straight,the apex of all 3 truncate dorsally or with an apiculum up to 0.1 mm long. Samararoughly ellipticalin outline, 55-80 x 35-60 mm, sericeousto glabrescent, thc longesthairs (on the nut) 2-2.7 mm long, straight.appressed; fertile locule 2.5- 4 mm in diameter,ventral areole5.5-8.5 mm high; centraldorsal wing 10-18 mm wide, 20-30 mm high; lateral wings confluentbut often deeply emarginateat base, free at apex. each consistingof: 1) an outer membranoussemicircular wing 22-36 mm wide, 35-60 mm high. flat or slightlycorrugated, entire or repand 2) an inner corrugatedand deeplylobed wing 5-11mm wide. parallelto the centralwing and narrowerthan it; and 3) a sterilecavity betweenthe outer and inner wings.2-3 x 3-5 mm across,chambered, covered externally by 5-8 irregular paralleltransverse wingletsup to 2-7 mm wide, theseconfluent with the inner wing. Typr. SunrNnup.Upper Marowijne River, Kappler I 807(holotype : U ! ; isotypes: LE,! P!. Field Museum negative12717 (MICH!) showstwo specimensformerly at B. The one with leaves and flowers was an isotype of Hiraea rosea, but the other, comprisingonly fruits from Mtlinon in 1864,from French Guiana, was not a type.) Habitat and Phenolclgy.Primary and secondaryforests on terrafirma (always?) from near sea level to 700 m; collectedin flower from August to October and once in March, in fruit from August to November. Distribution.Suriname and northwesternFrench Guiana: see Fie. 1.

AnorrrclNnl SpEcrrutNsE,x,rmtNto. SuRrNnmr.. Brownsberg Naturc Park.90 km S of Paramaribo. Mazaroni Plareau.Mori & Bolten B-199(MICH, NY)l Leli' Mts. 17-5km SSE of Paramaribo.Mori & Bolren,g-5.51(MlCH, NY), Brownsberg.distr. Brokopondo, Sung LBB-11811(U): Tirpan;rhoniRiver. Versteeg720 (U).-FnrNcs CiurnN,r.Itany and Marouini. BAFOC Servic'eF'orestier 2965 (U): St. Laurent toward Paul Isnard, Km,l0. Billiet & Jartlin 1704 (MICH); region of Paul Isnard, between Citron and Mont D6cou D6cou, Cr(mersS2l1 (MICH); edgeof the Mana IRiverl. Mdlinon [97] in 185a (P)l sinc lctc..Milinon in 1tt64(F. GH. NY, P); sine krc., Mdinon in 186-5(P); Maroni. St. Laurent. "parcclle Mdlinon [112] in 1t376(P)l Sinnamary,road from St. Elie. ARBOCEL." Prtvctst720 & 819 (both MICH), Riera.t62 (CAY). Sustre6057 (CAY. MICH). Sttstre6154 (CAY. P): St. Laurent. .Sttuhirouin ltJ96(P).

5. JubelinaripariaAdr.Juss. in Delessert,Icon. Sel.Pl.3: 19,pl. 32. 1837[lt]38]. Diplopterys riparia (Adr. Juss.) Nied., Arbeiten Bot. Inst. Konigl. Lyceums Hosianum Braunsbers.4: 20. 1912.

Stemsvelutinous to eventuallyglabrate, the hairs up to 0.8 mm long, mostly erect and -r Y-shaped. Lamina of larger leaves 1l-21 cm long,8-14 cm wide, aa 1990 W. R. ANDERSON:JUBELINA J-) broadly elliptical or obovate to rotund, cuneate, truncate, or rounded at base, rounded or abruptly short-acuminateat apex with the acumen up to 5 (-10) mm long, bearing below on each side (0-) 1-2 (-3) glandsnear baseand (0-) l-2 (-3) in a single row distally, persistently velutinous on both sides, lnore densely so below, the hairs Y-shaped; petiole B-I7 mm long, velutinous. Inflorescence velutinous;bracts and bracteolespink, 6-8.5 mm long, 4-5 mm wide, obovate, rounded at apex, appressed-tomentose;peduncle 2 mm long; pedicel 8-12 mm long, velutinous,thickened in fruit. Sepalspink, 5-7 mm long, I.5-2 mm wide, narrowly obovate, obtuse at apex, reflexed, distally inflated with aerenchyma, appressed-tomentoseon both sides,the anterior eglandular.the lateral4 all bearing 1 largepink gland 2-2.5 mm long, 2-2.2 mm wide, roundedand non-revoluteat its apex.Petals pink; anterior-lateral2 petalswith the claw 3-4 mm long, the limb 6-8 mm long, 5-6 mm wide, deeplyconcave, erose and eglandularat margin,abaxially denselysericeousi posterior-lateral 2 petalswith the claw 2.5-3 mm long, the limb 4.5-5.5 mm long, 4-4.5 mm wide, flat or somewhatconcave, glandular-fimbriate all around margin or only on distal /2, abaxrallyglabrous or sparselyto moderately sericeousin center;posterior petal glabrous,the claw 3-4 mm long, the limb 4.2-5 mm long, 4-4.5 mm wide, ovate or quadrate,flat, glandular-fimbriateall around margin. Filaments2.5-3.3 mm long, mostly longeroppositc sepals than petals,up to rk connate;anthers 0.8-1.3 mm long, largestopposite posterior-lateral petals, thc connectivesmoderately glandular-swollen.Ovary hispid; stylesglabrous or hispidat base,2-2.5 mm long, straightor the posterior2 slightlyarcuate, subequal with the anteriorslightly shorter than the posterior2,the apextruncate dorsally or with an apiculumless than 0.1 mm long. Szrmararoughly circular in outline.2I-29 mm in diamctcr,sericeous to glabrescent,the longesthairs (on the nut) 1.5-2 mm long, straight,appressed; fertile locule 3-5 x 5-8 mm across;ventral areole8-10 mm high; centraldorsal wing 8-10 mm wide,23-26 mm high; lateralwings conflu- ent at base,free at apcx, eachconsisting of: 1) an outer membranoussemicircular wing 5-7 mm wide. 20-25 mm high, flat or corrugated and rcpand or coarsely dentate at margin, 2) an inner repand or coarselydentate wing 5-9 mm wide, parallelto thc central dorsalwing and narrower than it to just as wide; and 3) a greatlyinflated sterile cavity between outer and inner wings,7-10 mm across,open or chambered,covered externally by many irregularoutgrowths up to 6 mm long, thesescattered or conflucntinto lobed, gencrallytransverse winglets. TypE.FRENcs GureNn. Oiapock and Oyes Rivers,Leprieur in 1832(holotype: P-JUI:isotypes: F! G. P!). Habitat and Phenology.Y'6rzea and riverine forcstsat low elevations;collected in flower in September,November, February,and March, in fruit in December and March. Distribution.AmapA, Brazrl.and adjacentPard and FrenchGuiana; see Fig. 1.

Auor rr

As noted above in the discussionof genericcharacters and trends,the fruit of Jubelino riparia seemsto representthe culmination of an evolutionary shift from wind dispersaltcl water dispersal. Its membranous wings are much reduced, the 34 CONTR.UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 sterile cavities are enlarged, and there are many irregular outgrowths on the sa- mara. The samara seems most unlikely to fly far, but it should float quite well, buoyed by the large air-filled cavities,and the extra outgrowthsof the lateral wings should retard wetting. The embryo is significantlylarger in this speciesthan in "I. rosea;note that the fertile locule is 3-5 x 5-8 mm acrosshere, 2.5-4 mm across there. Christiane Anderson (pers. comm.) has found that the embryo is always much larger in speciesof Srlgmaphyllon that have shifted from wind to water dispersal,and I have seen the same tendency in other groups of Malpighiaceae.I interpret this to be an evolutionary responseto releasefrom the constrainton the weight of the embryo imposed by wind dispersal.Natural selectioncould be ex- pected to favor a larger, heavier embryo with more stored reservesif that change did not decreasethe efficacyof dispersal.

6. Jubelinamagnifica W. R. Anderson,Mem. New York Bot. Gard.32:228. 1981. Fig. 4.

Stems velutinous to eventually glabrate, the hairs up to 1 mm long, erect, fusiform, distally bifurcate with the short branches + erect and often unequal. Lamina of larger leaves18-28 cm long, 10-18 cm wide, broadly ovate or elliptical, rounded at base, abrupty acuminateat apex with the acumenoften up to 20 (-25) mm long, bearing (4-) 5-15 scatteredglands below on each side, persistently velutinouson both sides,more denselyso below, the hairs Y-shaped;petiole 9-20 (-25) mm long, velutinous.Inflorescence velutinous; bracts and bracteolespink, 6- 8 mm long, 3-4 mm wide, obovate or elliptical, broadly obtuseor rounded at apex, appressed-tomentoseor subsericeous;peduncle 0-2.5 mm long; pedicel 8-15 mm long, velutinousor tomentose,much thickenedin fruit. Sepalspink, 6.5-7 mm long, 1.8-2.5 mm wide, narrowly obovate or oblong, rounded at apex, reflexed, distally much inflated with aerenchyma, tomentose on both sides. the anterior eglandular,the lateral4 all bearing1 largewhite gland 2-3.5 mm long, 1.5-3 mm wide, rounded and non-revolute at its apex. Lateral 4 petals pink, abaxially sericeous,the anterior-lateral2 with the claw 2.5-3.5 mm long, the limb 6-7 mm long, 7-9 mm wide, deeply concave,erose at margin and eglandularor bearinga few glandsat baseiposterior-lateral 2 petalswith the claw 1.7-3 mm long, the limb 6-7 mm long, 5-6.5 mm wide, flat or somewhatconcave, fimbriate or glandular- fimbriate all around margin; posterior petal white with pink tints, glabrous, the claw2.5-3.5 mm long, the limb 5-7.5 mm long, 3-4.5 mm wide, ovate,elliptical, or obovate, flat, glandular-fimbriateall around margin. Filaments 1.8-3 mm long, longer opposite sepalsthan petals, up to lz connate; anthers0.9-1.8 mm long, longest opposite posterior-lateral petals, the connectivesmoderately glandular- swollen,especially opposite sepals.Ovary hispid; stylesglabrous, 2-3 mm long, straight or the posterior 2 arcuate, subequal,the apex truncate dorsally or with a hook up to 0.1 (-0.3) -- long. Samarasubcircular,40-50 (-75) mm in diameter, sericeousto glabrescent,the longest hairs (on the nut) 1-2 mm long, straight, appressed;fcrtile locule 7-8 mm in diameter, ventral areole lI-17 mm high; cen- tral dorsalwing 5-10 (-15) mm wide,25-30 (-35) mm high; lateralwings free at base or confluent but deeply emarginate, free at apex, each consistingof: 1) an outer membranoussemicircular wing 10-15 (-30) mm wide,40-45 (-60) mm high, corrugated and repand at margin; 2) an inner, strongly plicate or corrugated, coarselydentate wing ca B-72 (-17) mm widc, parallelto and hiding the central dorsal wing; and 3) an inflated sterile cavity between outer and inner wings, 4-7I 1990 W. R. ANDERSON:JUBELINA 35

'a t.r,k ,;*. .r-l 1a \.\l \1

ii

1r- 1"1

ic b

FIG. 4. Jttbelinamagnit'ica: a, flowering branch. x0.5; b, detail. abaxialsurface of Ieaf. xl0; c, hair frcrm abaxial surface of leaf. x25: d, stem hairs. x25; e, flower, x2.5; f, gynoecium, x10; g. samara,abaxial view, x0.5;h. samara,cross section, x0.5. Drawn by Karin Douthit, a-f from Maguire er al. -16748,g and h from Wrdack & Adderley 43579. 36 CONTR.UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 mm across,chambered, covered externally by 7-10 irregular,often lobed,generally transversewinglets 3-7 (-ll) mm wide. Chromosomenumber: n : 10 (countedin Anderson 13361). TVps. VENEzupln. Terr. Fed. Amazonas: Rfo Yaciba, Rfos Pacimoni-Yatua. Casiquiare,Maguire et a\.36748 (holotype: NY!;isotypes: MICH!NY! US!VEN!). Habitat and Phenology.Climbing in treeson terrafirmabut near lowland rivers, at elevationsof 100-150 m; collected in flower and fruit from November to June. Distribution.Known only from Amazonas,Venezuela; see Fig. l.

AoolrtoNnt- Spl,ctlr'Ns ExevrNr.o. VENEz.unln.Terr. Fcd. Amazonas: Depto. Rfo Negro, Rfo Mawarinuma. Neblina Basc Camp. (l'49'-50"N.66'9',10"W, Anderson 1-l-161(MICH. VEN). Liesner 1-58179(MICH). 17159(MO), between Rfo Mawarinuma and headwatcrsof Rio Baria. 0'-52'N.66"15'W. ()entrv <9Srein 47223 (MICH): 8 km NE of San Carlos de Rfo Negro, Lie.sner,9B12(MICH. MO); from Los Tamboresof Comisi6n de Limite to Cano Erubichi on Rio Baria. 1T2'N.66'20'W. Liesner17117 (MICH): t1,pelocalitv. Muguire et al. 36694(MICH, NY. US, VEN). 36694-A(NY), 3674-1(MICH.NY. US, VEN): uppermost Rfo Yatua, Maguire et al. 12601(NY); Rio Siapabetween Raudal Callincta ancl SaftoGallinctit.Wuduck & Adderlev 4-1579(MICH. NY. US. VEN).

The sole collcction from near San Carlos de Rfo Negro, Liesner 8842, differs qualitatively and quantitativelyfrom all other known collectionsof this species.The generallyvelutinous vesture of its stemshas an understoryof apprcssedhairs. The leavesbear rather few glandsbelow, perhaps only thrcc on eachside. Thc posterior stylesare arcuate,and all three stylesbear well-devcloped dorsal hooks at the apex. The samarais larger in almostall dimensionsthan is typical,these extremes being given abovc in parentheses.Liesner 8842 docs not seemto be sufficiently distinct to merit taxonomic recognition,but it does suggestthat the descriptiongiven above may have to be expandedsignificantly as this speciesbecomes better known. An- other possibility is that Liesner 8842 resultedfrom hybridization bctween Jubelina magnifca and "/.grisebachiana, which grows near where 8842 was found. Genetic influencc from J. grisebachianacould account for all the unusual features noted above, except perhaps the size of the apical-dorsalhooks on the styles. Further fieldwork in thc vicinity of San Carlos de Rfo Negro may clarify the statusof this anomalousplant. Jubelinamagnifica seems to havc begun the evolutionaryshift toward dispersal by water.but not to havegone nearlyas far in reductionof the samarawings as has J. riparia. Like the latter species,this one also has an enlargedembryo. See the discussionunder J. riparia.

EXCLUDED SPECIE,S

Jubelina nicaraguensrsGrisebach, Vidensk. Mcddel. Dansk Naturhist. Foren. Kj6benhavn 1t153:48-49. 1854.-The correct name for this is Callaeum nicaroguense(Grisebach) Small, N. Amer. Flora 25 128. 1910.See Johnson's re- ccnt revisionof Callaeum(1986).

ACKNOWLE,DGME,NTS

This rcsearchand its publication were supportedin part by Grants No. DEB-S103522 and BSR- t37(X)3-10from the National ScienceFoundation to The University of Michigan. The Expedition ro the ParqueNacional Scrranfa La Neblina was organizcdand directed by the Fundacicinpara cl Desarrrollodc las ClicnciasFfsicas. Matemdticas y Naturalcs of Venezuela.with the support of the Ministerio de 1990 W. R. ANDERSON:JUBELINA 37

Educaci6n. Consejo Nacional de InvestigacionesCientfficas y Tecnol6gicas,and the Instituto Nacional de Parques,under the coordination of Dr. Charles Brewer-Cariasand with the collaborationof the National ScienceFoundation and the National GeographicSociety of the United States.Figure 3 was originally published in Brittonia 28: 411. 1976.and Figure 4 in Memoirs of The New York Botanical Garden 32:.229. 1981. Specimcnsbelonging to the following herbariawere studied.and I sendmv thanks to the curatorsof thosecollcctions: A. BM. CAY. COL. DUKE. F. G. GH. IAN. INPA. K. LE. MICH. . MO. NY. P. SEL. US. VEN.

LITERATURE CITED

Andcrson. W. R. 1975.Dicella, a genus of Malpighiaceaenew to Colombia. Acta Amaz. 5:279-283. 1978.Byrsonimoideae, a new subfamilyof the Malpighiaceae.Leandra 7:5-18. 1977. 1980.Ectopoptervs, a new genus of Malpighiaceaefrom Colombia and Peru. Contr. Univ. Mich. Herb.l4: ll-t-5. 1981.Malpighiaceae. ln The Botany of the Guayana Highland, Xl. Mem. New York Bot. Gard. 32: 2l-30-5. Cuatrecasas.J. 19-58.Prima Flora Colornbiana.2. Malpighiaceac.Webbia 13:343-66,1. Gates, B. 1982.A monographof Banisteriopsisand Diplopterys, Malpighiaceae.Flora Neotropica 30: l-

l,,trnr.L,"n,O. M. 1986.Revision of the neotropicalgenus Callaeum(Malpighiaceae). Syst. Bot. 11:335- 3,s3. Lowrie, S. R. 1982.The palynology of the Malpighiaceaeand its contribution to family systematics. Ph.D. dissertation,pp. i-xii. 1-35,1,The Universityof Michigan. Niedcnzu, F. 1912. Malpighiaceae americanae I. Arbeiten Bot. Inst. Konigl. Lvceums Hosianum Braunsberg4: 3-34. 192tt.Malpighiaceae. ln Das Pflanzenreiclr,cd. A. Engler. IV. l4l:1-870. Leipzig: Wilhelm Engelmann.